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Notes and Comments • Braz. J. Biol. 86 • 2026 • https://doi.org/10.1590/1519-6984.305836 linkcopy

Open-access Alpha and beta diversity reveal the conservation value of a protected riparian forest in the Northern Pantanal

A diversidade alfa e beta revelam a importância da conservação de uma floresta ripária protegida no Pantanal Norte

SCIMAGO INSTITUTIONS RANKINGS

Abstract

Riparian vegetation plays a key role in maintaining biodiversity and regulating ecological processes in the Pantanal, one of the most dynamic ecosystems and one of the most sensitive to seasonal hydrological fluctuations. This study assessed the alpha and beta diversity of a riparian area located in the Sesc Pantanal Private Natural Heritage Reserve (RPPN Sesc Pantanal) through phytosociological parameters, diversity indices, spatial dispersion patterns, and floristic similarity among six sampling plots. A total of 170 individuals were recorded, distributed across 19 species and 12 families. Alpha diversity showed values consistent with those reported for riparian formations in the Pantanal (H’ = 2.48 and J’ = 0.84), with Fabaceae standing out as the most representative family. The rarefaction curve and the Chao1 estimator (22 species) indicated sampling sufficiency and a low number of undetected species. Beta diversity was high, with low overall floristic similarity among plots, as evidenced by clusters formed only at lower similarity levels—approximately 20% in the Bray–Curtis index and 44% in the Sørensen index. Only plots P1 and P2 showed higher similarity, while P5 stood out due to its greater dissimilarity. The results indicate a structurally stable and functionally preserved vegetation, primarily influenced by local hydrological variations, reinforcing the relevance of the RPPN Sesc Pantanal for flora conservation and for monitoring environmental changes in the Northern Pantanal.

Keywords:
wetlands; phytosociology; conservation; floristic similarity; spatial dispersion

Resumo

A vegetação ripária exerce papel fundamental na manutenção da biodiversidade e na regulação dos processos ecológicos no Pantanal, um dos ecossistemas mais dinâmicos e sensíveis às variações hidrológicas sazonais. Este estudo avaliou a diversidade alfa e beta de uma área ripária localizada na RPPN Sesc Pantanal, por meio de parâmetros fitossociológicos, índices de diversidade, padrões de dispersão espacial e similaridade florística entre seis parcelas amostrais. Ao todo, foram registrados 170 indivíduos, distribuídos em 19 espécies e 12 famílias. A diversidade alfa mostrou valores compatíveis com formações ripárias do Pantanal (H’ = 2,48 e J’ = 0,84), com destaque para Fabaceae como família mais representativa. A curva de rarefação e o estimador Chao1 (22 espécies) indicaram suficiência amostral e presença reduzida de espécies não detectadas. A diversidade beta foi elevada, com baixa similaridade florística geral entre as parcelas, evidenciada pelos agrupamentos formados apenas em níveis menores de similaridade, aproximadamente 20% no índice de Bray–Curtis e 44% no índice de Sørensen. Apenas P1 e P2 apresentaram maior similaridade, enquanto P5 destacou-se pela maior dissimilaridade. Os resultados indicam uma vegetação estruturalmente estável, funcionalmente preservada e influenciada principalmente pelas variações hidrológicas locais, reforçando a relevância da RPPN Sesc Pantanal para a conservação da flora e o monitoramento de mudanças ambientais no Pantanal Norte.

Palavras-chave:
áreas úmidas; fitossociologia; conservação; similaridade florística; dispersão espacial

1. Introduction

The Pantanal, the largest continuous floodplain in the world, is recognized as a World Natural Heritage Site and Biosphere Reserve by UNESCO due to its high biodiversity and ecological importance (UNESCO, 2010). It is located within the Upper Paraguay River Basin and extends across the Brazilian states of Mato Grosso and Mato Grosso do Sul, as well as portions of Bolivia and Paraguay (Tomas et al., 2019, 2025).

The floristic composition of the Pantanal is characterized by a complex mosaic of phytophysiognomies that includes grasslands, savannas, and forest formations (Cunha et al., 2023; Soriano et al., 2020). This diversity results from the influence of multiple adjacent biomes—such as the Cerrado, Amazon, Atlantic Forest, and Chaco (Tomas et al., 2022), combined with the regulating effect of seasonal flood pulses (Junk et al., 2014). An estimated 2,000 plant species occur in the Pantanal (Pott et al., 2011), of which 234 are arboreal species (Jardim Botânico do Rio de Janeiro, 2025).

Despite its ecological relevance, this ecosystem has been significantly impacted by human activities, including alterations in the carbon cycle, land-use and land-cover change, and the occurrence of large-scale wildfires, such as those recorded in 2020 and 2024, which reached historic levels (Alho et al., 2019; Libonati et al., 2020; Alencar et al., 2024).

Among the main consequences of these impacts is the alteration of the Pantanal’s hydrological dynamics, reflected in reductions in the extent of flooded areas as well as in the duration and intensity of inundation events (Caballero et al., 2025). In this context, riparian formations are particularly sensitive to hydrological changes, as such shifts compromise both the floristic diversity and the ecological functioning of these environments (Oliveira et al., 2014), which are essential for maintaining the integrity of associated aquatic and terrestrial ecosystems.

Given this scenario, forest inventories and phytosociological surveys are essential for flora conservation, as they provide information on the horizontal structure of plant communities, which is fundamental for studies on forest dynamics (Felfili et al., 2005; Junk et al., 2014). A widely used approach for interpreting these data is the analysis of alpha and beta diversity.

Alpha diversity, assessed at a local scale, is an important measure for understanding the complexity of plant communities, including species richness and the degree of uniformity in the distribution of individuals within a given environment (Moura et al., 2021; Whittaker, 1972; Carvalho and Felfili, 2011). Beta diversity, in turn, evaluates variations in floristic composition among sampling units, reflecting the degree of similarity or differentiation among plots and allowing the identification of spatial heterogeneity in vegetation (Ferreira Júnior et al., 2008; Della Coletta and Silveira Vasconcelos, 2022; Whittaker, 1972).

In this context, the objective of this study was to assess the alpha and beta diversity of a riparian vegetation area in the Northern Pantanal by calculating phytosociological parameters, diversity indices, spatial dispersion indices, and similarity indices among plots, in order to address the following questions:

  1. Does the riparian vegetation exhibit structural and floristic composition patterns consistent with those described for riparian formations in the Pantanal?

  2. Is there significant spatial heterogeneity in floristic composition among the sampled plots?

  3. Is the studied area in good ecological conservation status?

2. Materials and Methods

2.1. Study area

The study area is located in the northern region of the Pantanal, within the Sesc Pantanal Private Natural Heritage Reserve (RPPN Sesc Pantanal). The reserve is situated in the municipality of Barão de Melgaço, Mato Grosso State, with central geographic coordinates ranging from 16°28’ S to 16°50’ S and 56° W to 56°30’ W (Figure 1).

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Figure 1
Location of the study area, distribution of the sampled plots and photographic record of the riparian environment in the Northern Pantanal. Note: the blue polygon represents the delimitation of the Pantanal biome.

The riparian vegetation where the sampling plots were established is located along the banks of the Cuiabá River (Figure 1), one of the main watercourses driving the hydrological dynamics of the Northern Pantanal (Luz et al., 2017). The regional climate is classified as Aw according to the Köppen classification system, characterized by a well-defined dry season from May to October and a rainy season from November to April (Alvares et al., 2013). Mean annual temperature is approximately 26.1 °C, and annual precipitation reaches 1,400 mm (Danelichen et al., 2019).

The soil in the study area is classified as an Ebânico Vertisol according to the Brazilian Institute of Geography and Statistics (IBGE, 2025), characterized by high clay content and a dark subsurface horizon (Santos et al., 2025). Local soil analyses indicated a clay content of 50.65%, organic matter content of 9.57 g dm−3, and pH (H2O) of 5.05 (Agostini, 2026).

According to the flood-pulse regime, the region falls within the functional unit “Periodically terrestrial areas,” subclass “Areas with polyspecific forests” (Cunha et al., 2023), characterized by alternating periods of inundation and soil exposure, remaining flooded for only a few weeks annually, which directly influence vegetation composition and structure.

2.2. Data collection and processing

Data collection was conducted in April 2025 through the establishment and measurement of the arboreal component in six rectangular plots measuring 20 m × 30 m (600 m2), totaling 0.36 hectares of sampled area (Figure 1). The plots were distributed along approximately 250 m of the Cuiabá River margin in a riparian forest stretch. Plot distance from the river margin ranged from approximately 30 to 50 m, while the distance between adjacent plots ranged from approximately 20 to 40 m.

Sampling was carried out in April considering regional hydrological and phenological conditions, which are important for accurate species identification, as well as the accessibility of the study area during the sampling period, since access to the site is exclusively by boat.

The floristic survey followed the methodology proposed by Felfili et al. (2005), in which adult tree individuals with a diameter at 1.3 m above the ground greater than 5 cm (d1,3 > 5 cm) were included. Botanical identification was carried out in the field, and unidentified specimens were collected, pressed, and later confirmed at the central herbarium of the Federal University of Mato Grosso. Taxonomic classification followed the Angiosperm Phylogeny Group IV (2016).

Data processing and the calculation of the parameters described in the following sections were performed using Microsoft Excel and PAST software (Hammer et al., 2001).

2.3. Alpha diversity

Alpha diversity of the community was assessed using phytosociological parameters, including density, frequency, dominance, cover value, and importance value. The Importance Value Index (IVI), calculated as IVI = RD + RF + RDo, where RD is relative density, RF is relative frequency, and RDo is relative dominance, was used to identify the species with the greatest ecological importance within the community (Felfili et al., 2011; Magurran, 2013; Souza et al., 2024; Pereira Gomes et al., 2020; Lopes da Rocha et al., 2025). The Cover Value Index (IVC) was calculated as IVC = RD + RDo (Felfili et al., 2011).

In addition, the Shannon diversity index (H’) and Pielou’s evenness index (J’) were calculated, both of which relate species richness to the distribution of individuals among species (Shannon, 1948; Pielou, 1966; 1969; Magurran, 2013). Shannon index values typically range from 1.5 to 3.5, while Pielou’s index varies from 0 to 1, with 1 indicating maximum evenness (Magurran, 2003; Margalef, 1972).

Spatial dispersion patterns of the species were evaluated using the dispersion indices of McGuinnes and Fracker & Brischle, which classify species distribution within the sampled area as random, aggregated, uniform, or showing tendencies toward clustering (Fracker and Brischle, 1944; McGuinnes, 1934; Brower et al., 1998; Melo et al., 2024; Oliveira et al., 2021; Almeida Andrade et al., 2022).

Sampling sufficiency was evaluated through the construction of a rarefaction curve based on plot sampling effort (Sanders, 1968; Hurlbert, 1971), allowing verification of whether the number of sampling units was adequate to represent the floristic richness of the area.

Additionally, the Chao1 richness estimator (Chao, 1987, 1984) was calculated, an index based on the abundance of rare species that estimates total expected richness if sampling effort were sufficiently high (Silva, 2022; Gotelli and Chao, 2013).

2.4. Beta diversity

Beta diversity was assessed based on floristic similarity among the sampled plots using hierarchical cluster analysis (dendrograms). The Sørensen index, which relies on presence–absence data (Sorensen, 1948; Reang et al., 2026; Reddy et al., 2025), and the Bray–Curtis index, which incorporates species abundance, were employed (Bray and Curtis, 1957; Gupta et al., 2025; Silva, 2022).

To interpret the resulting clusters, a Phenon line (cutoff threshold) was set at 60% (0.6) similarity (Milani and Braz, 2012; Sokal and Rohlf, 1962). These indices allow the identification of internal variations in species composition and potential environmental gradients within a single vegetation type, making them useful tools for detecting local-scale heterogeneity (Oliveira et al., 2008; Silva Pinheiro and Ferreira, 2022; Magurran, 2013).

3. Results

3.1. Alpha diversity

A total of 170 tree individuals were sampled, belonging to 19 species—three identified only to the genus level—totaling 19 genera and 12 botanical families (Table 1). The area contained 472.22 trees per hectare and a basal area of 15.94 m2/ha.

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Table 1
Phytosociological parameters of the arboreal component in a riparian forest of the Northern Pantanal, Barão de Melgaço, Mato Grosso, Brazil.

Regarding community diversity, the Shannon index (H’) was 2.48, and Pielou’s evenness index (J’) was 0.84.

Fabaceae was the family with the highest number of species (four), followed by Rubiaceae (two), with the remaining families represented by a single species each. Fabaceae also showed the highest cover value (44.67%) and the highest importance value (41.51%).

The four species with the highest importance values were Pseudalbizzia inundata, Banara arguta, Zygia inaequalis, and Ocotea sp., which together accounted for 50.74% of the importance value index (IVI). The species Salacia elliptica, Cupania vernalis, Alibertia sp., Spondias mombin, and Cordia glabrata showed the lowest IVI values in the community, each represented by only one individual. Among these, three occurred exclusively in plot P1.

Regarding spatial dispersion, most species exhibited a random pattern according to both the McGuinnes and Fracker & Brischle indices (Table 2). Using the McGuinnes index, six species were classified as aggregated, twelve as random, and one as showing a tendency toward clustering. The Fracker & Brischle index indicated a higher number of species with aggregated (six) and clustered (five) patterns, while eight species presented a random distribution.

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Table 2
Spatial dispersion of arboreal species in a riparian forest of the Northern Pantanal, Barão de Melgaço, Mato Grosso, Brazil, according to the McGuinnes and Fracker & Brischle indices.

The species rarefaction curve showed an increasing trend with the addition of sampled plots, without reaching full stabilization at the end of the sampling effort (Figura 2).

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Figure 2
Rarefaction curve of arboreal species based on the number of sampled plots in the riparian vegetation of the Northern Pantanal. Note: the solid line represents observed richness, while dashed lines represent the standard deviation.

However, there was a progressive reduction in the slope of the curve, suggesting an approach to a plateau and indicating that additional plots would likely result primarily in the recording of rare species.

The estimated number of species according to the Chao1 index was 21.78, indicating a potential richness of approximately 22 species for the assessed community.

3.2. Beta diversity

The floristic similarity dendrograms based on the Bray–Curtis and Sørensen indices showed high cophenetic correlation coefficients (r = 0.9785 and r = 0.8863, respectively), indicating good representativeness of the similarity matrices (Figure 3).

thumbnail of Figura 3
Figura 3
Cluster dendrograms of floristic similarity among plots (P1–P6), based on Bray–Curtis and Sørensen indices. The red dashed line indicates the Fenon cut-off line.

In the dendrogram based on the Bray–Curtis index, all plots were grouped at low similarity levels, around 20%, whereas in the Sørensen dendrogram, this grouping occurred at approximately 44% similarity.

Considering a phenetic cutoff level, two clusters were observed in the Bray–Curtis dendrogram and three clusters in the Sørensen dendrogram (Figure 3). In both indices, the grouping of P1 and P2 stood out, indicating greater floristic and structural similarity between these plots compared to the others. Conversely, P5 showed the greatest floristic dissimilarity, being the last to group with the remaining plots.

4. Discussion

4.1. Alpha diversity

The estimated number of trees per hectare and basal area were consistent with those reported for other riparian formations in the Pantanal, which range from 145 to 675.47 individuals per hectare and basal areas between 17.25 and 46.41 m2/ha (Damasceno-Junior, 1997; Moretti et al., 2013; Negrelle, 2013). Similarly, the Shannon index obtained in this study fell within the range reported for riparian vegetation in the biome, from 2.14 to 2.67, while Pielou’s evenness showed a relatively high value, above the reference value of 0.81 previously described for the region, indicating low dominance by a few species (Damasceno-Junior, 1997; Moretti et al., 2013; Negrelle, 2013).

Variations in these parameters reflect the natural heterogeneity of riparian forests in the Pantanal, which are strongly influenced by seasonal flood pulses that regulate species distribution, establishment, and development, shaping vegetation structure and floristic composition over time (Damasceno-Junior and Pott, 2021).

Fabaceae stood out as the most representative family, a recurrent pattern in studies conducted in the Pantanal (Lima Júnior, 2007; Negrelle, 2013; 2016), reinforcing its characterization as the richest and most widely distributed family in the biome (Pott et al., 2011; Pott and Pott, 2021). The species with the highest importance values accounted for more than half of the community’s total IVI and are among the most common species along riverbanks and old river channels in the Pantanal (Santos Vila da Silva et al., 2021).

Among these, Pseudalbizzia inundata (Fabaceae) had the highest relative importance value (13.78%), a pattern expected for a pioneer species widely distributed in periodically flooded areas (Pott and Pott, 1994; Santos Vila da Silva et al., 2021). Its dispersal mechanisms, including autochory and hydrochory, facilitate colonization of wet and marginal environments (Carvalho, 2014; Santos Vila da Silva et al., 2021).

Banara arguta (Salicaceae) and Zygia inaequalis (Fabaceae) also stood out, with IVI values of 12.65% and 12.54%, respectively. Both species are widely distributed in wet and flooded areas of the Pantanal and the Amazon, with dispersal syndromes compatible with the fluvial dynamics of their habitats, such as hydrochory and zoochory (Costa-Pereira et al., 2011; Santos Vila da Silva et al., 2021; Lorenzi, 1998; Pott and Pott, 1994; Muniz, 2020).

These species exhibited aggregated or clustering dispersion patterns, which is expected in Pantanal environments, where communities dominated by one or a few highly adapted species are common due to restrictive environmental conditions, particularly the flood regime (Borges et al., 2014; Manabe and Silva, 2010; Damasceno-Junior and Pott, 2021; Gris et al., 2020; Manrique-Pineda et al., 2021).

In contrast, the five species represented by only one individual can be considered rare within the community (Hernández, 2010; Rios et al., 2023; Silva, 2022). Low abundance levels may reflect ecological preferences for environmental conditions different from those predominant in the area, such as well-drained soils or specific nutrient characteristics, which can limit establishment in environments subject to periodic flooding (Begon and Townsend, 2023; Williams et al., 2009; John et al., 2007; Damasceno-Junior and Pott, 2021; Scremin-Dias et al., 2011).

The rarefaction curve showed a trend toward stabilization, suggesting that most of the area’s species richness had already been sampled and that additional sampling effort would mainly result in the detection of rare species (Schilling and Batista, 2008; Gotelli and Colwell, 2001; Rolim and Nascimento, 1997; Hernández, 2010). The Chao1 richness estimator supports this interpretation, indicating that only a few additional species would be recorded with increased sampling effort, reinforcing the sufficiency of the survey.

The similarity between the structural and floristic patterns observed in this study and those reported in surveys conducted more than a decade ago (Stape et al., 2011) suggests stability in the local riparian vegetation. This persistence may be related to the high adaptive capacity of native species to natural variations in the hydrological regime (Junk et al., 2014; Menezes Mendes et al., 2022).

Although the impacts of climate change on flooding cycles and vegetation structure have already been documented in other areas of the Pantanal (Justino et al., 2024), the results indicate that, to date, this portion of the RPPN Sesc Pantanal maintains a functional ecological composition, demonstrating resilience potential that should be considered in long-term monitoring and conservation strategies.

4.2. Beta diversity

The clustering of plots only at low similarity levels, below the Phenon line, demonstrates weak floristic similarity among sampling units and, consequently, high beta diversity. This pattern indicates species turnover along the analyzed gradient, except for P1 and P2. This result is consistent with the ecological dynamics of riparian environments, which are naturally heterogeneous systems in which hydrological variation contributes to floristic dissimilarity even at small spatial scales (Silva et al., 2020; Scremin-Dias et al., 2011).

The consistent clustering of P1 and P2 in both indices suggests not only shared species but also similar abundance structures, particularly in the case of the Bray–Curtis index.

This high similarity may be associated with spatial proximity and similar environmental conditions, possibly related to the same hydrological influence regime, factors known to determine vegetation composition and structure in riparian environments of the Pantanal (Junk et al., 2014; Soininen et al., 2007).

In contrast, P5 exhibited the highest floristic dissimilarity relative to the other plots, potentially due to its greater distance from the watercourse (Figure 1), reflecting an environmental gradient that influences water availability and, consequently, the composition of tree species (Guilherme et al., 2020; Silva et al., 2020). Additionally, the occurrence of a wildfire near the area in August 2020 (INPE, 2026) may have contributed to the observed differentiation.

In this context, the high beta diversity indicates that the plots are complementary to one another, each contributing a distinct portion of the floristic diversity. Thus, conserving isolated points would be insufficient to represent the total richness of the local riparian vegetation, reinforcing the importance of the protection strategies adopted in the RPPN Sesc Pantanal.

Overall, the patterns of alpha and beta diversity suggest that the studied riparian vegetation is in good conservation status, maintaining structural and functional characteristics typical of preserved environments. This reinforces the importance of protected areas, such as the RPPN Sesc Pantanal, for maintaining biodiversity and serving as reference sites for monitoring environmental changes in riparian ecosystems of the Northern Pantanal.

Acknowledgements

The authors thank the Scientific Initiation Program of the Federal University of Mato Grosso and the Research Support Foundation of the State of Mato Grosso (FAPEMAT) for granting the research scholarship.

We also thank the Nature Conservation Laboratory of the Federal University of Mato Grosso for institutional support and the infrastructure provided.

We express our gratitude to the Sesc Pantanal Private Natural Heritage Reserve for providing transportation and logistical support during field activities.

Data Availability Statement

The dataset analyzed or generated during this study is available upon request from the corresponding author.

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Editor:

Takako Matsumura Tundisi

Publication Dates

  • Publication in this collection
    17 Aug 2026
  • Date of issue
    2026

History

  • Received
    24 Feb 2026
  • Accepted
    28 June 2026
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This is an Open Access article distributed under the terms of the Creative Commons Attribution license (https://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
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