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2019, The Bulletin of the Ecological Society of America
AI
Some nymphalid butterflies obtain their nutrients from fermented fruits or plant saps, forming a guild known as "fruit-feeding butterflies," which serve as biological indicators. This study provides a comprehensive dataset comprising 7,062 records of 279 species across 122 locations in the Atlantic Forest biome, contributing significantly to the understanding of their diversity and distribution. The findings aim to support conservation initiatives and enhance studies related to various ecological processes.
Global Ecology and Biogeography, 2007
Aim To evaluate frugivorous butterflies as indicators of forest disturbance in the Brazilian Atlantic Forest.Location The study area is located in the municipality of Cotia, São Paulo State, south-eastern Brazil (23°35′−23°50′ S, 46°45′−47°15′ W).Methods Sampling was done at four sites inside a large forest block, the Morro Grande State Reserve, and in five forest fragments in an adjacent fragmented landscape. Butterflies were sampled with portable traps, baited with a fermented mixture of banana and sugar cane juice. Sampling was carried out during the period most favourable for the capture of frugivorous butterflies in south-eastern Brazil.Results All richness-related results indicated no effect of forest fragmentation on the frugivorous butterfly guild, concurring with the suggestion of appreciable resistance of Atlantic Forest butterflies to habitat modification. However, species composition discriminated between the two landscapes, indicating that fragmentation may have effects beyond simple species richness. When species were analysed individually, clear patterns of distribution emerged, with some species that were very abundant in the fragmented landscape being practically absent in the continuous landscape, and vice versa. This pattern seems consistent even for some subfamilies.Main conclusion Our findings support the usefulness of the frugivorous butterfly guild as a biological indicator of forest disturbance effects in one of the world's most threatened ecosystems.
Studies on Neotropical Fauna and Environment, 2020
Recently, the availability of cheap and effective survey methods increased the number of forest canopy studies. Several studies have described butterfly communities in the upper forest stratum. However, these studies were all conducted in tall forests (20-60 m canopy height). Here we aim to assess the stratification of fruit-feeding butterfly species in two vertical strata of a semi-deciduous seasonal forest. In addition, we present a field guide to assist in future butterfly monitoring. This study was carried out in two areas of semi-deciduous seasonal forest in the Serra da Bodoquena National Park, Mato Grosso do Sul State, Brazil. Sampling monthly for a year both the, canopy and the understory, we recorded 4229 individuals of 65 species. The species community of fruit-feeding butterflies showed a clear vertical stratification. The composition differed significantly between the two vertical strata, with a higher species richness and abundance in the understory, but less diversity compared to the canopy. Species were distributed heterogeneously between the understory and the canopy in this forest of lower canopy height. Considering published literature of the Mato Grosso do Sul State, the Serra da Bodoquena is one of the most species rich areas with regard to fruit-feeding butterflies.
Journal of Animal Ecology, 2012
1. Documenting species abundance distributions in natural environments is critical to ecology and conservation biology. Tropical forest insect faunas vary in space and time, and these partitions can differ in their contribution to overall species diversity. 2. In the Neotropics, the Central American butterfly fauna is best known in terms of general natural history, but butterfly community diversity is best documented by studies on South American fruit-feeding butterflies. Here, we present the first long-term study of fruit-feeding nymphalid species diversity from Central America and provide a unique comparison between Central and South American butterfly communities.
Papéis Avulsos de Zoologia (São Paulo)
Deforestation has negative impacts on diversity and community patterns of several taxa. In the eastern Amazon, where much deforestation is predicted for the coming years, forests patches may be essential to maintain the local biodiversity. Despite increasing concerns about the conservation of threatened areas, few studies have been performed to analyze the communities of diversified groups, such as insects, in the eastern Amazon. Here, we investigated species diversity and community structure of fruit-feeding butterflies, a well-known bioindicator group, in a threatened remnant of an eastern Amazonian forest located on Maranhão Island, northeastern Brazil. Fruit-feeding butterflies were sampled monthly for one year. Diversity and evenness indices, richness estimators, rarefaction curve, and rank-abundance plot were used to describe community structure in the study area. We captured 529 fruit-feeding butterflies in four subfamilies, 23 genera and 34 species. The three most abundant s...
Research Square (Research Square), 2022
Introduction: Diet and temporal specialists, especially those in tropical seasonal environments, require synchronous phenology with hosts. Species with life-history-traits that covary with temporal specialization may be particularly vulnerable to climate-driven changes in phenology. In the Cerrado, larger fruit-feeding butter ies species tend to be less abundant and more seasonal than their smallerbodied relatives. It is important to consider how ecological process and phylogenetic history are associated with these attributes. Aim/Methods: Here we evaluated the effects of climate and body size on the seasonality of abundance of fruit-feeding butter ies as well as their association with natural history traits including diet breadth and synchrony with fruits. Results: The distribution of abundances of these butter ies species was not uniform throughout the year, was associated with body size and shaped by habitat type and climate. Overall, hot and wet months were associated with higher abundance and species richness of these nymphalids in the Cerrado. Lineages of large-bodied species were less abundant, more seasonal and more generalized in larval diet breadth. In addition, these large-bodied clades were more tightly synchronized with fruit phenology. Discussion: Our ndings support the hypothesis that large and less abundant species to occur in a narrow temporal window and synchronized with fruit phenology. Implications for insect conservation: Particular attention should be given for the conservation of largerbodied temporal specialist species as they are vulnerable to phenology mismatching as a result of rapid global warming, but they will also face additional pressures with the advancing deforestation in the Cerrado.
PloS one, 2017
We provide the first description of the effects of local vegetation and landscape structure on the fruit-feeding butterfly community of a natural archipelago of montane rainforest islands in the Serra do Espinhaço, southeastern Brazil. Butterflies were collected with bait traps in eleven forest islands through both dry and rainy seasons for two consecutive years. The influence of local and landscape parameters and seasonality on butterfly species richness, abundance and composition were analyzed. We also examined the partitioning and decomposition of temporal and spatial beta diversity. Five hundred and twelve fruit-feeding butterflies belonging to thirty-four species were recorded. Butterfly species richness and abundance were higher on islands with greater canopy openness in the dry season. On the other hand, islands with greater understory coverage hosted higher species richness in the rainy season. Instead, the butterfly species richness was higher with lower understory coverage...
Forest Ecology and Management, 2019
Large areas in tropical countries were converted into Eucalyptus plantations for pulp production. Although these plantations are structurally more similar to native ecosystems than traditional short-lived crops, they can be less suitable for more sensitive species. Thus, it is important to know how they can harbor native biodiversity and contribute for its conservation in highly fragmented landscapes. In this work, we compared fruit-feeding butterfly assemblages from Eucalyptus plantations, forest fragments immersed in Eucalyptus plantations and samples inside a continuous forest tract on an extremely rich and threatened area of Atlantic forest in Brazil. We found that plantations harbor a less diverse assemblage of fruit-feeding butterflies, with low richness and a few, very abundant species. Samples placed in plantations were more similar to each other in species composition than those taken from fragments or continuous forests, although the dissimilarities among forest fragments are similar to those found among continuous forest samples. The occurrence of some very abundant species, mostly grassfeeding (Satyrines), differentiate the plantations from forests plots. In common between plantations and forests there were a few other species, notably some associated with second growth (Biblidinae and Satyrinae) and others with strong flying capabilities (Charaxinae). The small fragments harbored a significant portion of the regional butterfly diversity, and this reinforces the importance of actions to preserve them and to increase landscape connectivity for butterfly conservation purposes. It is clear that Eucalyptus sp. plantations cannot substitute forests for a vast majority of fruit-feeding butterflies, but it is better than other land use practices, such as pastures and sugar cane plantations, in sustaining part of this fauna and acting as a potential corridor.
Zoologia, 2019
Due to the important ecological role of Nymphalidae as consumers of fermented fruits, excrement, exudates of decomposed plants and animals, as well as bioindicators of environmental quality that attract the interest of the general public, this study aimed to characterize the fauna of frugivorous butterflies at the Serra Azul State Park – PESA (Mato Grosso, Brazil) with regards to composition, taxonomic richness and spatial distribution. Collections were carried out in 2014 from six 250 m plots in a RAPELD module (5x5 km). We defined five sampling points in each plot and placed a Van Someren-Rydon trap (VSR), with a bait made from banana fermented in cane juice, positioned 1 m above the ground for 24 hours. We collected 204 specimens of Nymphalidae from five subfamilies, 22 genera, and 40 species. Satyrinae was the most abundant and species-rich subfamily. The community structure varied spatially, with the gallery forest presenting the greatest richness, diversity, and equitability, ...
Journal of Applied Ecology, 2007
1. Secondary forests growing on cleared lands and tree plantations are becoming increasingly widespread land-uses in the tropics. Previous studies are divided on the conservation importance of these habitats for tropical forest butterflies. 2. We use a robust sampling design, accounting for both seasonality and vertical stratification, to examine fruit-feeding butterflies (Nymphalidae) in patches of secondary forest and Eucalyptus plantation 2-3 orders of magnitude larger than those previously sampled. 3. We recorded 10 587 butterflies and 128 species in 3200 trap-days. Species richness was highest in primary forest and lowest in plantations, while butterfly abundance showed the opposite response. All habitats were distinct in terms of community structure. 4. There was a significant interaction between habitat and season based on richness and abundance metrics, although not based on community structure. Secondary forest exhibited higher observed richness than primary forest in the peak of the dry-season, but not at other times of the year. This observation could explain the lack of consensus in previous studies, as those reporting higher richness in secondary forest only sampled during the dry-season. 5. In general, habitat quality appeared to be more important than the surrounding landscape in determining butterfly community structure. However, the community structure of the strong-flying Charaxinae was related to the amount of primary forest in surrounding landscape. There was very poor congruence between the response patterns of richness and abundance among different butterfly subfamilies. 6. Linear regressions between resource availability and butterfly abundance showed a strong influence of leaf phenology in both primary and secondary forest, but no influence of fruit phenology. 7. Synthesis and applications : A lack of seasonal replication and small sampling sizes may have led previous studies to overemphasize the conservation importance of secondary forest and plantations for butterflies. We show that these habitats are significantly poorer than primary forest in terms of number of species, and hold distinct butterfly communities. Although quantifying the number of species restricted to primary forest remains difficult, these results cast doubt on the presumption that secondary habitats will provide refuge for many of the species being lost through deforestation. We therefore strongly urge measures that prioritize the conservation of remaining primary forests where they still exist.
Currently, a large-scale restoration project aims to restore around 15 million hectares of Atlantic Forest in Brazil. This will increase forest cover and connectivity among remnant sites as well as restore environmental services. Currently, studies on recovery of fauna in restored areas of the Atlantic Forest are practically nonexistent. To address this knowledge vacuum, our study compares diversity patterns of fruit-feeding butterflies in three forest areas with different restoration ages (11, 22, and 54 years), and uses a native forest area as reference. Results showed butterfly communities in maturing restored areas becoming more similar to the ones found in the native forest, with an increase in the proportional abundance of forest species, and a decrease of edge and grassland species. Moreover, we found a higher diversity among sites at the intermediate restoration age, with a community composed of both grassland and forest species. Butterfly species composition differed significantly among sites, showing interesting patterns of potential species replacement over time. Our results indicate that, although restored sites were located in a fragmented landscape, they provide suitable habitats for recolonization by fruit-feeding butterfly assemblages. Hence, restored areas can be considered important habitat for forest animal species, increasing local biodiversity and, possibly, restoring some of the ecosystem services provided by them.
The Cerrado Biome comprises many habitats, the most common being the savanna type. This biome presents an intense dry season that imposes difficulties on organism survival. The Cerrado’s wet season is the period of greatest abundance and richness of species, including the Lepidoptera. In Brazil, most studies on Nymphalidae seasonality have been conducted in biomes other than the Cerrado. Thus, the pattern of the Nymphalidae seasonality in the Cerrado is unknown. The aim of this study was to describe the temporal dynamics of these butterflies in both savanna and gallery forest habitats in the Cerrado, comparing our findings to those previously reported from other Brazilian biomes, and to discuss the effects of weather and habitat structure on temporal dynamics. Thirty bait-traps were distributed in 2 vertical strata (understory and canopy) in each habitat type (savanna and gallery forest; thus, 60 bait-traps were distributed during 6 d each month from Jul 2012 to Jun 2013. In total, 3,459 individuals of 62 species were captured. The transition between the wet and dry season presented the greatest abundance and species richness and was the period of lowest evenness in the Nymphalidae community. Two distinct assemblages were found in the savanna and gallery forest. Moreover, greater butterfly species turnover was found over time in the gallery forest than in the savanna habitat. These results highlight the importance of climatic seasons, plant phenology, and habitat structure as important predictors of the temporal dynamics of these butterflies in the Cerrado Biome. Furthermore, the floristic and structural distinctiveness of the savanna and gallery forest, and the biological requirements of the butterfly species, could explain a large portion of the species turnover observed in the gallery forest.
Acta Amazonica
Rainfall is one of the most influential factors driving insect seasonality in the Amazon region. However, few studies have analyzed the temporal dynamics of fruit-feeding butterflies in the Brazilian Amazon, specially in its eastern portion. Here, we evaluated the diversity patterns and temporal distribution of fruit-feeding butterflies in a remnant of eastern Amazon forest in the Baixada Maranhense, northeastern Brazil. Specifically, we tested whether fruit-feeding butterflies are temporally structured and whether rainfall influences species richness and abundance. Butterflies were collected with baited traps in both the rainy and dry seasons for two consecutive years. In total, we captured 493 butterflies belonging to 28 species, 15 genera and eight tribes. Three species comprised about half of the overall abundance, and Satyrinae was the most representative subfamily. The fruit-feeding butterfly assemblage showed a strong temporal structure during the second year of sampling, but...
Nature and Conservation, 2023
The present study aimed to describe the composition of the butterfly community in relation to the altitudinal gradient in the Reserva Particular do Patrimônio Natural (RPPN) Alto-Montana, Serra da Mantiqueira, Minas Gerais, Brazil, and to provide a list of species for the area. We collected samples in the RPPN Alto-Montana along an altitudinal gradient from 1400 to 2100 m, between the dry and rainy seasons of 2018 and 2019. During this period, the sampling method utilizing Van-Someren Rydon traps totaled 3,936 hours and the effort using sweep nets totaled 246 hours. A total of 1,253 butterflies distributed across 124 species and six families of diurnal butterflies were observed (Hesperiidae, Lycaenidae, Nymphalidae, Papilionidae, Pieridae, and Riodinidae). Nymphalidae was the most representative family, followed by Hesperiidae, Pieridade, Lycaenidae, Papilionidae and Riodinidae. Moreover, we recorded 37 species with only one individual (singletons) and 20 species with only two individuals (doubletons), totalling 57 species, which corresponds to 46% of all sampled richness. The rarefaction curve did not reveal a tendency toward stabilization. However, the indices showed slightly higher values for the 124 species sampled. The analysis performed using the Bootstrap estimator predicted a total of 143.22 species (± 10.87 SE), with a further 19 additional species than observed. Chao 1 predicted 153.42 (± 11.82 SE), and Jackknife 1 predicted 164.00 (± 16.29 SE) species, with 29 and 40 additional species than the observed, respectively. Our study contributes to the knowledge of butterfly biodiversity in Serra da Mantiqueira and reveals a high species richness for the RPPN Alto Montana, especially considering the relatively small area. In addition, our study provides the first inventory of butterflies for the RPPN Alto Montana, thus supporting further studies investigating the butterfly richness in the Serra da Mantiqueira region. Finally, our findings of endemic, rare, and endangered butterfly species highlight the relevance of further conservation strategies to be considered for the Protected Area's Management Plan.
Florida Entomologist, 2015
The Cerrado Biome comprises many habitats, the most common being the savanna type. This biome presents an intense dry season that imposes difficulties on organism survival. The Cerrado's wet season is the period of greatest abundance and richness of species, including the Lepidoptera. In Brazil, most studies on Nymphalidae seasonality have been conducted in biomes other than the Cerrado. Thus, the pattern of the Nymphalidae seasonality in the Cerrado is unknown. The aim of this study was to describe the temporal dynamics of these butterflies in both savanna and gallery forest habitats in the Cerrado, comparing our findings to those previously reported from other Brazilian biomes, and to discuss the effects of weather and habitat structure on temporal dynamics. Thirty bait-traps were distributed in 2 vertical strata (understory and canopy) in each habitat type (savanna and gallery forest; thus, 60 bait-traps were distributed during 6 d each month from Jul 2012 to Jun 2013. In total, 3,459 individuals of 62 species were captured. The transition between the wet and dry season presented the greatest abundance and species richness and was the period of lowest evenness in the Nymphalidae community. Two distinct assemblages were found in the savanna and gallery forest. Moreover, greater butterfly species turnover was found over time in the gallery forest than in the savanna habitat. These results highlight the importance of climatic seasons, plant phenology, and habitat structure as important predictors of the temporal dynamics of these butterflies in the Cerrado Biome. Furthermore, the floristic and structural distinctiveness of the savanna and gallery forest, and the biological requirements of the butterfly species, could explain a large portion of the species turnover observed in the gallery forest.
EntomoBrasilis, 2024
Conservation Units are areas legally established by the government with the goal of conserving territory and its natural resources. Given the limited knowledge about the invertebrate biodiversity in the Northern Atlantic Forest, including its legally protected areas, the present study aimed to inventory butterfly species in two Conservation Units of the Northern Atlantic Forest in Paraíba, Brazil: Engenho Gargaú Private Natural Heritage Reserve (RPPN Gargaú) and Mata do Xém-xém State Park (PE Xém-xém), both located in the metropolitan region of João Pessoa. We conducted the collections every two months, over three consecutive days, in six expeditions from February 2013 to April 2014 in RPPN Gargaú and in four expeditions from August 2014 to April 2015 in PE Xém-xem. Additionally, a two-day collecting expedition was carried out in August 2021 in RPPN Gargaú. We employed two sampling methods: Van Someren-Rydon traps and entomological nets. A total of 212 species (2,841 specimens) were recorded, 158 (1,867 specimens) in RPPN Gargaú and 129 (974 specimens) in PE Xém-xém. Overall, Hesperiidae was the richest family (81 spp.), followed by Nymphalidae (70), Riodinidae (22), Lycaenidae (22), Pieridae (12), and Papilionidae (4). Out of the total species, 48 are new records for Paraíba, and seven for northeast Brazil. Notably, the record of Morpho menelaus eberti Weber, 1963 in RPPN Gargaú, a subspecies classified as Critically Endangered (CR), stands out. The results emphasize the need to protect the forest remnants in the region, as well as to develop management and monitoring actions for butterflies and other invertebrates.
2010
The Atlantic Forest deserves special attention due to its high level of species endemism and degree of threat. As in other tropical biomes, there is little information about the ecology of the organisms that occur there. The objectives of this study were to verify how fruit-feeding butterflies are distributed through time, and the relation with meteorological conditions. Species richness and Shannon index were partitioned additively at the monthly level, and b diversity, used as a hierarchical measure of temporal species turnover, was calculated among months, trimesters, and semesters. Circular analysis was used to verify how butterflies are distributed along seasons and its relation with meteorological conditions. We sampled 6488 individuals of 73 species. Temporal diversity of butterflies was more grouped than expected by chance among the months of each trimester. Circular analyses revealed that diversity is concentrated in hot months (September-March), with the subfamily Brassolinae strongly concentrated in February-March. Average temperature was correlated with total abundance of butterflies, abundance of Biblidinae, Brassolinae and Morphinae, and richness of Satyrinae. The present results show that 3 mo of sampling between September and March is enough to produce a nonbiased sample of the local assemblage of butterflies, containing at least 70 percent of the richness and 25 percent of abundance. The influence of temperature on sampling is probably related to butterfly physiology. Moreover, temperature affects resource availability for larvae and adults, which is higher in hot months. The difference in seasonality patterns among subfamilies is probably a consequence of different evolutionary pressures through time.
Biological Journal of The Linnean Society, 1999
To test the hypotheses that butterflies in an intact lowland rainforest are randomly distributed in space and time, a guild of nymphalid butterflies was sampled at monthly intervals for one year by trapping 883 individuals of 91 species in the canopy and understory of four contiguous, intact forest plots and one naturally occurring lake edge. The overall species abundance distribution was well described by a log-normal distribution. Total species diversity ( -diversity) was partitioned into additive components within and among community subdivisions ( -diversity and -diversity) in vertical, horizontal and temporal dimensions. Although community subdivisions showed high similarity (1--diversity/ -diversity), significant -diversity existed in each dimension. Individual abundance and observed species richness were lower in the canopy than in the understory, but rarefaction analysis suggested that the underlying species richness was similar in both canopy and understory. Observed species richness varied among four contiguous forest plots, and was lowest in the lake edge plot. Rarefaction and species accumulation curves showed that one forest plot and the lake edge had significantly lower species richness than other forest plots. Within any given month, only a small fraction of total sample species richness was represented by a single plot and height (canopy or understory). Comparison of this study to a similar one done in disturbed forest showed that butterfly diversity at a naturally occurring lake edge differed strongly from a pasture-forest edge. Further comparison showed that species abundance distributions from intact and disturbed forest areas had variances that differed significantly, suggesting that in addition to extrapolation, rarefaction and species accumulation techniques, the shapes of species abundance distributions are fundamental to assessing diversity among sites. This study shows the necessity for long-term sampling of diverse communities in space and time to assess tropical insect diversity among different areas, and the need of such studies is discussed in relation to tropical ecology and quick surveys in conservation biology.
Biota Neotropica, 2019
This study aimed to present a list of the species of frugivorous butterflies occurring in Atlantic Forests, in the Conservation Units: National Forest of Chapecó (FLONA), Ecological Station of Mata Preta (ESEC) and State Park of Araucárias (PAEAR) and adjacent forest fragments, located in the western region of the state of Santa Catarina. Three samplings were conducted between December 2017 and March 2018, totaling 24 days of collection in each sampling area. Van Someren-Rydon traps were used to capture frugivorous butterflies. There were 4,231 frugivorous butterflies belonging to four subfamilies, 12 tribes and 49 species. In all, 37 species of frugivorous butterflies were sampled in FLONA and 29 in adjacent forest fragments. In ESEC, 29 species and 33 in adjacent forest fragments. In PAEAR, 33 species and 28 in adjacent forest fragments. Of the total species registered, 15 species are new records for the state of Santa Catarina and 11 are new records for the western region of the state. The most abundant species for FLONA were: Manataria hercyna (Hübner, 1821) and Hermeuptychia sp. In ESEC, were Hermeuptychia sp. and Yphthimoides ordinaria (Freitas, Kaminski & Mielke, 2012). In PAEAR, greater abundance of Forsterinaria quantius (Godart, 1824) and Eryphanes reevesii (Doubleday, 1849) were verified. For the adjacent forest fragments to Conservation Units, there was a greater abundance of Hermeuptychia sp., Moneuptychia soter (Butler, 1877), Morpho epistrophus (Fabricius, 1796) e Forsterinaria quantius (Godart, 1824). Satyrinae presented higher richness (S = 34) and abundance (90.58%) in all areas sampled. The rarefaction and extrapolation curves for the Conservation Units and adjacent forest fragments showed a greater rise in the FLONA and PAEAR sampling units and their adjacent forest fragments. The estimated sampling coverage for Conservation Unit and forest fragments was above 97%. The richness calculated through the Jackknife 1 estimator, for the FLONA and PAEAR samplings, presented a value of 50.75 and 37.09, respectively. The fauna of frugivorous butterflies from this region, first investigated in areas of Conservation Units, showed to be expressive and well represented in the Atlantic Forest Biome, indicating its potential as a refuge for biodiversity.
A survey of 6,000 trap/hours using fruit-bait traps was conducted, in order to characterize the community of fruit-feeding butterflies and their seasonal variation in a semiarid area of NE Brazil, which exhibits a highly seasonal rainfall regime. The community was composed of 15 species, the four most abundant comprising more than 80% of the total individuals. In the first sampling month, 80% of the species had already been recorded. A strong positive correlation was found between butterfly abundance and precipitation of the sampled year. The highly seasonal pattern of the studied community may be related to the availability of both larvae and adult food resources. The endemic Fountainea halice moretta presented the highest positive correlations with other butterfly species and with total butterfly abundance and illustrates well the seasonal variation of the whole community. Hamadryas februa was the only species trapped in all sampling months, probably due to the ability of adults to aestivate during the dryer months. The open and more disturbed sample area showed higher abundance of butterflies in the rainy season and lower abundance in the dry season. Both abundance and richness were affected by season in a manner that the rainy period had significantly more individuals and species.
To test the hypotheses that fruit-feeding nymphalid butterflies are randomly distributed in space and time, a community of fruit-feeding nymphalid butterflies was sampled at monthly intervals for one year by trapping 6690 individuals of 130 species in the canopy and understory of four forest habitats: primary, higraded, secondary, and edge. The overall species abundance distribution was well described by a lognormal distribution. Total species diversity (diversity) was partitioned into additive components within and among community subdivisions (diversity and-diversity) in vertical, horizontal and temporal dimensions. Although community subdivisions showed high similarity (1−-diversity/-diversity), significant diversity existed in each dimension. Individual abundance and observed species richness was lower in the canopy than in the understory. However, rarefaction analysis and species accumulation curves revealed that canopy had higher species richness than understory. Observed species richness was roughly equal in all habitats, but individual abundance was much greater in edge, largely due to a single, specialist species. Rarefaction analysis and species accumulation curves showed that edge had significantly lower species richness than all other habitats. Samples from a single habitat, height and time contained only a small fraction of the total community species richness. This study demonstrates the feasibility, and necessity, of large-scale, long-term sampling in multiple dimensions for accurately measuring species richness and diversity in tropical forest communities. We discuss the importance of such studies in conservation biology.
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