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Article

Vertical Stratification of Butterfly Assemblages Persists in Highly Disturbed Forest Fragments of the Brazilian Atlantic Forest

by
Denise B. Silva
1,
André V. L. Freitas
2,
Oscar F. Junior
1 and
Jessie P. Santos
2,*
1
Faculdades Integradas de Fernandópolis, Fundação Educacional de Fernandópolis, Fernandopolis 15608-380, Sao Paulo, Brazil
2
Departamento de Biologia Animal, Instituto de Biologia, Universidade Estadual de Campinas, Campinas 13083-862, Sao Paulo, Brazil
*
Author to whom correspondence should be addressed.
Diversity 2024, 16(10), 608; https://doi.org/10.3390/d16100608
Submission received: 3 September 2024 / Revised: 20 September 2024 / Accepted: 22 September 2024 / Published: 1 October 2024
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Abstract

:
Vertical stratification is a property of forest habitats related to the differential distribution of organisms according to the variation in the conditions, from the understory to the canopy. Here, we aimed to test whether butterfly assemblages from highly disturbed forests maintain the pattern of vertical stratification. We hypothesized that degraded forests would not exhibit vertical stratification due to the low variation in the microhabitat conditions along the vertical gradient, resulting from the canopy openness. To test this, we sampled fruit-feeding butterflies with bait traps, alternately disposed between the understory and canopy of three secondary forest fragments in a very fragmented Atlantic Forest landscape, for one year. We found that the vertical strata differed in terms of species composition, with a high contribution by the nestedness component on the beta diversity spatial variation. The understory assemblages had a higher abundance and were more diverse than the upper stratum. We demonstrated that vertical stratification is maintained even in disturbed forests; however, this does not necessarily provide support for a good quality and functioning ecosystem in these habitats. The butterfly assemblages recorded here are a subset of the species pool that inhabits conserved remnants. Thus, even being represented by species commonly found in disturbed habitats, the dynamic of vertical stratification of assemblages remains.

1. Introduction

Land use involves the conversion of natural habitats into landscapes of anthropic usage and is recognized as the primary factor behind biodiversity loss [1,2]. Modified habitats undergo several changes to their properties that can significantly impact the biodiversity that resides in them. In tropical and subtropical regions, land use is responsible for the fragmentation of natural forest areas [3]. The fragmentation divides entire forest ecosystems into small patches, increasing edge effects [4,5]. In addition, deforestation reduces the canopy cover, which increases the temperature and reduces humidity [6]. These changes in the microclimate conditions consequently lead to significant changes in the composition of local assemblages [7].
A key property in terms of the maintenance of high diversity in forest ecosystems is the vertical gradient of the conditions that allow the habitat partition between co-occurring species. The differential distribution of species along vertical forest strata is known as vertical stratification, a pattern recognized as important for several animal groups [8,9,10,11]. Previous studies indicate that butterfly species consistently partition the vertical forest strata regarding their foraging habits, to such a point that the understory and canopy can be considered two independent ecological assemblages [12,13,14,15,16]. Although forest clearings and edges allow canopy species to reach the ground [16], low-impact activities, such as the opening of small trails for example, are not sufficient to promote changes in vertical stratification patterns [17]. On the other hand, forest remnants within highly fragmented landscapes have modified habitat conditions that could potentially disrupt the vertical distribution of assemblages, but this assumption requires investigation.
The Brazilian Atlantic Forest (AF) is a tropical hotspot that has been suffering from intense degradation since the colonization period [18]. Deforestation processes in the AF are clearly influenced by altitude [19], since the larger forest remnants are in terrains with high elevation [18]. In contrast, the natural vegetation of flatter terrains has changed substantially due to the expansion of agricultural and urban areas [20,21]. This is the case in the western limits of the AF, where the regional landscape was massively converted for livestock grazing, crop cultures, and other agricultural activities [21]. Notwithstanding, most of the remaining natural vegetation in the region is located in riparian protected areas, or forest fragments smaller than 50 ha that are highly isolated from each other [20]. Consequently, this region is biologically depleted, frequently pointed out as a species-poor area [22,23] and limited in terms of ecosystem services provision compared to other Atlantic Forest regions [23,24].
In the current study, we sampled butterfly assemblages from disturbed forest fragments within a heavily modified landscape in the Atlantic Forest to investigate whether they exhibit vertical stratification. We hypothesized that due to the significant alterations to these habitats, there would be reduced variation in the environmental conditions across the vertical strata, resulting in relatively uniform butterfly assemblages. Thus, we expect that canopy and understory assemblages do not differ in terms of their community parameters, such as their abundance, species richness, diversity, and species composition.

2. Materials and Methods

2.1. Study Area

We performed the study in three forest remnants of a secondary semideciduous seasonal forest in the Turmalina municipality, Sao Paulo State, southeastern Brazil (Figure 1A). According to the Atlantic Forest integral delimitation, this region is near the western limits of the biome [25]. The region consists of a mosaic of forest fragments, riparian forests, and transitional areas between the Cerrado savannas and the Atlantic Forest, within a severely fragmented landscape. The climate in this region is classified as humid mesothermal, with dry winters and wet summers. The hot season spans from October to March, with milder months from April to September; the mean annual temperature is 26.5 °C and the annual rainfall ranges from 875 to 1475 mm [26]. The fragments were highly disturbed, with a very low open canopy (5 to 7 m high and emergent trees taller than 10 m) and large clearings allowing a large amount of sunlight to reach the understory (Figure 1C).

2.2. Butterfly Sampling

We conducted monthly butterfly samplings from September 2017 to August 2018. We selected three forest fragments, in order to establish one linear transect (about 400 m length) in each of them. To compare understory and canopy assemblages, we installed 10 Van-Someren bait traps, at least 30 m apart from each other, in each transect (a total of 30 traps), alternating between the understory (1.5 m above the ground; Figure 1B) and the canopy (from about 6 m in height). We adopted alternate disposition of the traps to avoid interference due to the attractivity of the neighboring traps to species from the same stratum [12]. The bait consisted of a mixture of banana and sugar cane juice, which was fermented for 48 h before sampling. The traps were checked every 24 h, with the bait replaced at each visit. We kept all the traps simultaneously open in the field (four complete sampling days for each trap per month). We collected all the butterfly specimens in entomological envelopes and carried out the mounting and identification in the Laboratory of Ecology and Systematics of Butterflies at Unicamp in Campinas, Brazil.

2.3. Data Analyses

We assessed the differences in the butterfly assemblages between the vertical strata, using the species richness, abundance, diversity, and species composition as parameters in our analyses. First, we compared the abundance data with generalized linear mixed models (GLMMs) with Poisson distribution, evaluating the direct effect of the vertical strata, the sampled month, and their interaction (treated as fixed effects). Transects were used as random factors in our model. For representing the differences in terms of abundance, we plotted the mean abundance of each vertical stratum per month. These analyses were performed with vegan and lme4 packages in R software version 4.4.1 [27,28,29].
To test whether the species richness differed among the vertical strata, we calculated interpolation and extrapolation curves (with 999 bootstrap resampling), based on the number of individuals and the sample coverage [30,31]. The combined use of these approaches allows more robust inferences and guarantees sampling quality in comparisons with the species richness [32]. We also performed diversity profile curves, using Hill numbers to test the differences in diversity between the understory and canopy. The order of diversity indicates its sensitivity in regard to common and rare species; the most common species are favored as the q-value increases, then for values less than unity (q < 1), the diversity profile gives more weight to the rare species [33]. Orders of diversity (q values) are analogous to species richness (q = 0), the Shannon index (q = 1), and Simpson’s index (q = 2). Interpolation/extrapolation rarefactions and diversity profiles were calculated using the iNEXT package [31].
We used the abundance matrix with Hellinger-transformed data to calculate a similarity matrix with the Bray–Curtis distance. Then, we carried out betadisper analysis with 999 permutations, to check whether there was any heterogeneity in the variance between the groups. Finally, we performed a PERMANOVA to test the differences in the species composition between the vertical strata. We used non-metric multidimensional scaling (NMDS) to visually inspect the formed groups. These analyses were performed using the vegan package [27]. To identify species associations with each vertical stratum, we used the IndVal index [34]. This analysis is based on species fidelity and specificity, with fidelity being the sample estimate of the probability of finding the species in sites belonging to a given site group. The specificity component is a sample estimate of the probability that the surveyed site belongs to the target site group, given that the species has been found. The IndVal analysis was performed with the indicspecies package [34].
We applied the SDR simplex approach to partition the beta diversity of each vertical stratum and between them [35]. This method is based on the Ruzicka dissimilarity matrix and divides beta diversity into three components: the similarity of the species abundance between pairs of sites (S), the abundance difference (D), which indicates the loss or gain of abundance in the sites, and the number of individuals in the replacement species, plus the number of individuals in the species replaced between two sites (R) [35]. In addition, the SDR simplex approach can be used to obtain the nestedness component (N), which is the sum of the similarity and abundance difference (N = S + D), and that corresponds to the inverse of the replacement. The sum of the replacement (R) and abundance difference (D) results in the total dissimilarity (β-diversity) between a pair of sites (β = R + D), while the similarity (S) is 1 minus the total dissimilarity (S = 1 − β). The sum of the three components is always equal to 1 (S + D + R = 1), thus the components can be interpreted as percentages, considering the relative importance of each component regarding beta diversity [35]. Finally, we used permutation tests (1000 permutations) to check the significance of the components. The SDR analysis was performed using the vegan package [27].

3. Results

In total, we captured 5303 individuals, distributed across 46 butterfly species, from four subfamilies (Table S1). The most representative subfamily was Biblidinae and the three most abundant species, Eunica tatila, Callicore astarte, and Eunica maja, belonged to this subfamily. The most abundant species, E. tatila, represented about 75% of the total abundance in the assemblages (N = 3972), being dominant in both vertical strata. However, understory assemblages had higher butterfly abundance (N = 3121) than canopy assemblages (N = 2182). The GLMM analysis revealed a significant effect of the vertical strata in terms of the abundance distribution and detected variations in the abundance between different months (Table 1; Figure 2). The hot season months had higher abundance than the milder months and, in some cases, the abundance did not differ between the vertical strata within the sampled month (Figure 2).
Despite the higher abundance in the understory, the species richness was statistically the same in both vertical strata (Figure 3). We recorded 14 species captured exclusively in the understory, while nine were only captured in canopy traps. Of all the recorded species, 12 were singletons, with six only captured once in the canopy and six only captured once in the understory. Although the species richness did not differ between the vertical strata, understory assemblages were more diverse when considering the more weight applied for common species, according to the diversity profiles (Figure 4).
We confirmed the homogeneity in the variance among the groups in the betadisper analysis (p = 0.916), thus the species composition of fruit-feeding assemblages was distinct between the vertical strata and due to the differences in the group positions (F = 10.93; p = 0.001). The canopy and understory formed two groups in the ordination space of the NMDS, with good quality (Figure 5; stress = 0.178). According to the SDR simplex approach, the most relevant component of beta diversity was the similarity among the assemblages (Figure 6); however, the similarity index values differed statistically, being higher in the understory (F = 7.64; p = 0.007). The replacement component contributed less than 20% in terms of both vertical strata, which means that they had up to 80% of nested assemblages (Figure 6). For the pairwise comparison between the canopy and understory traps, we found that the mean similarity contributed to more than half of the variation between the two strata (S = 0.53). This means that the canopy and understory share species with similar abundance. The replacement (R = 0.17) and abundance difference (D = 0.29) components had low contributions (Figure S1). However, these values in terms of abundance variation and turnover were sufficient to attest to the difference in species composition between the vertical strata.
We detected two species indicators of canopy habitats (Callicore sorana and Haematera pyrame) and eight indicators of understory habitats (Table 2). At least three understory species presented high values of specificity and fidelity (Archaeoprepona demophon, Eunica maja, and Hamadryas chloe). This means that the probability of capturing these species in the understory stratum is high and the traps that sampled these species also have a high probability of belonging to forest understory samplings.

4. Discussion

In the present study, we tested whether the vertical stratification in fruit-feeding butterfly assemblages is maintained in highly disturbed forests. Although the species richness did not differ between the vertical strata, we found a higher abundance and diversity in understory assemblages. Additionally, we observed distinct species compositions in the canopy and the understory, with the nestedness component significantly contributing to the beta diversity in both vertical layers. These findings underscore the importance of the vertical structure of forest habitats in maintaining butterfly diversity, but also raise concerns about the consequences of forest landscape fragmentation.
This is the first study to report on the vertical stratification of butterfly assemblages distributed in a highly disturbed Atlantic Forest landscape. There is no consensus on which vertical stratum has the most species richness in tropical forests, but understory assemblages usually have higher abundance values in warm neotropical forests [16,36,37]. Montane forests are an exception in this regard, when the canopy experiences more stable temperatures throughout the year, especially in the cold season [13,38]. Butterfly assemblages from our study site presented parameters that resemble those observed in warm seasonal forests, with a low range of temperature variation between the vertical strata and over time [37]. Therefore, the difference in the mean temperature between the vertical strata does not seem to limit the stratification dynamics of the species in the present study, where the absence of a cold season provides high temperatures in both strata throughout the year.
The landscape where the present study was carried out is highly modified, mostly dominated by agriculture and cattle ranching, with less than 5% of forest remnants [39]. The low connectivity among the forest remnants in this region hampers the dispersion of forest species among the fragments, thus the biodiversity patterns are more related to the fragment size [40]. In a recent niche modeling analysis, about 50 fruit-feeding butterfly species were estimated to be present in the studied region, according to climatic variables. However, considering only the landscape features, such as the forest cover percentage and functional connectivity, the richness predicted falls to a maximum of 20 species [24]. The combined models (climatic and landscape) predicted a range of 50 to 100 species, which represents a medium species richness number considering the total fruit-feeding species recorded in the Atlantic Forest [24]. Accordingly, the reported richness of 46 butterfly species found in the studied region nearly matches the prediction of the combined models. But, more importantly, it demonstrates the pervasive effects of landscape modification on the study site in contrast with other AF regions with higher species richness.
We found that canopy assemblages had higher dissimilarity than those in the understory, but the nestedness component significantly contributed to the beta diversity in both strata. Previous studies in the Atlantic Forest have demonstrated that understory assemblages have higher species turnover, relating this pattern to a phylogenetic constraint on some butterfly clades that have evolved in the understory of forests [13,38]. For example, the Morphini and Brassolini tribes are adapted to closed forests and their life cycles are temporally synchronized with the seasons offering high resource availability [41]. In our study, we recorded no specimens from the Morphini tribe and only three species of Brassolini, of which the most abundant is frequently captured in the canopy (Opsiphanes invirae). Despite belonging to the clade that evolved in closed forests, O. invirae feeds on tall palm trees (Table S1) and frequently flies high in open areas at twilight, which explains its frequency in canopy traps. Conversely, other Brassolini species sampled in the present study site, such as Caligo illioneus, feed mostly on understory plants and low palms (Table S1) and present a low level of flight (usually below 2 m above ground) and were not captured in the canopy traps. Species trait evolution is the product of an intricate net of interactions; thus, the interpretation of patterns should not be based on a single trait. Nevertheless, we can assume that vertical stratification dynamics depend on the identity of the species that compose butterfly assemblages.
The butterfly assemblages recorded here are mostly represented by species commonly found in secondary formations or disturbed forests, such as species from the genus Hamadryas and the most abundant species, E. maja [42]. We found an association between these species and the understory, which highlights the quality of the forest habitat in the study area. Two species that had a high fidelity with canopy assemblages, C. sorana and H. pyrame, belong to the same tribe and are closely related in terms of their life history; individuals from these species are frequently sighted landed in wet mud and their larvae feed on some vine species (Sapindaceae, see Table S1), frequently found in the canopy of secondary forests [43].
Based on previous studies and the contribution made by our results, we elaborated insights to elucidate a pivotal question considering the scenario of fragmented forests: How much disturbance is necessary to disrupt the vertical stratification of butterfly assemblages? Although the disturbance intensity may compromise the heterogeneity of the forest structure, the vertical stratification patterns seem to be regulated dynamically. However, a factor that leads to a huge decrease in species from the assemblage may exceed this self-regulation threshold and possibly fade out the stratification pattern. This effect can be measured through the variation in the beta diversity component values. Here, we found that the high degree of nestedness in both the vertical strata indicates that butterfly communities consist of species commonly found in disturbed habitats [9,42,44]. As a result, these forest fragments have lower diversity and provide fewer ecosystem services when compared to conserved sites in the Atlantic Forest, even though the vertical stratification of butterfly assemblages remains.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/d16100608/s1, Table S1: List of fruit-feeding butterfly species recorded at the study site and their abundance in each vertical stratum. Larval host plant families of all the species are listed. In the column, the “aspect” refers to the plants used by the larvae; for example, although the host plants of H. clytemnestra and C. dirce are trees, larvae are only found when these plants are small shrubs. Figure S1: SDR simplex approach for spatial beta diversity of pairwise comparison between canopy and understory obtained through the Ruzicka dissimilarity matrix. The continuous lines inside each triangle represent the mean value for each component; the dashed lines represent the confidence interval (0.95).

Author Contributions

Conceptualization, D.B.S., A.V.L.F., O.F.J. and J.P.S.; methodology, D.B.S. and J.P.S.; software, J.P.S.; validation, D.B.S., A.V.L.F., O.F.J. and J.P.S.; formal analysis, J.P.S.; investigation, D.B.S., A.V.L.F. and J.P.S.; resources, D.B.S., A.V.L.F., O.F.J. and J.P.S.; data curation, D.B.S. and J.P.S.; writing—original draft preparation, D.B.S., A.V.L.F. and J.P.S.; writing—review and editing, D.B.S., A.V.L.F. and J.P.S.; visualization, D.B.S., A.V.L.F. and J.P.S.; supervision, J.P.S.; project administration, J.P.S.; funding acquisition, D.B.S., A.V.L.F., O.F.J. and J.P.S. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by the Conselho Nacional de Desenvolvimento Científico e Tecnológico—CNPq (Fellowship 304291/2020-0); FAPESP (grant 2021/03868-8); JPS researcher grant is supported by the Universidade Estadual de Campinas [Resolution GR-033/2023].

Institutional Review Board Statement

Not applicable.

Data Availability Statement

The data presented in this study are available on request from the corresponding author.

Acknowledgments

We thank the anonymous reviewers, the landowners of “Fazenda Palmeirinha”, Roberto, Renato, and Rosana for authorization to access the fragments and perform this work. Fernando Dias helped with the identification of several species of Charaxinae. Augusto H. B. Rosa helped during the fieldwork. The current study is registered in SISGEN (N°: A75DBE8).

Conflicts of Interest

The authors declare that there are no conflicts of interest. The funders had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript, or in the decision to publish the results.

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Figure 1. (A) Map with the location of the three sampling transects (red circles) projected onto the forest remnants in the study site. The orange outline represents the Atlantic Forest domain, according to the integrative limit proposed by Muylaert et al. 2018; (B) sampling transect with a bait trap placed in the understory; (C) a transect photo to illustrate the canopy structure of the sampling site.
Figure 1. (A) Map with the location of the three sampling transects (red circles) projected onto the forest remnants in the study site. The orange outline represents the Atlantic Forest domain, according to the integrative limit proposed by Muylaert et al. 2018; (B) sampling transect with a bait trap placed in the understory; (C) a transect photo to illustrate the canopy structure of the sampling site.
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Figure 2. Abundance of fruit-feeding butterflies recorded in each vertical stratum per month (mean ± SE).
Figure 2. Abundance of fruit-feeding butterflies recorded in each vertical stratum per month (mean ± SE).
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Figure 3. (a) Individual-based rarefaction curves comparing the species richness of fruit-feeding butterfly assemblages between the canopy and understory. The dashed line represents the interpolation where the number of individuals is the same for both vertical strata. The blue dashed line is the extrapolated number of individuals for the canopy. (b) Coverage-based curves comparing the species richness between the canopy and understory.
Figure 3. (a) Individual-based rarefaction curves comparing the species richness of fruit-feeding butterfly assemblages between the canopy and understory. The dashed line represents the interpolation where the number of individuals is the same for both vertical strata. The blue dashed line is the extrapolated number of individuals for the canopy. (b) Coverage-based curves comparing the species richness between the canopy and understory.
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Figure 4. Diversity profiles calculated with Hill numbers to compare the diversity between the canopy and understory in regard to different q values.
Figure 4. Diversity profiles calculated with Hill numbers to compare the diversity between the canopy and understory in regard to different q values.
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Figure 5. Non-metric multidimensional scaling (NMDS) demonstrating the difference in species composition between the canopy (circles) and understory (triangles).
Figure 5. Non-metric multidimensional scaling (NMDS) demonstrating the difference in species composition between the canopy (circles) and understory (triangles).
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Figure 6. SDR simplex approach for spatial beta diversity within each vertical stratum obtained through the Ruzicka dissimilarity matrix. Black lines inside each triangle represent the mean values for each component and the black points are the mean values for all the three components. Dashed lines represent the confidence intervals (0.95). (a) SDR simplex for pairwise canopy traps; (b) SDR simplex for pairwise understory traps.
Figure 6. SDR simplex approach for spatial beta diversity within each vertical stratum obtained through the Ruzicka dissimilarity matrix. Black lines inside each triangle represent the mean values for each component and the black points are the mean values for all the three components. Dashed lines represent the confidence intervals (0.95). (a) SDR simplex for pairwise canopy traps; (b) SDR simplex for pairwise understory traps.
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Table 1. Results of generalized linear mixed models (GLMMs), testing the effects of fixed factors on the abundance of butterfly assemblages. Transects were modeled as a random factor. SE = standard error; Z = parameter estimated.
Table 1. Results of generalized linear mixed models (GLMMs), testing the effects of fixed factors on the abundance of butterfly assemblages. Transects were modeled as a random factor. SE = standard error; Z = parameter estimated.
PredictorsEstimateSEZp Value
(Intercept)4.17420.183622.7<0.001
Understory−0.34830.1088−3.20.001
Month—February0.24630.09342.60.008
Month—March−0.69310.1212−5.7<0.001
Month—April−1.40600.1577−8.91<0.001
Month—May−0.79630.1255−6.3<0.001
Month—June−0.89920.1301−6.9<0.001
Month—July−0.30740.1075−2.80.004
Month—August−2.22700.2243−9.9<0.001
Month—September0.15830.09531.60.096
Month—October−0.24920.1057−2.30.018
Month—November0.60880.08696.9<0.001
Month—December0.80870.08419.6<0.001
Understory: February0.84520.13416.2<0.001
Understory: March0.83530.16635.0<0.001
Understory: April0.83070.21013.9<0.001
Understory: May0.67060.17483.8<0.001
Understory: June−0.70320.2415−2.90.003
Understory: July0.02900.16630.10.861
Understory: August1.01840.28373.5<0.001
Understory: September1.07030.13437.9<0.001
Understory: October0.40330.15512.50.009
Understory: November0.95800.12637.5<0.001
Understory: December0.66710.12495.3<0.001
Table 2. Indicator value index (IndVal) of indicative butterfly species in each vertical strata and their respective specificity and fidelity values.
Table 2. Indicator value index (IndVal) of indicative butterfly species in each vertical strata and their respective specificity and fidelity values.
SpeciesVertical StrataSpecificityFidelityIndValp Value
Archaeoprepona demophonUnderstory0.8670.8750.8440.001
Callicore soranaCanopy0.4001.0000.6320.026
Catoblepia berecynthiaUnderstory0.5330.9330.7060.008
Eunica majaUnderstory1.0000.9050.9510.001
Haematera pyrameCanopy0.3331.0000.5770.038
Hamadryas chloeUnderstory0.9331.0000.9660.001
Hamadryas februaUnderstory0.6661.0000.8160.002
Pareuptychia ocirrhoeUnderstory0.3331.0000.5770.033
Taygetis lachesUnderstory0.6660.9580.7990.002
Taygetis rufomarginataUnderstory0.3331.0000.5770.048
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MDPI and ACS Style

Silva, D.B.; Freitas, A.V.L.; Junior, O.F.; Santos, J.P. Vertical Stratification of Butterfly Assemblages Persists in Highly Disturbed Forest Fragments of the Brazilian Atlantic Forest. Diversity 2024, 16, 608. https://doi.org/10.3390/d16100608

AMA Style

Silva DB, Freitas AVL, Junior OF, Santos JP. Vertical Stratification of Butterfly Assemblages Persists in Highly Disturbed Forest Fragments of the Brazilian Atlantic Forest. Diversity. 2024; 16(10):608. https://doi.org/10.3390/d16100608

Chicago/Turabian Style

Silva, Denise B., André V. L. Freitas, Oscar F. Junior, and Jessie P. Santos. 2024. "Vertical Stratification of Butterfly Assemblages Persists in Highly Disturbed Forest Fragments of the Brazilian Atlantic Forest" Diversity 16, no. 10: 608. https://doi.org/10.3390/d16100608

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