Population, community, and ecosystem studies often focus on very specific parts of social-ecological systems, such as a selected species, a single habitat type, a unique ecological process, one or few environmental governance regimes or certain anthropogenic threat. Mostly, simplification is the only way to understand biodiversity and ecosystems given their inherent complexity. Nevertheless, expanding the range of analysis by including multiple systems’ elements, as well as the interaction among them, without losing precision, is wishful. Yet, integrative analyses of conservation that addresses both social and ecological complexity are rare.
One overlooked aspect of simplistic ecological models is landscape heterogeneity, which is defined as the qualitative and quantitative variation in landscape elements (Li and Reynolds 1994; 1995; Fahrig et al. 2011), including the variety and the spatial arrangement of land cover types compositional and configurational heterogeneities, respectively (Li and Reynolds 1995). The idea of islands composed of a single habitat type embedded in a non-habitat environment echoes in much of the ecological literature (Perfecto and Vandermeer 2008). However, landscape heterogeneity increases when more than one habitat type is considered and when habitat patches present a variety of shapes and sizes. Both compositional and configurational heterogeneities influence biotic and abiotic processes important for maintaining functional landscapes, including predation (Kauffman et al. 2007), pest control (Gardiner et al. 2009), pollination (Boscolo et al. 2017), and the movement of individuals (Romero et al. 2009).
Habitat area is traditionally known as the most important factor for conserving biodiversity and when addressing the conservation of heterogeneous landscapes, conservationists have often used the concept of representativeness. Habitat representativeness is defined as how different habitats are under or overrepresented by reserves relative to their area, in another words, the proportion of habitat under protection (Braz and Cavalcanti 2013; Austin and Margules 1986). Nevertheless, ecologists, biologists and environmental scientists are becoming aware that apart from protecting large habitat tracks, certain biological processes must be preserved in order to maintain landscape functionality and stability for species, including humans. Among those ecosystem processes, landscape connectivity has a key role to the long term persistence of populations in patchy landscapes (Taylor et al. 1993). Sub-populations (a population’s subset of individuals) connected via dispersal in highly connected landscapes are able to guarantee recolonization after possible local extinction events, a process known as “rescue effect” (Brown and Kodric-Brown 1977). Thus, conservationists should target the protection of landscape features that are important for landscape connectivity, particularly for functional connectivity, defined as how much a given landscape fosters of renders biological flux given its structure and the species responses to it (Metzger 2001).
Functional connectivity is fundamental for conservation and can be addressed in different ways, from regional to local levels, including the entire landscape or by connecting specific places, such as reserves or large habitat patches. Yet, assessing functional connectivity on heterogeneous landscape is challenging. Different land cover classes present diverse influences on species movements across landscapes and those classes most similar to species habitats can assist in the maintenance of the landscape functional connectivity (e.g., agroforests allow the movement of bird species between native forest patches) (Goulart et al. 2015). Among the methods for modeling and understanding functional connectivity, the integral index of connectivity (Saura and Pascual-Hortal 2007; Horta et al. 2018) provides good indicators of how much a given landscape or a habitat patch is important for functional connectivity. Another approach is least-cost corridors, which stands out for identifying routes that may act as ecological corridors between habitat patches (Goulart et al. 2015; Bhakti et al. 2021; Graviola et al. 2021). As to habitat area, the concept of representativeness could be adapted to the understanding of how different governance regimes protect connectivity for biodiversity. This issue could be particularly useful in the understanding and management of heterogeneous and complex landscapes.
Heterogeneous landscapes are also often subjected to diverse governance regimes, from strictly protected, sustainable-use, and the matrix, which increases complexity for conservation scholars and practitioners. We here define matrix as the area that is not under legal protection, which is generally composed of a set of anthropogenic managements, natural and semi-natural areas, although we are aware of the other possible definitions of this term (Forman 1995; Kupfer et al. 2006; Perfecto et al. 2009). These diverse governance regimes have different impacts on nature conservation or degradation, and while many studies have devoted to measuring how much each regime effectively protect habitats, some researchers argue that multiple strategies acting in an integrated form are best (Blackman 2015; Lima et al. 2020). In this sense, the mosaic composed of different governance regimes is a promising option. In Brazil, the Conservation Mosaics is a national policy that combines management of multiple strict conservation reserves (from local to national parks) together with sustainable-use reserves [called Environmental Protection Areas (EPAs)] in association with matrixes’ stakeholders. This is an effort to integrate governance networks, at the same time that optimizes and maximizes conservation strategies. Due to the fact that most ecological processes are not restricted to individual conservation units, but act on the entire landscape, the strategy of Conservation Mosaics may be better fitted into ecological scales, and thus more successful in conserving biodiversity and ecosystems at long term (Bergsten et al. 2014).
Apart from those issues, multiple drivers of species loss are rarely considered together in a single framework. Instead, studies focus on isolated threats, such as deforestation (Nolte et al. 2013), fire (Nepstad et al. 2006), exotic eucalypt afforestation, agriculture intensification (Fernandes et al. 2016; Goulart et al. 2016; Goulart et al. 2016) or mining on habitats (Sonter et al. 2017) or species (Pena et al. 2017). All these impacts reduces habitat area and function connectivity for wildlife. Yet, to have a complete picture of the biodiversity threats, there is a need to understand how these impact act synergistically.
The Espinhaço mountain range, located in the Southeast of Brazil, is notable for its biological importance, natural heterogeneity, social complexity, as well as the diversity of threats. The western part of the mountain range is characterized by the Cerrado biome, a savanna-like environment, while the eastern slope is dominated by a semi-deciduous tropical forest, the Atlantic Forest biome (see reviews in Fernandes 2016b). Both of these biomes are considered global hotspots for conservation (Mittermeier et al. 1998). The upper parts of the mountain range are covered by the rupestrian grassland, a mixture of rocky outcrops, bushlands and grasslands, considered a Biosphere Reserve by UNESCO due to its species and ecosystems richness (Miola et al. 2021; Fernandes et al. 2020; 2018). All the three environments forest, cerrado (both are written in minor case letters to distinguish the ecosystem type from the biome, which is written in upper case), and rupestrian grassland, are inhabited by their specific biological community, among which, birds distinguish due to their large diversity and degree of endemism (Vasconcelos 2008; Hoffmann et al. 2020). Parallel to this fact, the governance regimes are also very diverse, and the region is composed of a network of strictly protected and sustainable-use reserves, surrounded by a matrix formed by a variety of land covers and management types (from managed pastures to natural habitat patches).
Within the Espinhaço mountain range lies the Mosaic of Conservation Units of the Espinhaço: Alto Jequitinhonha - Serra do Cabral, hereafter Espinhaço Mosaic (EM) (www.icmbio.gov.br/portal/images/stories/mosaicos/planejamento-espinhaco.pdf). Conservation Mosaic is a model that seeks for integrating management of reserve networks via a participatory basis, in order to conserve biodiversity, ecosystems and foster sustainable development adopted by the Brazilian government since 2010 (http://www.mma.gov.br/areas-protegidas/instrumentos-de-gestao/mosaicos). The EM was built under a participatory basis including the State Forest Institute (IEF), Ministry of Environment, Chico Mendes Institute for Biodiversity Conservation, together with the NGOs Biotropicos Institute and Conservation International (https://www.icmbio.gov.br/portal/images/stories/mosaicos/planejamento-espinhaco.pdf).
As the other parts of the Espinhaço mountain range, the EM is composed of a diversity of vegetation types (mostly forest, cerrado and rupestrian grasslands), all of which inhabited numerous species of endemic and endangered birds. The EM is subjected to multiple governance regimes (from strictly protected reserves, sustainable-use reserves, to the matrix), as well as is under a set of anthropogenic threats, such as afforestation, road development, mining activities, urban expansion, fires (Pena et al. 2017; Fernandes et al. 2018; Fernandes et al. 2016; Sonter et al. 2013). The physical, environmental, biological and social aspects of this region make it a good case study for integrating the analysis of biodiversity conservation in a highly complex landscape.
Given the above mentioned, we propose an analytical framework that contributes to the management of complex landscapes by understanding the interactive effects of anthropogenic threats and governance regimes on landscape traits that are determinant for biodiversity conservation. We then apply the framework to the management of bird conservation in the EM. To this end we selected the main landscape traits that are important for maintaining such diversity (habitat area, integral landscape connectivity, connectivity among strictly protected reserves), main governance regimes (matrix, strictly protected reserves, sustainable-use reserves) and main threats that are known to occur in the region (fire, mining and wood-cover loss).
More specifically, this study aims at assessing the following objectives (Fig. 1):
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Compare the area representativeness of the different governance regimes in terms of covering habitat area.
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Compare the representativeness of the different governance regimes in terms of protecting landscape integral connectivity .
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Compare the representativeness of the different governance regimes in terms of protecting connectivity among strictly protected reserves.
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Compare the occurrence of different threats on habitat area.
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Compare the occurrence of different threats on the landscape integral connectivity.
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Compare the occurrence of different threats on the connectivity among strictly protected reserves.