Research Article |
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Corresponding author: Cássio Cardoso Pereira ( cassiocardosopereira@gmail.com ) Academic editor: François Gillet
© 2025 Angélica Aparecida Ávila, Cássio Cardoso Pereira, Crislaine Paula de Oliveira, Carlos A. Ordóñez-Parra, Sérgio Gualberto Martins, Gislene Carvalho de Castro.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Ávila AA, Pereira CC, de Oliveira CP, Ordóñez-Parra CA, Martins SG, de Castro GC (2025) Forgotten guardians of biodiversity in the Atlantic Forest: seed rain confirms the ecological role of hedgerows in landscape connectivity. Plant Ecology and Evolution 158(3): 350-357. https://doi.org/10.5091/plecevo.160271
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Background and aims – Hedgerows may play a crucial ecological role in maintaining connectivity in fragmented Atlantic Forest landscapes. However, little is known about their contribution to ecological processes, particularly seed rain.
Material and methods – To test this, we conducted a study in Ritápolis, Minas Gerais, Brazil, comparing a 500-meter-long hedgerow corridor with a 10.5-hectare forest fragment. We installed 20 seed collectors in each area and monitored them for a year, testing the hypothesis that hedgerow corridors have equivalent levels of litter, fruit, and seed production compared to preserved forest fragments.
Key results – The results revealed that hedgerows can present greater fruit richness and higher litter dry weight than adjacent forest fragments. Furthermore, the other parameters evaluated in these hedgerows were equivalent to those observed in fragmented areas, reinforcing their importance in biodiversity conservation in rural landscapes. These findings directly support the hypothesis tested, highlighting hedgerows as essential structures for maintaining ecological processes, increasing environmental connectivity, and mitigating the impacts of forest fragmentation.
Conclusion – This study contributes to the understanding of the role of ecological connectivity in hedgerows, paving the way for further research that can expand this knowledge.
ecological connectivity, ecosystem resilience, environmental governance, fragmentation mitigation, habitat restoration, leaf litter, seed dispersal, sustainable land management, vegetation corridors
A notable feature of the south-eastern Brazilian landscape is the presence of hedgerows, which emerged from the spontaneous establishment of vegetation in ditches constructed by slaves during European colonization, particularly in the 18th and 19th centuries (
Since the construction of these hedgerows to the present day, the Atlantic Forest has undergone extensive land-use transformations, giving rise to major urban centres and some of Brazil’s most productive agricultural lands (
Beyond their role in maintaining landscape connectivity, hedgerows may actively contribute to ecosystem functionality by influencing key ecological processes (
To develop effective conservation strategies, it is necessary to understand the functional aspects of these ecosystems. Current research efforts should incorporate indicators that elucidate the processes that precede plant establishment. Seed rain, for example, can be a critical factor in landscape regeneration, influencing immediate and long-term ecosystem recovery through species recruitment and establishment (
Despite this potential, hedgerows remain conceptually and functionally distinct from conventional ecological corridors, which have received considerable attention in conservation science, often through planned restoration or the identification of existing forest fragments with formal ecological functions (
In this study, we compared litter, fruit, and seed production between a hedgerow and a preserved Atlantic Forest fragment in Minas Gerais, Brazil. Our goal was to determine whether hedgerows contribute equivalently to the maintenance of key ecological processes, particularly seed rain. We hypothesized that hedgerows exhibit similar levels of litter production, as well as fruit and seed abundance and richness, compared to forest fragments, emphasizing their potential ecological role in fragmented landscapes. To our knowledge, this is the first study to examine these ecosystem functioning variables across these two distinct habitat types. To test this, we installed 20 seed collectors at each site, monitoring litter, fruit, and seed deposition over the course of one year.
The present study was conducted in the municipality of Ritápolis, Minas Gerais, Brazil, in a 500 m long and 4 to 5 m wide hedgerow and a preserved forest fragment of approximately 10.5 hectares near the Floresta Nacional de Ritápolis (FLONA), which covers 89 ha (
Forty 1-m2 seed collectors were systematically installed in the study area, with 20 seed collectors for each vegetation type and maintaining a distance of 50 m from the edge of the fragment to avoid edge effects. We tried to distribute the 20 seed collectors along the 500 m length of the hedgerow corridor, keeping them spatially separated at a distance of about 20–25 m from each other. The seed collectors were installed 30 cm above the ground to avoid sampling herbaceous seeds and small shrubs.
The collector’s content was collected fortnightly over a year, from July 2012 to June 2013. Each sample was placed in plastic bags, identified by collector and sampling date. After the collection, the samples underwent screening to separate diaspores (fruits and seeds) from other plant material (branches and leaves) with the aid of a stereomicroscope. Each fraction was dried at 70°C for 48 h and then weighted in a precision balance to obtain its dry weight. Abundance, richness, and dry weight data from each collector were summed over the year to capture seasonal variation and the actual accumulation of species and biomass. For fruit and seed richness, each species was counted only once, ensuring a more accurate comparison between habitats (Table
Total values of abundance and richness of fruits and seeds, and dry mass of fruits, seeds, leaves, and branches found for each of the 40 collectors installed in hedgerows (H, n = 20) and forests (F, n = 20) from July 2012 to June 2013, in the municipality of Ritápolis, Minas Gerais, Brazil. DW = dry weight.
| Habitat | Seed collectors | Fruit abundance | Fruit richness | Seed abundance | Seed Richness | DW of seeds (mg) | DW of fruits (mg) | DW of leaves and branches (mg) | Fruit diameter/length ratio |
| F | 1 | 43 | 19 | 9 | 7 | 0.34 | 1.33 | 1396.91 | 0.77 |
| F | 2 | 162 | 25 | 29 | 8 | 1.23 | 8.01 | 1281.49 | 0.48 |
| F | 3 | 259 | 31 | 19 | 7 | 1.10 | 6.65 | 870.63 | 0.57 |
| F | 4 | 239 | 36 | 26 | 17 | 3.65 | 15.62 | 761.87 | 0.51 |
| F | 5 | 312 | 20 | 46 | 7 | 0.31 | 5.06 | 1103.77 | 0.63 |
| F | 6 | 78 | 26 | 6 | 5 | 0.49 | 0.82 | 655.78 | 1.08 |
| F | 7 | 810 | 52 | 141 | 16 | 3.28 | 42.70 | 826.52 | 0.85 |
| F | 8 | 320 | 33 | 405 | 15 | 1.34 | 13.26 | 1045.90 | 0.66 |
| F | 9 | 1480 | 41 | 1600 | 19 | 1.13 | 13.56 | 2433.80 | 0.74 |
| F | 10 | 580 | 45 | 180 | 14 | 0.42 | 5.17 | 752.25 | 0.69 |
| F | 11 | 169 | 31 | 25 | 12 | 1.37 | 1.85 | 717.65 | 0.77 |
| F | 12 | 401 | 41 | 47 | 17 | 1.33 | 15.21 | 993.78 | 0.65 |
| F | 13 | 111 | 28 | 53 | 19 | 2.28 | 1.21 | 824.94 | 0.47 |
| F | 14 | 50 | 23 | 21 | 12 | 1.05 | 5.20 | 886.67 | 0.48 |
| F | 15 | 97 | 32 | 23 | 10 | 0.53 | 0.53 | 1156.43 | 0.54 |
| F | 16 | 216 | 39 | 14 | 9 | 0.62 | 1.63 | 1467.99 | 0.60 |
| F | 17 | 178 | 32 | 62 | 16 | 1.90 | 1.14 | 849.37 | 0.59 |
| F | 18 | 405 | 61 | 17 | 10 | 0.68 | 8.22 | 478.52 | 0.31 |
| F | 19 | 336 | 57 | 27 | 11 | 6.54 | 6.45 | 972.63 | 0.74 |
| F | 20 | 227 | 37 | 23 | 12 | 0.55 | 8.56 | 809.07 | 0.62 |
| H | 1 | 116 | 34 | 59 | 24 | 1.77 | 27.82 | 1370.64 | 0.27 |
| H | 2 | 410 | 59 | 85 | 23 | 1.58 | 85.63 | 3277.71 | 0.95 |
| H | 3 | 440 | 71 | 57 | 28 | 3.02 | 109.53 | 1801.97 | 0.95 |
| H | 4 | 354 | 49 | 32 | 12 | 1.58 | 18.65 | 1911.84 | 0.72 |
| H | 5 | 238 | 26 | 9 | 8 | 0.22 | 11.02 | 1108.40 | 0.25 |
| H | 6 | 549 | 64 | 11 | 10 | 0.20 | 51.89 | 1635.67 | 0.51 |
| H | 7 | 1205 | 62 | 23 | 15 | 0.92 | 34.01 | 1885.60 | 1.22 |
| H | 8 | 967 | 59 | 125 | 13 | 1.07 | 28.30 | 1237.35 | 0.94 |
| H | 9 | 376 | 42 | 12 | 7 | 1.12 | 21.86 | 4347.21 | 0.71 |
| H | 10 | 257 | 35 | 10 | 6 | 0.39 | 12.12 | 1055.27 | 0.75 |
| H | 11 | 205 | 56 | 56 | 11 | 1.65 | 15.09 | 785.56 | 0.73 |
| H | 12 | 566 | 81 | 77 | 23 | 3.12 | 40.82 | 1907.50 | 0.57 |
| H | 13 | 128 | 46 | 24 | 12 | 2.70 | 13.28 | 1494.48 | 0.51 |
| H | 14 | 159 | 37 | 10 | 3 | 1.08 | 7.95 | 712.17 | 1.04 |
| H | 15 | 557 | 47 | 101 | 14 | 0.93 | 8.39 | 770.21 | 0.69 |
| H | 16 | 330 | 45 | 118 | 7 | 1.26 | 58.49 | 1399.90 | 0.81 |
| H | 17 | 324 | 46 | 71 | 17 | 5.10 | 39.88 | 1439.27 | 0.63 |
| H | 18 | 295 | 42 | 20 | 8 | 1.27 | 40.44 | 1208.20 | 0.78 |
| H | 19 | 152 | 45 | 79 | 21 | 3.03 | 32.91 | 939.40 | 0.39 |
| H | 20 | 549 | 68 | 56 | 11 | 1.00 | 69.09 | 1859.57 | 0.41 |
We fitted generalised linear models (GLMs) to investigate differences in the richness and abundance of fruits and seeds, and the dry weight of fruits, seeds, leaves, and branches between the two habitats (forest fragment and hedgerow). We used models with a negative binomial distribution for the richness and abundance data, while for dry weight data, we used a Gamma distribution with a log link. In all these models, the response variable was the values of abundance, richness, or dry weight of fruits, seeds, leaves, and branches found in each of the 20 collectors of each habitat collected fortnightly over a year, from July 2012 to June 2013. The explanatory variable was the type of habitat (see collection data in Table
We found significant differences in fruit richness between the two habitats (χ2 = 11.87, p < 0.001), with the hedgerow exhibiting higher richness (50.70 ± 3.12 SE) than the forest fragment (35.45 ± 2.59 SE) (Fig.
The dry weight of branches and leaves was statistically higher in the hedgerow (1607.40 mg ± 193.81 SE) than in the fragment (1014.30 mg ± 92.95 SE) (χ2 = 8.950, p = 0.005) (Fig.
Finally, there was no significant difference in the diameter/length ratio of the fruits between the hedgerow (0.69 ± 0.06 SE) and the forest fragment (0.64 ± 0.04 SE) (p > 0.05).
Our results revealed that hedgerows can present greater fruit richness and higher dry weight of litter than adjacent forest fragments. Furthermore, the other parameters evaluated in these hedgerows were equivalent to those found in fragmented areas, reinforcing their importance in biodiversity conservation in rural landscapes. These findings directly address the central aim of this study, demonstrating that hedgerows, which developed over narrow trenches manually excavated by enslaved people to demarcate land boundaries, are capable of sustaining ecological processes comparable to those observed in preserved forest fragments. Over time, these trenches were colonized by vegetation and evolved into continuous forested strips, now functioning as living fences embedded within agricultural landscapes. The patterns observed in ecosystem functioning indicators reinforce the capacity of these structures to maintain key processes within fragmented landscapes, validating the study’s hypothesis and emphasizing their role in nutrient cycling, species recruitment, and ecosystem regeneration. This study is the first to assess ecosystem functioning indicators, with special emphasis on seed rain, in hedgerows within tropical landscapes. These findings offer a novel contribution to understanding the role of such structures in ecological connectivity and conservation planning.
Due to their narrow structure and distinct environmental exposure, hedgerows promote conditions that facilitate seed deposition and subsequent nutrient retention (
The increased fruit richness observed in these corridors reinforces their possible role as key areas for interactions between fauna and flora, which could promote ecological processes fundamental to the connectivity of fragmented landscapes (
The relevance of hedgerows goes beyond biodiversity conservation, as they also contribute to forest regeneration and help mitigate the impacts of environmental fragmentation and climate change (
The conservation of hedgerows should be treated as a priority in environmental governance strategies, especially in regions where remaining native vegetation is scarce and highly fragmented, such as in Minas Gerais (
Our study contributes to the understanding of the role of hedgerows in ecological connectivity, paving the way for further investigations that can expand this knowledge even further. Future research may deepen the analysis of how floristic composition influences the functionality of these corridors, as well as explore more precisely the specific role of different disperser groups. Additionally, assessing the effectiveness of public policies aimed at protecting these structures could provide more refined guidelines for their conservation. Advancing in these directions will support more effective conservation strategies, further strengthening the importance of hedgerows in biodiversity preservation.
This paper is a tribute to the first author, Angélica Aparecida Ávila, who died in the Brumadinho dam collapse on January 25, 2019. Angélica was a brilliant student whose passion for research inspired those around her. We hope that this study will not only be a valuable contribution to the understanding of seed rain but also a way to keep Angélica’s legacy alive, bringing comfort and recognition to her family and friends. Angélica Aparecida Ávila set up the field experiment and collected the data with financial support from FAPEMIG. Crislaine Paula de Oliveira thanks FAPEMIG (N° 12984/2024) for grant support. Cássio Cardoso Pereira, Crislaine Paula de Oliveira, Sérgio Gualberto Martins, and Gislene Carvalho de Castro thank the Universidade Federal de São João del-Rei (UFSJ) for its continuous support. We would like to thank Tatiana Sozzi Miguel and Sabrina Carvalho for their help in organizing the data.