Abstract
The integrity of natural landscapes is affected by human actions, mainly by the intensification and expansion of agriculture. Factors such as fragment size and the structure of the environment can determine changes in the structure and composition of bird assemblages. In this study we evaluated the bird species composition in three structurally different forest environments, defined as the Core areas, Edge areas, and Buffer areas. The surveys were performed in the Natural Park of Sertão (NPS) and its surroundings in the southern limit of the Atlantic Forest, southern Brazil. To record species composition of birds, the point count method was used. The bird species were categorized according to feeding habits, habitat use, and sensitivity to forest fragmentation. A total of 131 species of birds distributed in 18 orders and 38 families were recorded. The species composition varied between the three areas and there was a significant difference in diversity between the Core and Edge areas and the Core and Buffer areas. Omnivorous species were found more in the Buffer areas than in the Core areas. Species that use the Broad habitat were more frequent in the Buffer and Edge areas than in the Core areas. Species that use the Forest habitat were found more in the Core areas than in the Buffer areas. Most recorded species (66%) have low sensitivity to forest fragmentation. The assemblage patterns found in this study, notably the differences among the three areas in diversity and species composition, can be explained by the ecological traits and the sensitivity levels of birds to forest fragmentation, which in turn seem to reflect different forest structures in the NPS and its surroundings. Although the Edge and Buffer areas have greater diversity, the Core areas showed great importance in maintaining species that are more sensitive to forest fragmentation. Even the largest fragments (such as the NPS), considering the regional context, may have bird species that are widely distributed and less sensitive to forest fragmentation. Forested habitat species may no longer occur or be restricted to the core area of the fragments. For conservation of bird fauna in the NPS, the maintenance of the core areas is essential, especially for those species that require a structurally preserved environment.
Keywords: Forest fragmentation, Edge effect, Bird conservation, Community ecology, Agricultural matrix
BACKGROUND
Forest degradation and fragmentation processes are widely distributed in the world and are especially associated with the expansion of human development frontiers (Haddad et al. 2015; Pendrill et al. 2022). The integrity of natural landscapes is affected by human actions, mainly by the intensification and expansion of agriculture (Solórzano et al. 2021). These changes in land use and cover can reduce the quantity and quality of natural habitats (Hill and Curran 2003). Therefore, the degradation of natural landscapes promotes changes in biodiversity, ecosystem stability, and resilience in the face of disturbances (Fahrig 2003).
Decreasing area and increasing fragmentation reduce species richness (Ibáñez et al. 2014; Haddad et al. 2015) and the diversity of functional characteristics of biological communities (Bregman et al. 2016; Luz et al. 2019). For example, forest fragmentation increases the edge effect and, consequently, the number of generalist species in forest fragments (Menke et al. 2012). In extreme situations, in small patches, local extinction of certain groups of species may occur, such as those restricted to conserved forest habitats (Devictor et al. 2008).
Forest fragmentation promotes changes in landscapes that may reflect on the structure and composition of bird assemblages (Anjos and Boçon 1999; Silva et al. 2017). Factors such as fragment size and structure of the environment can determine the presence or absence of species (Fischer and Lindenmayer 2007). Larger size and circular shape, with less edge effect, contribute to a larger central area of the fragment with a higher degree of conservation (Laurance et al. 2002).
The structure of the environment also influences the diversity of birds in a region (Anjos et al. 2019). The degree of tolerance of each species to changes in its environment varies according to its ability to adjust to habitat conditions (Silva et al. 2017; Mikolaiczik et al. 2019). Thus, different areas of a fragmented forest may have differences in species composition, depending on the degree of alteration generated by the effect of the matrix habitat. The maintenance of connectivity is associated with the physical arrangement of the landscape and the permeability of the matrix, that is, the ability of the species to overcome the matrix habitat and to move between the fragments (Metzger and Décamps 1997; Tischendorf and Fahrig 2000). In this way, the maintenance of populations of bird species in fragmented environments depends on the relationship between the quality and structure of the forest fragment with the size and isolation of the fragment (Boscolo and Metzger 2011).
In order to understand the mechanisms that influence the dynamics of species, in addition to the structural factors of the fragments and the matrix, it is also necessary to consider the traits of the species (Uezu and Metzger 2011). The greater the structural and microclimatic differences, the lower the probability of species sensitive to fragmentation being able to use matrix or fragment edges (Stouffer et al. 2006). The effects of forest fragmentation and environmental structure on bird assemblages also show regional variations, which probably reflects the complex relationship between habitat modification and the characteristics of each region (Restrepo et al. 1997; Silva et al. 2017; Anjos et al. 2019; Mikolaiczik et al. 2019).
The Atlantic Forest is one of the most biodiverse biomes in the world; however, it is also one of the most threatened (Ribeiro et al. 2009; Marques et al. 2021). The remaining area of this biome with natural forest cover corresponds to only 13.1% of that of the original forest (Fundação SOS Mata Atlântica, INPE 2019). The forest cover is mainly distributed in small and isolated fragments, composed of forests of various ages and in different stages of regeneration (Lira et al. 2021), incorporated into a matrix of degraded areas, pastures, agricultural and urban areas (Joly et al. 2014).
The Natural Park of Sertão (NPS) is a protected area located at the southern limit of the Atlantic Forest distribution (Slaviero 2014). During the 1960s, the region where the NPS is located, while originally covered by forest, became highly fragmented by nearby logging and agricultural expansion. The advance of agricultural cultivation in the region, with an emphasis on soybean, corn, and wheat crops, increased human pressure in the areas surrounding the Park (Slaviero et al. 2014).
Understanding how the forest structure influences the bird assemblages, mainly in highly fragmented regions inserted in an agricultural matrix, is essential for the conservation of birds, especially in the Atlantic Forest, with its high endemism and high anthropogenic pressure (Barbosa et al. 2017). In this study, we evaluated the avifauna assemblage patterns in the NPS and its surroundings in order to answer the following questions: How does the bird species composition vary between different forest structures in the NPS? Are the patterns found associated with the ecological traits of the species and their sensitivity to forest fragmentation?
MATERIALS AND METHODS
Study area
The study was carried out in the Natural Park of Sertão (NPS) and its surroundings, located in the municipality of Sertão, in the north of Rio Grande do Sul state, Brazil (Fig. 1). The area has slightly undulating relief, with an approximate altitude of 650 m (SERTÃO 2015). The climate has an average annual temperature of 17.5°C and an average annual rainfall of 1,800 mm, with well-distributed rainfall throughout the year. Originally covered by the Atlantic Forest biome, the landscape of the study area is characterized by the predominance of Mixed Ombrophilous Forest (Oliveira-Filho et al. 2015). The NPS consists of two forest fragments totaling 590.80 ha (Slaviero 2014). The study was performed in the largest fragment (513 ha) and in the forest remnants in its surroundings. The surroundings of the NPS are characterized by the predominance of rural properties with an agricultural economy (SERTÃO 2015), in which small forest fragments occur, which may or may not be connected to the NPS (Fig. 1).
Fig. 1.
Study area transects and sampling points in the Natural Park of Sertão (NPS), southern Brazil. Core areas 1, 2 and 3; Edge areas 1, 2, and 3; and Buffer areas 1, 2 and 3.
Bird surveys
The surveys were performed in structurally different forest environments, defined as the Core area, Edge area, and Buffer area of the NPS. The areas differ structurally, mainly in the height, volume, and average diameter of the trees, in addition to the richness of plant species and the stage of regeneration, which are influenced by the edge-core gradient (Slaviero et al. 2014).
The core areas of the NPS have higher vegetation, dense canopy, and older trees. In these areas, the advanced regeneration stage predominates, representing the most preserved sites of the NPS (Slaviero et al. 2014; SERTÃO 2015).
The edge areas in the NPS represented the transition between forest vegetation and the agricultural matrix (Slaviero et al. 2014). These areas are characterized by the intermediate stage of regeneration close to the agricultural matrix, and the advanced stage when closer to the core area of the fragment. In the NPS, the edge effect on forest vegetation is estimated at up to 250 meters, and the edge areas show greater plant species richness than the core area, mainly with species from the regenerating stratum (Slaviero et al. 2014).
The forest fragments in the buffer areas (within a radius of 1 km from the edge of the NPS) are small (< 10 ha) and are isolated from the largest fragment (SERTÃO 2015). These fragments are elongated, narrow (ranging from 50 to 160 meters in width), and usually associated with streams. Fragments are subject to the edge effect, which in most cases affects the entire fragment area. Vegetation tends to be lower and less diverse than in the other study areas. In these areas, the intermediate regeneration stage predominates, with sites of the early regeneration stage at the edges (SERTÃO 2015).
In each environment, with different forest structures, three sample transects were established, according to the following definitions: Core area transects (Core area 1, Core area 2, and Core area 3) inside the NPS, with at least 250 meters from the edge of the fragment; Edge transects (Edge area 1, Edge area 2, and Edge area 3), situated between 10 and 30 meters from the edge of the NPS (into the fragment); and Buffer transects (Buffer area 1, Buffer area 2, and Buffer area 3), in forest fragments around the NPS, distant 5001000 meters from the edge of the NPS. All transects, regardless of area, were at least 500 meters apart (Fig. 1).
For each transect, four sampling points were established, 100 meters apart, totaling 300 meters in length. A distance of up to 100 meters is recommended so that individuals are not counted more than once and to generate sample independence (Santos Junior et al. 2016; Mammides et al. 2016).
To record species richness and abundance of birds, the point count method was used (Blondel et al. 1970). Two observers remained at each point for 15 minutes and recorded birds sighted and/or heard. Birds within a radius of up to 30 meters from the counting point and only those active in the evaluated environment were recorded (e.g., birds in flight were not recorded). By this method, each counting point is considered as a sample unit, and was used to calculate the Punctual Abundance Index (PAI) (Blake 2007).
The surveys were carried out from November 2020 to February 2021, which corresponds to the breeding season of most bird species. Observations were performed between 5:45 am and 9:45 am. Two transects were sampled per day, alternating the start time between transects and sample areas. Each transect was sampled eight times, totaling 64 samples.
Birds recorded during displacements outside the sample transects were considered Occasional Encounter (OE) and were not used in comparative analyses between areas. Photographic records and vocalizations, when possible, were performed and used to identify or confirm the identification of species. Species were identified with the help of field guides and specialized literature (Sick 2001; Sigrist 2014; Meller 2017; Jacobs and Fenalti 2020). The nomenclature and taxonomic order adopted followed the proposal by the Brazilian Committee of Ornithological Records (Pacheco et al. 2021).
The bird species recorded were categorized according to their feeding habits, habitat use and sensitivity to forest fragmentation. The ecological traits of feeding habits and habitat use of the birds were determined according to descriptions in the literature (Willis 1979; Sick 2001). Feeding habit categories for birds in the NPS and its surroundings were: Carnivorous (CAR, captures and consumes other animals, mainly vertebrates), Detritivorous (DET, consumes carcasses of dead animals), Frugivorous (FRU, consumes mainly fruits), Granivorous (GRA, consumes mainly grains and seeds), Insectivorous (INS, specializes in the consumption of insects), Nectarivorous (NEC, consumes mainly nectar), and Omnivorous (OMN, wide and varied diet, able to consume different foods). The following habitat use categories were recorded for birds in the NPS and its surroundings: Broad (BRO, can occupy different habitats, including anthropic areas),
Open area (OA, mainly occupies open areas such as grasslands and savannas), Wetlands (WET, occurs mainly in flooded areas, such as ponds and rivers), Forest edge (FE, occupies transition areas between forested and open areas), and Forest (FOR, occupies the interior of forested areas). Birds were categorized with High, Medium or Low sensitivity to forest fragmentation as described primarily by Stotz et al. (1996) and Anjos (2006).
Data analysis
To compare the similarity between species composition, in different structures of forest environments, a Cluster Analysis (Cluster-UPGMA) was calculated using the Bray-Curtis Index. Also, we used the Jaccard similarity coefficient (Sij) to compare the similarity between areas. Data normality was assessed using the Shapiro-Wilk test and homogeneity using the Levene test. Comparisons between areas (Core area, Edge area, and Buffer area) for the number of species and for the ecological traits of food habits and habitat use (number of species recorded per transect and per sampling) were made using an analysis of variance (One-way ANOVA) and the Tukey post-hoc test. Comparisons of sensitivity to forest fragmentation (number of species recorded in each category per area) were performed using the Kruskal-Wallis test and Dunn post-hoc test. The diversity between the three areas was compared using the Shannon H' diversity index. To test whether H' values obtained in each area differ from each other, the t-test for specific diversity was used. Analyses were performed using the PAST 4.06 program (Hammer et al. 2001).
RESULTS
A total of 131 bird species distributed in 18 orders and 38 families were documented (Appendix 1). Of these, 106 were recorded in the transects and another 25 as occasional encounters. The total number of species observed represents 6.64% of the birds recorded in Brazil (Pacheco et al. 2021) and 18.42% of the birds recorded in the Rio Grande do Sul state (Jacobs and Fenalti 2020). The most represented families were Thraupidae (N = 18) and Tyrannidae (N = 16).
The largest number of bird species was recorded in the Buffer area (N = 74) followed by the Edge area (N = 73) and the Core area (N = 59). Although the number of species in the Core area was lower, there was no difference in the number of species recorded between the three areas (F = 0.012; p = 0.98; Fig. 2). The species diversity index, considering the three sampling areas, was H' = 3.93. Among the sampled areas, the greatest diversity was presented in the Edge area (H' = 3.89), followed by the Buffer area (H' = 3.85) and the Core area (H' = 3.65). There was a significant difference in the diversity index between the Core and Edge areas (t = -2.55; p = 0.01), and between the Core and Buffer areas (t = 2.21; p = 0.02). There was no difference between the Edge and Buffer areas (t = 0.40; p = 0.68).
Fig. 2.
Number of bird species recorded in the three sampling areas (Core area, Edge area, and Buffer area) in the Natural Park of Sertão (NPS), southern Brazil. Middle line (mean), boxes (mean ± standard deviation), and vertical bars (mean ± conf. interval).
Species composition varied among the three areas analyzed. Cluster analysis showed that the species composition of the Core areas differs from that of the other two areas (Fig. 3). Only 35 species (33.02%) of the 106 species found were recorded in the three sample areas. On the other hand, 41 species (38,68%) were exclusive to one area: nine species were found only in the Core area, 14 only in the Edge area, and 18 only in the Buffer area. The similarities were higher between the Edge and Buffer areas (S = 0.67; 50 species), followed by the Core and Edge areas (S = 0.65; 44 species), and the lowest similarity was observed between the Core and Buffer areas (S = 0.59; 41 species).
Fig. 3.

Dendrogram based on Bray-Curtis dissimilarity to species composition in the three sampling areas (Core area, Edge area, and Buffer area) in the Natural Park of Sertão (NPS), southern Brazil.
The average Punctual Abundance Index (PAI) was 0.18. All species with PAI > 0.18 (N = 22) occurred in the Edge and Buffer areas. Two of these species were not recorded in the Core area. When considering the PAI per sample area, the Buffer area showed more species with PAI > 0.18 (N = 16), followed by the Edge area (N = 14) and Core area (N = 12).
There was a difference in the average number of species with certain ecological traits between the areas. Omnivorous species were found more in the Buffer area than in the Core area. Broad habitat species were more frequent in the Buffer and Edge areas than in the Core area. Forest habitat species were found more in the Core area than in the Buffer area. There were no significant differences for the other analyzed ecological traits (Table 1; Appendix 2).
Table 1.
Comparisons of species richness per feeding habit and habitat use of bird species among the tree sampling areas (Core area, Edge area, and Buffer area) through Analysis of Variance with Tukey post-hoc test, Natural Park of Sertão (NPS), southern Brazil
Most recorded species (N = 67; 66%) had low sensitivity to forest fragmentation, followed by medium sensitive species (N = 32; 32%), and only two species showed high sensitivity (2%). No indication of the degree of sensitivity to forest fragmentation was found for six species (Appendix 1). Species with low sensitivity predominated in the three sampling areas (Fig. 4). The number of species in the study sites differed based on degree of sensitivity to forest fragmentation. Low sensitivity species were more commonly found in the Buffer area than in the Core area (H = 7.68, p = 0.02). Medium sensitivity species were more commonly found in the Core and Edge areas than in the Buffer areas (H = 7.26, p = 0.02). The number of highly sensitive species, due to their low number, did not vary between areas (H = 3.20, p = 0.09; Table 2; Appendix 2). No threatened species were recorded in NPS during this study.
Fig. 4.
Number of bird species per sensitivity level to forest fragmentation in the three sampling areas (Core areas, Edge areas, and Buffer areas) in the Natural Park of Sertão (NPS), southern Brazil.
Table 2.
Comparisons of bird species sensitivity level to forest fragmentation among the tree sampling areas (Core areas, Edge areas, and Buffer areas) through Kruskal-Wallis with Dunn post-hoc test, Natural Park of Sertão (NPS), southern Brazil
DISCUSSION
The number of species recorded (N = 131) seems to reflect the richness of birds for the NPS and its surroundings. In previous studies carried out at the NPS, which served as the basis for the management plan of the Park (Rezende and Agne 2014), a few more species were recorded than in our study (N = 154), through sampling fieldwork and the addition of bibliographic data.
The number of species recorded in the NPS and its surroundings is similar to that observed in other protected areas also located in fragmented landscapes with intense agricultural use in the south of the Atlantic Forest. In the Passo Fundo National Forest (1358 ha; 30 km from the NPS), characterized by the Mixed Ombrophilous Forest in different stages of regeneration, 129 bird species were recorded (PLANO DE MANEJO DA FLORESTA NACIONAL DE PASSO FUNDO 2011). In the Mata do Rio Uruguai Teixeira Soares Municipal Natural Park (431 ha; 65 km from the NPS), Mikolaiczik et al. (2019) recorded 145 bird species, evaluating the influence of forest regeneration stages on the richness and species composition in the transition between Seasonal Deciduous Forest and Mixed Ombrophilous Forest. In Fritz Plaumann State Park (717.48 ha; 82 km from the NPS), also under the influence of the Seasonal Deciduous Forest, 221 bird species were recorded (field and bibliographic survey) in the park and buffer zone (PLANO DE MANEJO FASE II DO PARQUE ESTADUAL FRITZ PLAUMANN 2014).
In forest fragments outside the protected areas, but also in fragmented landscapes at the south of the Atlantic Forest, the number of species remains similar. In a forest area in the municipality of Augusto Pestana (236.4 ha; 180 km from the NPS), 126 species of birds were recorded among the Core, Edge, and Buffer areas (Jacoboski et al. 2014a). In a forest fragment of about 200 ha (185 km from the NPS), 87 bird species were recorded, considering the Core and Edge areas of the forest fragment (Jacoboski et al. 2014b). A richness of 165 bird species was recorded by Teixeira and Pardini (2009) in the municipality of Frederico Westphalen (area of about 100 ha; 120 km from the NPS) in a landscape formed by a mosaic of small forest fragments and characterized by agriculture.
Our results showed that the Core areas differs from the Edge and Buffer areas due to the difference in the diversity index, as well as in the species composition. On the other hand, the species composition was similar between the Edge and Buffer areas. This fact may be associated with the structural similarities of the Edge and Buffer areas (see description in Methods and in Slaviero et al. 2014). The low percentage of species that occurred in the three environments (33%) and the percentage of species that occurred in just one area (38%) confirm the differences in diversity and species composition.
Bird responses to human interventions range from those that have benefited from habitat changes and increased their populations, to those that are excluded locally from environments (Marini and Garcia 2005). In the Edge and Buffer areas of the NPS, more species were found with the highest Punctual Abundance Index (PAI). According to Forman and Godron (1986), the intermediate disturbance of an area tends to promote heterogeneous landscape, and the absence of disturbance tends to a homogenization of the landscape. The Edge and Buffer areas seem to have more heterogeneity of environments/substrates (Slaviero et al. 2014), which allows for a greater occurrence of species, with a greater abundance of birds with flexibility in behavior and broad environmental tolerance.
The results regarding the ecological traits of the species reinforce the influence of the different structures of the environments on the distribution of birds in the NPS and its surroundings. Omnivory is a common trophic category in Neotropical birds, mainly in birds from open or forested areas under anthropic influence (Willis 1979; Sick 2001). A large number of omnivorous species is a characteristic of smaller forest fragments, as well as areas of secondary vegetation, due to the fact that omnivorous species adjust more easily to these types of environments (Mikolaiczik et al. 2019).
The high number of omnivorous species recorded in the Edge and Buffer areas of the NPS may be related to the forest structure of these environments. The alterations in the forest structure generated by the edge effect may provide an environment with broad characteristics of resources and habitats (Colles et al. 2009). The ability to use different food resources may favor omnivorous species in areas of heterogeneous forest structure (Anjos et al. 2019), such as the species recorded in the Edge and Buffer areas in this study.
The high number of Forest habitat species in the Core areas, and of Broad habitat species in the Edge and Buffer areas, corroborate the proposal of the influence of the different structures of the vegetation cover on the bird fauna in the NPS and its surroundings. The Core areas have a more homogeneous vegetation structure, with a denser and higher canopy (Slaviero et al. 2014) and can be considered a refuge for species primarily from forest environments.
Different bird species have different levels of sensitivity to environmental structure changes. Clearly, there are species that show different sensitivities to forest fragmentation (Parker III 1996; Uezu et al. 2005; Martensen et al. 2008). However, it is important to consider that the sensitivity levels of bird species vary according to the differences in landscape fragmentation (Anjos 2006). Our results show a small number of species that are highly sensitive to forest fragmentation in the study area. The tendency of the most sensitive birds is to disappear over time as changes in the landscape result in further degradation of the environment.
The variety of structurally different environments can result in a diversity of environmental resources and favor species that are less sensitive to forest fragmentation. On the other hand, greater homogeneity in the landscape may favor species sensitive to disturbances, which need large areas of internal habitat, far from the area of edge effect (Crooks et al. 2001; Bolger et al. 2001). The difference in the number of species of low sensitivity between the Core and Buffer areas, as well as the difference recorded in medium sensitive species between the Core and Buffer areas and between the Edge and Buffer areas, can be a consequence of the landscape structure on the occurrence of species in each sample area analyzed in the NPS.
Although the Edge and Buffer areas have greater diversity, the Core areas showed great importance in maintaining species that are more sensitive to forest fragmentation. If the most conserved habitat areas are reduced, birds that are more demanding in relation to the quality of the environment will disappear over time (Leck 1979; Whitmore 1997). On the other hand, if this fragmentation occurs, numbers of less sensitive species may increase, such as we observed among omnivorous species and those that use broad habitat types in the Edge and Buffer areas. Even the largest fragments, considering the regional context, such as the NPS, may have bird fauna mainly composed of widely distributed species that are less sensitive to forest fragmentation. Species that prefer forest habitats may no longer occur or be restricted to the core areas of the fragments with reduced populations. This may be indicated by the small number of species with high sensitivity. These species may have already been excluded due to the absence or low availability of preserved environments.
CONCLUSIONS
In conclusion, the present work shows that the diversity and species composition in the three sampled areas seem to be explained by the ecological traits and the sensitivity of bird species to forest fragmentation, which in turn seems to reflect different forest structures in the NPS and its surroundings. For conservation of bird fauna in the NPS, the maintenance of the core areas is essential, especially for species that require a structurally preserved environment.
Supplementary materials
Bird species recorded (X) in the tree sampling areas (Core, Edge and Buffer areas) in the Natural Park of Sertão (NPS), southern Brazil. Occasional encounter (OE). Feeding habits (FH): Carnivorous (CAR), Detritivorous (DET), Frugivorous (FRU), Granivorous (GRA), Insectivorous (INS), Nectarivorous (NEC), Omnivorous (OMN). Habitat use (HU): Broad (BRO), Forest (FOR), Forest edge (FE), Open area (OA), Wetland (WET). Sensitivity level to forest fragmentation (Sen): Low (L), Medium (M), High (H), -The authors agree with the suggestion. Punctual Abundance Index (PAI). Species without PAI value were recorded by Occasional encounter.
Number of species (N), mean and standard deviation (mean ± SD) by ecological traits and the level of sensitivity in the tree sampling areas (Core areas, Edge areas, and Buffer areas) in the Natural Park of Sertão (NPS), southern Brazil.
Acknowledgments
We are grateful to the Federal University of Fronteira Sul (UFFS) for providing logistical support. Valuable help in fieldwork was provided by Renato Betiol and Natural Park of Sertão team. We thank Dr. Caroline Muller comments and suggestions on previous versions of the manuscript. This study was financed in part by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior–Brasil (CAPES)–Finance Code 001.
Footnotes
Authors’ contributions: All authors contributed to the study conception, design, and data analysis. Data collection was performed by CB and PH. The first draft of the manuscript was written by CB, and all authors commented on previous versions of the manuscript. All authors read and approved the final manuscript.
Competing interests: CB, MH and PH declare that they have no conflicts of interest.
Availability of data and materials: The data supporting the findings of this study are available within the article and its supplementary materials.
Consent for publication: Not applicable.
Ethics approval consent to participate: Not applicable.
References
- Anjos L. 2006. Bird species sensitivity in a fragmented landscape of the Atlantic Forest in Southern Brazil 1. Biotropica 38:229–234. doi:10.1111/j.1744-7429.2006.00122.x.
- Anjos L, Boçon R. 1999. Bird communities in natural forest patches in southern Brazil. Wilson Bull 111:397–414.
- Anjos L, Bochio GM, Medeiros HR et al. 2019. Insights on the functional composition of specialist and generalist birds throughout continuous and fragmented forests. Ecol Evol 9:6318–6328. doi:10.1002/ece3.5204. [DOI] [PMC free article] [PubMed]
- Barbosa KVC, Knogge C, Develey PF, Jenkins CN, Uezu A. 2017. Use of small Atlantic Forest fragments by birds in Southeast Brazil. Perspect Ecol Conserv 15:42–46. doi:10.1016/j.pecon.2016.11.001.
- Blake JG. 2007. Neotropical forest bird communities: a comparison of species richness and composition at local and regional scales. Condor 109:237–255. doi:10.1650/0010-5422(2007)109[237:NF BCAC]2.0.CO;2.
- Blondel J, Chessel-Ferry C, Frochot B. 1970. La méthode des indices ponctuels d ´abondance (I.P.A.) ou des relevés d ´avifaune par “stations d ´écoute.” Alauda 38:55–71.
- Bolger DT, Scott TA, Rotenberry JT. 2001. Use of corridor-like landscape structures by bird and small mammal species. Biol Conserv 102:213–224. doi:10.1016/S0006-3207(01)00028-3.
- Boscolo D, Metzger JP. 2011. Isolation determines patterns of species presence in highly fragmented landscapes. Ecography 34:1018–1029. doi:10.1111/j.1600-0587.2011.06763.x.
- Bregman TP, Lees AC, MacGregor HEA, Darski B, de Moura NG, Aleixo A, Barlow J, Tobias JA 2016. Using avian functional traits to assess the impact of land-cover change on ecosystem processes linked to resilience in tropical forests. P R Soc B-Biol Sci 283:20161289. doi:10.1098/rspb.2016.1289. [DOI] [PMC free article] [PubMed]
- Colles A, Liow LH, Prinzing A. 2009. Are specialists at risk under environmental change? Neoecological, paleoecological and phylogenetic approaches. Ecol Lett 12:849–863. doi:10.1111/j.1461-0248.2009.01336.x. [DOI] [PMC free article] [PubMed]
- Crooks KR, Suarez AV, Bolger DT, Soulé ME. 2001. Extinction and Colonization of Birds on Habitat Islands. Conservation Biology 15:159–172. doi:10.1111/j.1523-1739.2001.99379.x.
- Devictor V, Julliard R, Jiguet F. 2008. Distribution of specialist and generalist species along spatial gradients of habitat disturbance and fragmentation. Oikos 117:507–514. doi:10.1111/j.00301299.2008.16215.x.
- Fahrig L. 2003. Effects of habitat fragmentation on biodiversity. Annu Rev Ecol Evol Syst 34:487–515. doi:10.1146/annurev.ecolsys.34. 011802.132419.
- Fischer J, Lindenmayer DB. 2007. Landscape modification and habitat fragmentation: a synthesis. Global Ecol Biogeogr 16:265–280. doi:10.1111/j.1466-8238.2007.00287.x.
- Forman RTT, Godron M. 1986. Landscape Ecology. John Wiley, New York, USA.
- Fundação SOS Mata Atlântica, INPE. 2019. Atlas dos Remanescentes Florestais da Mata Atlântica: Período 2017-2018. São Paulo.
- Haddad NM, Brudvig LA, Clobert J et al. 2015. Habitat fragmentation and its lasting impact on Earth’s ecosystems. Sci Adv. doi:10.1126/sciadv.1500052. [DOI] [PMC free article] [PubMed]
- Hammer Ø, Harper DAT, Ryan PD. 2001. PAST: paleontological statistics software package for education and data analysis. Paleontologia Electronica 4:1–9.
- Hill JL, Curran PJ. 2003. Area, shape and isolation of tropical forest fragments: effects on tree species diversity and implications for conservation. J Biogeogr 30:1391–1403. doi:10.1046/j.13652699.2003.00930.x.
- Ibáñez I, Katz DSW, Peltier D, Wolf SM, Connor Barrie BT 2014. Assessing the integrated effects of landscape fragmentation on plants and plant communities: the challenge of multiprocessmultiresponse dynamics. J Ecol 102:882–895. doi:10.1111/13652745.12223.
- Jacoboski L, Santos E, Ramos N. 2014a. Estrutura trófica da avifauna do Mato do Silva, fragmento de floresta estacional decidual, Chiapetta, Rio Grande do Sul. Revista da Biologia 12:22–28. doi:10.7594/revbio.12.02.04.
- Jacoboski LI, Oliveira TA, Bianchi V, Hartz SM. 2014b. Comparação da riqueza e composição de aves no interior e na borda em um fragmento de Floresta Estacional Decidual. Revista Biociências 2:40–51.
- Jacobs F, Fenalti P. 2020. Guia de identificação: Aves do Rio Grande do Sul. Aratinga, Pelotas.
- Joly CA, Metzger JP, Tabarelli M. 2014. Experiences from the Brazilian Atlantic Forest: ecological findings and conservation initiatives. New Phytologist 204:459–473. doi:10.1111/nph.12989. [DOI] [PubMed]
- Laurance WF, Lovejoy TE, Vasconcelos HL et al. 2002. Ecosystem decay of Amazonian forest fragments: a 22-year investigation. Conserv Biol 16:605–618. doi:10.1046/j.1523-1739.2002.01025. x.
- Leck CF. 1979. Avian extinctions in an isolated tropical wet-forest preserve, Ecuador. Auk 96:343–352. doi:10.1093/auk/96.2.343.
- Lira PK, Portela R de CQ, Tambosi LR. 2021. Land-cover changes and an uncertain Future: Will the Brazilian Atlantic forest lose the chance to become a hopespot? In: The Atlantic Forest. Springer International Publishing, Cham, pp. 233–251.
- Luz CCS, Ramos AWP, Silva GJO. 2019. Natural and environmental vulnerability of the Jauru-Mato Grosso river hydrographic basin, Brazil. Raega -O Espaço Geográfico em Análise 46:176–187.
- Mammides C, Kounnamas C, Goodale E, Kadis C. 2016. Do unpaved, low-traffic roads affect bird communities? Acta Oecologica 71:14–21. doi:10.1016/j.actao.2016.01.004.
- Marini MA, Garcia FI. 2005. Conservação de aves no Brasil. Megadiversidade 1:95–102.
- Marques MCM, Trindade W, Bohn A, Grelle CEV. 2021. The Atlantic Forest: An Introduction to the Megadiverse Forest of South America. In: The Atlantic Forest. Springer International Publishing, Cham, pp. 3–23.
- Martensen AC, Pimentel RG, Metzger JP. 2008. Relative effects of fragment size and connectivity on bird community in the Atlantic Rain Forest: Implications for conservation. Biol Conserv 141:2184–2192. doi:10.1016/j.biocon.2008.06.008.
- Meller DA. 2017. Guia de identificação: Aves da região noroeste do Rio Grande do Sul. Tenondé, Santo Ângelo.
- Menke S, Böhning-Gaese K, Schleuning M. 2012. Plant-frugivore networks are less specialized and more robust at forest-farmland edges than in the interior of a tropical forest. Oikos 121:1553–1566. doi:10.1111/j.1600-0706.2011.20210.x.
- Metzger J-P, Décamps H. 1997. The structural connectivity threshold: An hypothesis in conservation biology at the landscape scale. Acta Oecologica 18:1–12. doi:10.1016/S1146-609X(97)80075-6.
- Mikolaiczik NM, Barreto MS, Hartmann MT, Hartmann PA. 2019. Bird fauna in secondary forest stages: a study in a southern Brazilian protected area. Oecologia Australis 23:261–279. doi:10.4257/oeco.2019.2302.06.
- Oliveira-Filho AT, Budke JC, Jarenkow JA et al. 2015. Delving into the variations in tree species composition and richness across South American subtropical Atlantic and Pampean forests. Journal of Plant Ecology 8:242–260. doi:10.1093/jpe/rtt058.
- Pacheco JF, Silveira LF, Aleixo A et al. 2021. Annotated checklist of the birds of Brazil by the Brazilian Ornithological Records Committee—second edition. Ornithology Research 29:94–105. doi:10.1007/s43388-021-00058-x.
- Parker III TA. 1996. Birds and vegetation distribution and threat. In: D.F. S, Parker III TA, Fitzpatrick JN, Moskovits DK (eds) Neotropical Birds: Ecology and Conservation. University of Chicago Press, p. 502.
- Pendrill F, Gardner TA, Meyfroidt P et al. 2022. Disentangling the numbers behind agriculture-driven tropical deforestation. Science 377:eabm9267. doi:10.1126/science.abm9267. [DOI] [PubMed]
- PLANO DE MANEJO DA FLORESTA NACIONAL DE PASSO FUNDO. 2011. Florianópolis, Santa Catarina. Instituto Chico Mendes de Conservação da Biodiversidade. Socioambiental Consultores Associados Ltda, Florianópolis.
- PLANO DE MANEJO FASE II DO PARQUE ESTADUAL FRITZ PLAUMANN. 2014. Santa Catarina. Fundação do Meio Ambiente (FATMA). Volume I: Plano Básico. FATMA Caipora Cooperativa para Conservação da Natureza, Florianópolis.
- Restrepo C, Renjifo LM, Marples P. 1997. Frugivorous birds in fragmented neotropical montane forests: landscape pattern and body mass distribution. In: Tropical forest remnants: ecology, management and conservation of fragmented communities, pp. 171–189.
- Rezende EL, Agne CE. 2014. Avifauna do Parque Natural Municipal de Sertão. In: Tedesco CD, Zanella N (eds) Parque Natural Municipal de Sertão. Universidade de Passo Fundo, Passo Fundo, pp. 90–100.
- Ribeiro MC, Metzger JP, Martensen AC et al. 2009. The Brazilian Atlantic Forest: How much is left, and how is the remaining forest distributed? Implications for conservation. Biol Conserv 142:1141–1153. doi:10.1016/j.biocon.2009.02.021.
- Santos Junior PCA, Marques FC, Lima MR, Anjos L. 2016. The importance of restoration areas to conserve bird species in a highly fragmented Atlantic forest landscape. Nat Conservação 14:1–7. doi:10.1016/j.ncon.2016.03.001.
- SERTÃO. 2015. Plano de Manejo do Parque Natural Municipal de Sertão –RS. In: Prefeitura Municipal de Sertão (Secretaria Municipal de Agricultura e Desenvolvimento Econômico). Available at: https://www.sertao.rs.gov.br/pagina/561/parquenatural-municipal-de-sertao. Accessed 6 Sept. 2021.
- Sick H. 2001. Ornitologia Brasileira, 1st edn. Nova Fronteira, Rio de Janeiro.
- Sigrist T. 2014. Guia de Campo: Avifauna Brasileira. São Paulo.
- Silva VP, Deffaci AC, Hartmann MT, Hartmann PA. 2017. Birds around the road: effects of a road on a Savannah bird Community in Southern Brazil. Ornitol Neotrop 28:119–128.
- Slaviero LB. 2014. Criação do Parque Natural Municipal de Sertão. In: Tedesco CD, Zanella N (eds) Parque Natural Municipal de Sertão. Universidade de Passo Fundo, Passo Fundo, pp. 13–26.
- Slaviero LB, Budke JC, Cansian RL. 2014. As florestas do Parque Natural Municipal de Sertão. In: Tedesco CD, Zanella N (eds) Parque Natural Municipal de Sertão. Universidade de Passo Fundo, Passo Fundo, pp. 41–68.
- Solórzano A, Brasil LSC de A, de Oliveira RR. 2021. The Atlantic Forest Ecological History: From Pre-colonial Times to the Anthropocene. In: Marques MCM, Grelle CEV (eds) The Atlantic Forest. Springer International Publishing, Cham, pp. 25–44.
- Stotz DF, Fitzpatick JF, Parker TA, Moskovits DK. 1996. Neotropical Birds: Ecology and Conservation. University of Chicago Press, Chicago, USA.
- Stouffer RJ, Broccoli AJ, Delworth TL et al. 2006. GFDL’s CM2 Global Coupled Climate Models. Part IV: Idealized Climate Response. J Clim 19:723–740. doi:10.1175/JCLI3632.1.
- Teixeira AMG, Pardini R. 2009. Time-lag in biological responses to landscape changes in a highly dynamic Atlantic Forest region. Biol Conserv 142:1166–1177. doi:10.1016/j.biocon.2009.01.033.
- Tischendorf L, Fahrig L. 2000. On the usage and measurement of landscape connectivity. Oikos 90:7–19. doi:10.1034/j.1600-0706. 2000.900102.x.
- Uezu A, Metzger JP. 2011. Vanishing bird species in the Atlantic Forest: relative importance of landscape configuration, forest structure and species characteristics. Biodivers Conserv 20:3627–3643. doi:10.1007/s10531-011-0154-5.
- Uezu A, Metzger JP, Vielliard JME. 2005. Effects of structural and functional connectivity and patch size on the abundance of seven Atlantic Forest bird species. Biol Conserv 123:507–519. doi:10.1016/j.biocon.2005.01.001.
- Whitmore TC. 1997. Tropical forest disturbance, disappearance, and species loss. In: Laurance WF, Bierregaard Jr RO (eds) Tropical Forest Remnants: Ecology, Management, and Conservation of Fragmented Communities. University of Chicago Press, Illinois, pp. 3–12.
- Willis EO. 1979. The composition of avian communities in remanescent woodlots in southern Brazil. Pap Avulsos Zool 33:1–25.
Associated Data
This section collects any data citations, data availability statements, or supplementary materials included in this article.
Supplementary Materials
Bird species recorded (X) in the tree sampling areas (Core, Edge and Buffer areas) in the Natural Park of Sertão (NPS), southern Brazil. Occasional encounter (OE). Feeding habits (FH): Carnivorous (CAR), Detritivorous (DET), Frugivorous (FRU), Granivorous (GRA), Insectivorous (INS), Nectarivorous (NEC), Omnivorous (OMN). Habitat use (HU): Broad (BRO), Forest (FOR), Forest edge (FE), Open area (OA), Wetland (WET). Sensitivity level to forest fragmentation (Sen): Low (L), Medium (M), High (H), -The authors agree with the suggestion. Punctual Abundance Index (PAI). Species without PAI value were recorded by Occasional encounter.
Number of species (N), mean and standard deviation (mean ± SD) by ecological traits and the level of sensitivity in the tree sampling areas (Core areas, Edge areas, and Buffer areas) in the Natural Park of Sertão (NPS), southern Brazil.





