Agricultural fields structure affects landscape perception and movement decisions of understory birds

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The ability of forest-dependent species to traverse non-forested areas, such as sugarcane and Eucalyptus plantations (i.e. matrix), is crucial for maintaining population connectivity. This study investigates how matrix composition and gap distance influence the gap-crossing behavior of two understory bird species: the Variable Antshrike ( Thamnophilus caerulescens ) and the Sooty-fronted Spinetail ( Synallaxis frontalis ). We conducted playback experiments in ten sinuous forest fragments within the Brazilian Atlantic Forest, each surrounded by either sugarcane or Eucalyptus plantations. Birds were presented with two route options to reach the opposite edge: (1) a shorter path crossing the matrix or (2) a longer path along the forest edge. We hypothesized that: (a) birds would prefer routes within the forest edge over crossing the matrix; (b) the likelihood of gap crossing would decrease with increasing distance; and (c) Eucalyptus plantations would be more permeable than sugarcane. Edge-to-edge distances ranged from 30 to 130 meters. Control trials within continuous forest were also conducted for comparison. Results indicated a strong preference for forested routes ( T. caerulescens : 16% matrix crossings; S. frontalis : 10%). Additionally, a negative correlation was observed between gap distance and crossing probability, with longer distances deterring matrix crossings. Contrary to our expectations, sugarcane plantations exhibited higher permeability than Eucalyptus plantations. These findings highlight the importance of matrix composition in facilitating or impeding avian movement and emphasize its relevance for landscape-level conservation strategies.​ Atlantic Forest Thamnophilus caerulescens Synallaxis frontalis matrix permeability landscape connectivity habitat fragmentation Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction Anthropogenic activity drives habitat loss and fragmentation, leading to patch isolation that negatively influences ecological processes such as fauna movement (Ricketts 2001 , Boscolo and Metzger 2011 , Doherty et al. 2021 ) and gene flow (Carvalho et al. 2015 ). This isolation may be influenced by the presence of different land cover types, such as urban areas, agriculture, pasture, and commercial forest plantations that surround habitat patches (Lindenmayer et al. 2006 , Giubbina et al. 2018 ). Some animals may use such areas as complementary habitats that provide additional resources such as food and breeding sites (Castellón and Sieving 2006 , Driscoll et al. 2013 ). Nevertheless, these anthropogenic land cover types offer lower habitat quality compared to native habitats (Dunning et al. 1992 ). Highly hostile land covers act as barriers to movement between patches for some species, restricting population dispersal through the landscape (Antongiovanni and Metzger 2005 , Mortelliti et al. 2014 , Da Silveira et al. 2016 , Giubbina et al. 2018 ). The movement of an organism plays a key role in biodiversity patterns and determines species interactions and distributions (Schweiger et al. 2012). Faunal movement shapes genetic diversity (Shafer et al. 2012, Moraes et al. 2018), the structure and dynamics of metapopulations (Revilla et al. 2004), metacommunities (Vanschoenwinkel et al. 2008, Mokany et al. 2012), and entire ecosystems (Ellis-Soto et al. 2021 ). The landscape composition (i.e., proportion of land use types) and configuration (i.e., size, shape, and spatial arrangement of landscape elements) define landscape connectivity, which enables us to investigate how the interaction of landscape elements influences animal movement capacity and, consequently, species survival (Luque et al. 2012 ). Understanding how the configuration of landscape elements affects movement decisions and species behavior in the interpatch matrix is crucial for developing effective conservation planning, as it strongly influences habitat selection, genetic structure, and population viability (Baguette et al. 2017). Biofuels are rapidly expanding over large areas (White et al. 2008 ), and sugarcane plantations are among the most important crops for ethanol production in Brazil (Martines-Filho et al. 2006 , Rudorff et al. 2010 ). Silviculture is also growing quickly, and in Brazil, Eucalyptus plantations dominate vast regions (Kröger 2006). Some studies suggest that forestry land cover is more permeable to forest species—i.e., it facilitates faunal movement—compared to agricultural areas (Renjifo 2001 , Anderson et al. 2007 ). In this study, we evaluate the movement decisions of two forest-dependent understory bird species and, particularly, their willingness to travel through agricultural land cover (i.e., the matrix) in response to playback calls. Birds had two distinct routes to follow the playback stimulus: either through the matrix or along the forest edge. We expected that birds would respond to playback by moving through the forest edge rather than the matrix, which is assumed to present greater resistance to their movements (Da Silveira et al. 2016 ). Individuals were expected to travel more frequently across shorter distances than longer ones, due to increased vulnerability to predation associated with distance and land cover type (Yoder et al. 2004 ). Additionally, we predicted that birds would prefer to cross Eucalyptus patches over sugarcane due to the former’s vegetation structure, which may offer better protection from predators (Watling et al. 2011 , Biz et al. 2017 ). Methods Study Region The study sites are located in the Brazilian Atlantic Forest, a threatened biome that holds approximately 1–8% of all flora and fauna species in the world (Silva and Casteleti 2003 ). This biome was once one of the largest rainforests in the Americas and is known for its high environmental heterogeneity. However, it has suffered intense fragmentation—today, only 12% of the native vegetation remains, represented mainly by second-growth forest fragments isolated from each other (average distance 1,440 m) and surrounded by a mosaic of different anthropogenic land uses such as sugarcane and Eucalyptus plantations (Ribeiro et al. 2009 , Joly et al. 2014 ). We selected 10 landscapes containing a main forest fragment (< 2 ha in size), five of which were surrounded by a sugarcane matrix (approximately 2 meters high, with no elements such as isolated trees that couls act as stepping stones), and five immersed in an Eucalyptus matrix (3–5 years old, 5 meters high, with no understory development, Fig. 1 ). To test the hypothesis that birds prefer to travel along forest edges rather than across the matrix, we selected fragments with different sinous configurations surrounded by matrix (Fig. 2 ). Movement Responses of Birds The playback technique is an effective method to assess the functional connectivity of animal populations (Awade and Metzger 2008 , Bélisle and Desrochers 2001). It has been successfully used to evaluate movement and landscape perception by individuals (Bélisle and Desrochers 2001, Awade and Metzger 2008 , Shimazaki et al. 2017 ). The focal species were the forest-dependent understory birds Thamnophilus caerulescens (Variable Antshrike, Thamnophilidae) and Synallaxis frontalis (Sooty-fronted Spinetail, Furnariidae). Both species respond well to playback and are easily detected in forest fragments (Anjos and Boçon 1999 ), enabling good sampling replication. They are insectivorous, commonly found in forest edges, occur in small forest fragments, and have been observed foraging in sugarcane and Eucalyptus (Sigrist 2013 ). However, they differ in microhabitat preference: while S. frontalis favors denser vegetation and is rarely seen, T. caerulescens occupies more open areas in the understory (Sick 1997 , Souza 2004 , Sigrist 2013 , Alexandrino et al. 2016 ). The experiment was conducted from September 2014 to March 2015, which mostly corresponds to the breeding season when vocal activity is higher. Playback calls were broadcast between sunrise and 10 a.m., and from 4 p.m. until sunset. Playbacks were emitted at forest edges in sites with sinuous configurations, prompting birds to choose between a longer route inside the forest or a shortcut across the matrix to reach the source of the playback (Bélisle & Desrochers 2002 ). Each playback consisted of four 60-second vocalization series followed by a 30-second silence (Boscolo et al. 2006 ). Once the bird was sighted, another observer broadcast the playback on the opposite edge. Two additional observers, positioned with binoculars, recorded the birds' behavior while avoiding interference (Fig. 2 ). The distance between forest edges across the sinuosities ranged from 30 to 130 meters, enabling the evaluation of how far individuals were willing to cross through the matrix. Each sinuosity allowed one experiment at a distinct distance (increments of 10 meters), totaling 110 playbacks in sugarcane matrix and 110 in Eucalyptus matrix for each bird species (440 total). Experiments were randomized across the 10 landscapes, and to avoid habituation, no playback was performed at nearby points on the same day. The forest-edge route was always at least 1.5 times longer than the matrix route (Fig. 2 ). We also conducted control trials within the forest fragments to assess birds’ response distances to playback inside the habitat. These were carried out similarly to the matrix trials, using a 60-second vocalization followed by 30 seconds of silence. After detecting a response and visually locating the bird, the next playback was emitted 10 meters farther into the forest interior. Control trials ended when individuals stopped following the stimuli. We conducted five control experiments per fragment per species, totaling 50 trials. These were performed only once per day at each fragment to ensure individuals were not sampled more than once. Data analysis Crossing of individuals through the matrix or the forest-edge route from one side to the opposite edge of the sinuosity (1 = crossed matrix; 0 = forest-edge) was used as a response variable. The explanatory variables are: a) type of matrix (sugarcane or Eucalyptus ); b) distance in meters between the opposite sides of forest edges; the response variable: preference ( i.e. within the matrix vs within the fragment). We performed analyses of variance based on an F-test and used the GLM function (General Linear Model) with binomial distribution of the response variable. All analyses were made in R 3.2.0 (R Development Core Team 2015 ). Results Both bird species exhibited similar movement responses across the landscapes. Thamnophilus caerulescens and Synallaxis frontalis preferred to travel along the forest edge rather than cross the matrix ( T. caerulescens : F = 194.7, p < 0.001, df = 1, N = 220; S. frontalis : F = 24.4, p < 0.001, df = 1, N = 220). Only 16% of T. caerulescens individuals (Fig. 3 a, left column) and 10% of S. frontalis individuals (Fig. 3 a, right column) chose to travel through the matrix. Birds did not opt to cross the matrix at distances greater than 100 m. The probability of matrix crossing was negatively correlated with the distance between opposite forest edges (Fig. 3 b, left and right columns). When comparing matrix types, both species showed similar trends, with a higher probability of crossing sugarcane than Eucalyptus ( T. caerulescens : F = 34.3, p < 0.001, df = 1, N = 220; S. frontalis : F = 22.8, p < 0.001, df = 1, N = 220; Fig. 3 b). In control trials, the maximum distance traveled by T. caerulescens within forest fragments was 210 m—more than double the maximum distance traveled through the matrix (Fig. 3 c). The average distance following playback inside the forest was 145 m (SD = 1.9), compared to 60 m (SD = 2.3) in the matrix. For S. frontalis , the maximum distance in the forest was 200 m and 100 m in the matrix, with an average of 125 m (SD = 1.4) in forest control trials, and 60 m (SD = 1.3) in the matrix. When comparing gap-crossing probabilities across matrix types and control trials (Fig. 3 c), we observed a negative relationship with distance in the matrix, in contrast to control experiments, where birds crossed longer distances ( T. caerulescens : F = 19.645, p < 0.01, df = 1, N = 270; S. frontalis : F = 21.686, p < 0.01, df = 1, N = 270). Although both species crossed sugarcane more frequently than Eucalyptus, they differed in their overall route choices. T. caerulescens crossed the matrix more frequently than S. frontalis (F = 97.6, p < < 0.001, N = 440; Fig. 4 ). Additionally, T. caerulescens was more likely to move through gaps up to 100 m than S. frontalis . For distances greater than 100 m, both species equally preferred the forest edge route over the matrix. Discussion Our results indicate that both matrices formed by young Eucalyptus and sugarcane plantations exhibit low permeability to both Thamnophilus caerulescens and Synallaxis frontalis . Birds preferred to move along forest edges rather than crossing shorter paths through sugarcane or Eucalyptus matrices, supporting our hypothesis that individuals favor routes within forest edges and tend to avoid crossing anthropogenic matrices. This behavior is likely related to trade-offs associated with the configuration of remaining U-shaped edges. Such trade-offs may include higher energy expenditure, increased time spent traveling through the matrix (Hinsley 2000 ), and potential territorial defense triggered by the playback stimulus. In the case of S. frontalis , this preference may also be linked to its habit of inhabiting dark and dense vegetation, likely due to thermal and light niche preferences (Jirinec et al. 2022 ). Comparing the two matrix types, sugarcane was more permeable than Eucalyptus, contradicting our initial hypothesis that forestry matrices are generally more permeable (Watling et al. 2011 , Eycott et al. 2012 , Biz et al. 2017 ). We believe this result is due to the specific characteristics of the Eucalyptus plantations studied, which lacked any understory vegetation. The presence of scattered native trees or even early-stage understory vegetation composed of herbaceous plants and shrubs can increase bird occupancy in Eucalyptus plantations (Millan et al. 2015 ). Thus, we propose that birds were more likely to move through sugarcane (1.5 m high) because, compared to understory-free Eucalyptus plantations, sugarcane may offer a more suitable structure for understory bird movement. Furthermore, sugarcane might pose lower predation risk, as birds can hide between plants (similar to cornfields; Biz et al. 2017 ), increasing its permeability relative to Eucalyptus plantations. To better understand this difference, we observed individual behaviors while crossing both matrices. In Eucalyptus plantations, individuals moved from the forest edge understory to the canopy (approximately 5 m high), likely due to the absence of lower vegetation layers. We hypothesize that this upward movement may require more energy. In contrast, movements through sugarcane were shorter and more direct, with birds either flying between plants or making a single flight within the understory from one forest edge to the opposite. These behavioral differences suggest that sugarcane requires less energetic effort, making it more permeable than Eucalyptus. The configuration of the landscape plays a key role in determining movement success. More sinuous forest edges can facilitate movement of forest-edge specialists across the landscape (Zollner and Lima 1999), while larger interpatch distances reduce dispersal success (Awade et al. 2017 , Cornelius et al. 2017 ). The playback technique reflects home-range movement patterns (Norris and Stutchbury 2001), which typically occur over short distances related to daily activities (Awade and Metzger 2008 ). Our findings suggest that forest birds may follow remaining edges to travel distances of up to 100 m between fragments. This threshold suggests a break in habitat continuity beyond this distance, limiting functional connectivity. It may represent a critical limit for connectivity, particularly given that the average distance between fragments in the study region is ~ 190 m (Valente and Vetorazzi 2003), and the average in the Atlantic Forest is even higher (~ 1,440 m; Ribeiro et al. 2011), far exceeding the observed movement capacity of the focal species. In this study, we considered functional connectivity as the probability of crossing gaps between edges through different matrix types. However, many other landscape and species-specific factors can influence connectivity, such as patch size and shape (Prugh 2009), habitat quality, juvenile dispersal, and sex-biased dispersal (Awade et al. 2017 ). Awade and Metzger ( 2008 ) found similar results for T. caerulescens in a pasture matrix, with a maximum travel distance of 80 m between fragments—comparable to our findings in sugarcane (100 m) and Eucalyptus (90 m). In Mexico, forest birds also showed a movement threshold of 100 m (Ibarra-Macias et al. 2011 ), and in New Zealand, juvenile forest birds ( Petroica longipes ) crossed gaps up to 110 m (Richard and Armstrong 2010 ). Using a similar design in temperate Canadian forests, Bélisle and Desrochers ( 2002 ) found that birds rarely crossed sinuosities longer than 25 m, suggesting a general limitation among forest species in crossing large gaps. Comparing the two species, both T. caerulescens and S. frontalis showed a very low likelihood of crossing matrices at distances beyond 100 m, although T. caerulescens did so more frequently. This difference may reflect the distinct thermal and light niche preferences of S. frontalis , which may make it more sensitive to open matrix environments compared to T. caerulescens . Our findings contribute to the understanding of bird movement by offering new insights into the role of matrix type in shaping daily home-range movements of forest-dependent birds. The results indicate that such movements are constrained by the presence and structure of the surrounding matrix. Notably, young Eucalyptus plantations were less permeable than sugarcane, contrary to prevailing assumptions (Watling et al. 2011 ). In fact, both matrix types limited bird movement, reducing functional connectivity between fragments and potentially restricting gene flow in fragmented landscapes. Given that both T. caerulescens and S. frontalis exhibited limited willingness to cross structurally simplified matrices—such as sugarcane fields and young Eucalyptus plantations—our findings highlight the importance of maintaining short distances between forest fragments (under 100 meters) to preserve functional connectivity for understory birds. In landscapes where native habitats are embedded within matrices composed of homogeneous, low-complexity vegetation, similar to intensive agriculture or silviculture, conservation planning should consider spatial arrangements that reduce gap distances to facilitate movement. Although both matrix types restricted bird movement, we observed a slightly higher crossing frequency in sugarcane. This may reflect differences in vegetation structure at ground level, suggesting that the physical configuration of matrix habitats can influence short-distance movement behavior. Therefore, conservation efforts in fragmented landscapes should prioritize spatial configurations that minimize gap distances and reduce structural resistance to movement, even within human-modified matrices, through measures such as planting live fences or small forest patches that function as stepping stones, effectively shortening distances between fragments. Declarations Funding This work was supported by Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) under grant number 1238144, and by Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) under grant number 2013/50421-2. Disclosure of potential conflicts of interest The authors declare that they have no conflict of interest. Research involving Human Participants and/or Animals This article does not contain any studies with human participants or animals that require ethical approval. Author Contribution All authors have read and approved the content of the manuscript. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-6463312","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":473500889,"identity":"6dcde664-4bca-48e4-bb40-f9bdd29e2255","order_by":0,"name":"Marina Furlan Giubbina","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA4UlEQVRIiWNgGAWjYDAC5gMMBxgbDvCAWQwMEjKEtbAlwLQAWUAtPERpYQBqAbJ4DEB8wlp029gvHubdcUfGnP3M51c3aix4GNgPH92AT4vZMZ6Cw7xnnvFY9uRus845BnQYT1raDbxa7vckHOZtO8xjcCB3m3EOG1CLBI8Zfi3HeKBazr95Zpzzjygt7AcgWm7kMD/ObSPOFoaDc4F+MbjxzIw5t0+Ch42gX46xP/7wdscde4PzyY8/53yrk+NnP3wMrxZYdIAAmwSYxK8cBNgfwFjMHwirHgWjYBSMgpEIANctT1ms73aDAAAAAElFTkSuQmCC","orcid":"","institution":"UNESP - Universidade Estadual Paulista","correspondingAuthor":true,"prefix":"","firstName":"Marina","middleName":"Furlan","lastName":"Giubbina","suffix":""},{"id":473500893,"identity":"a3ba10a1-cf3a-4729-b4b6-e7004f6c8647","order_by":1,"name":"Marco Aurélio Pizo","email":"","orcid":"","institution":"UNESP - Universidade Estadual Paulista","correspondingAuthor":false,"prefix":"","firstName":"Marco","middleName":"Aurélio","lastName":"Pizo","suffix":""},{"id":473500894,"identity":"33cba462-684d-4325-8be4-a9ef315e4399","order_by":2,"name":"Milton Cezar Ribeiro","email":"","orcid":"","institution":"UNESP - Universidade Estadual Paulista","correspondingAuthor":false,"prefix":"","firstName":"Milton","middleName":"Cezar","lastName":"Ribeiro","suffix":""}],"badges":[],"createdAt":"2025-04-16 12:08:12","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6463312/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6463312/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":85078975,"identity":"2b02afd5-b090-4b05-a19c-cae0ff741f13","added_by":"auto","created_at":"2025-06-20 17:14:31","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":199670,"visible":true,"origin":"","legend":"\u003cp\u003eStudied landscapes. São Paulo State, Brazil; the light green areas indicate the Atlantic forest remnants; light grey areas represent the forest fragments surrounded by sugarcane matrix; orange areas indicate forest fragments with \u003cem\u003eEucalyptus\u003c/em\u003ematrix.\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-6463312/v1/1aa4be54ae1670b7280d019e.png"},{"id":85079334,"identity":"5f5b0714-ba3c-4081-887c-6544eb7c0228","added_by":"auto","created_at":"2025-06-20 17:22:32","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":716378,"visible":true,"origin":"","legend":"\u003cp\u003eExperimental scheme: the playback was reproduced in the edge of one side of a sinuosity of a forest fragment. When an individual of the focal species was seen, the same sound was reproduced on the opposite side of the curve, giving the bird two routes to follow the playback: through the forest edge (longer distance) or through the matrix (shortest distance). The control trials were carried out within the forest patch, starting at the edge and moving towards the forest interior until individuals disperse (Bélisle and Desroschers 2002).\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-6463312/v1/246d769f443212b1b1314428.png"},{"id":85078167,"identity":"950d61e5-357a-4a71-b067-52f502200b6b","added_by":"auto","created_at":"2025-06-20 17:06:32","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":299311,"visible":true,"origin":"","legend":"\u003cp\u003eLeft Column: (A) Percentages of events that individuals crossed \u003cem\u003eEucalyptus \u003c/em\u003eor sugarcane matrix. (B) Crossing probabilities of individuals in function of distance in sugarcane and \u003cem\u003eEucalyptus\u003c/em\u003ematrices, where 1 is the passage through the matrix, and 0 is through forest edge. (C) Control trials within the forest: crossing probabilities as function of distance at the beginning and end of playback, where 1 is a successful following of playback; and in both matrices, 1 is a crossing through matrix, and 0, through the forest edge. Left Column: \u003cem\u003eThamnophilus caerulescens. \u003c/em\u003eRight Column: \u003cem\u003eSynallaxis frontalis.\u003c/em\u003e\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-6463312/v1/e2361fe1873da7b6ef10a36b.png"},{"id":85078171,"identity":"7dc30edc-149f-44e4-a928-2d8c0b994179","added_by":"auto","created_at":"2025-06-20 17:06:32","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":128770,"visible":true,"origin":"","legend":"\u003cp\u003eProbabilities of crossing the matrix following playback calls for the species \u003cem\u003eThamnophilus caerulescens \u003c/em\u003eand \u003cem\u003eSynallaxis frontalis\u003c/em\u003e, where 1 is the route through matrix, and 0, through forest edge.\u003c/p\u003e","description":"","filename":"4.png","url":"https://assets-eu.researchsquare.com/files/rs-6463312/v1/90009470c6dc111fa423e04f.png"},{"id":85080007,"identity":"67f60416-4d46-4bdd-84a4-d2f754850bf5","added_by":"auto","created_at":"2025-06-20 17:30:32","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1713080,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6463312/v1/cf5d12a4-d622-4d71-a7e4-6b54da803ccd.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Agricultural fields structure affects landscape perception and movement decisions of understory birds","fulltext":[{"header":"Introduction","content":"\u003cp\u003eAnthropogenic activity drives habitat loss and fragmentation, leading to patch isolation that negatively influences ecological processes such as fauna movement (Ricketts \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2001\u003c/span\u003e, Boscolo and Metzger \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2011\u003c/span\u003e, Doherty et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) and gene flow (Carvalho et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). This isolation may be influenced by the presence of different land cover types, such as urban areas, agriculture, pasture, and commercial forest plantations that surround habitat patches (Lindenmayer et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2006\u003c/span\u003e, Giubbina et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Some animals may use such areas as complementary habitats that provide additional resources such as food and breeding sites (Castell\u0026oacute;n and Sieving \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2006\u003c/span\u003e, Driscoll et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Nevertheless, these anthropogenic land cover types offer lower habitat quality compared to native habitats (Dunning et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e1992\u003c/span\u003e). Highly hostile land covers act as barriers to movement between patches for some species, restricting population dispersal through the landscape (Antongiovanni and Metzger \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2005\u003c/span\u003e, Mortelliti et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2014\u003c/span\u003e, Da Silveira et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2016\u003c/span\u003e, Giubbina et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe movement of an organism plays a key role in biodiversity patterns and determines species interactions and distributions (Schweiger et al. 2012). Faunal movement shapes genetic diversity (Shafer et al. 2012, Moraes et al. 2018), the structure and dynamics of metapopulations (Revilla et al. 2004), metacommunities (Vanschoenwinkel et al. 2008, Mokany et al. 2012), and entire ecosystems (Ellis-Soto et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). The landscape composition (i.e., proportion of land use types) and configuration (i.e., size, shape, and spatial arrangement of landscape elements) define landscape connectivity, which enables us to investigate how the interaction of landscape elements influences animal movement capacity and, consequently, species survival (Luque et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Understanding how the configuration of landscape elements affects movement decisions and species behavior in the interpatch matrix is crucial for developing effective conservation planning, as it strongly influences habitat selection, genetic structure, and population viability (Baguette et al. 2017).\u003c/p\u003e \u003cp\u003eBiofuels are rapidly expanding over large areas (White et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2008\u003c/span\u003e), and sugarcane plantations are among the most important crops for ethanol production in Brazil (Martines-Filho et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2006\u003c/span\u003e, Rudorff et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). Silviculture is also growing quickly, and in Brazil, Eucalyptus plantations dominate vast regions (Kr\u0026ouml;ger 2006). Some studies suggest that forestry land cover is more permeable to forest species\u0026mdash;i.e., it facilitates faunal movement\u0026mdash;compared to agricultural areas (Renjifo \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2001\u003c/span\u003e, Anderson et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2007\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn this study, we evaluate the movement decisions of two forest-dependent understory bird species and, particularly, their willingness to travel through agricultural land cover (i.e., the matrix) in response to playback calls. Birds had two distinct routes to follow the playback stimulus: either through the matrix or along the forest edge. We expected that birds would respond to playback by moving through the forest edge rather than the matrix, which is assumed to present greater resistance to their movements (Da Silveira et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Individuals were expected to travel more frequently across shorter distances than longer ones, due to increased vulnerability to predation associated with distance and land cover type (Yoder et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). Additionally, we predicted that birds would prefer to cross Eucalyptus patches over sugarcane due to the former\u0026rsquo;s vegetation structure, which may offer better protection from predators (Watling et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2011\u003c/span\u003e, Biz et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e"},{"header":"Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eStudy Region\u003c/h2\u003e \u003cp\u003eThe study sites are located in the Brazilian Atlantic Forest, a threatened biome that holds approximately 1\u0026ndash;8% of all flora and fauna species in the world (Silva and Casteleti \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2003\u003c/span\u003e). This biome was once one of the largest rainforests in the Americas and is known for its high environmental heterogeneity. However, it has suffered intense fragmentation\u0026mdash;today, only 12% of the native vegetation remains, represented mainly by second-growth forest fragments isolated from each other (average distance 1,440 m) and surrounded by a mosaic of different anthropogenic land uses such as sugarcane and Eucalyptus plantations (Ribeiro et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2009\u003c/span\u003e, Joly et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2014\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eWe selected 10 landscapes containing a main forest fragment (\u0026lt;\u0026thinsp;2 ha in size), five of which were surrounded by a sugarcane matrix (approximately 2 meters high, with no elements such as isolated trees that couls act as stepping stones), and five immersed in an Eucalyptus matrix (3\u0026ndash;5 years old, 5 meters high, with no understory development, Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). To test the hypothesis that birds prefer to travel along forest edges rather than across the matrix, we selected fragments with different sinous configurations surrounded by matrix (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eMovement Responses of Birds\u003c/h3\u003e\n\u003cp\u003eThe playback technique is an effective method to assess the functional connectivity of animal populations (Awade and Metzger \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2008\u003c/span\u003e, B\u0026eacute;lisle and Desrochers 2001). It has been successfully used to evaluate movement and landscape perception by individuals (B\u0026eacute;lisle and Desrochers 2001, Awade and Metzger \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2008\u003c/span\u003e, Shimazaki et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). The focal species were the forest-dependent understory birds \u003cem\u003eThamnophilus caerulescens\u003c/em\u003e (Variable Antshrike, Thamnophilidae) and \u003cem\u003eSynallaxis frontalis\u003c/em\u003e (Sooty-fronted Spinetail, Furnariidae). Both species respond well to playback and are easily detected in forest fragments (Anjos and Bo\u0026ccedil;on \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1999\u003c/span\u003e), enabling good sampling replication. They are insectivorous, commonly found in forest edges, occur in small forest fragments, and have been observed foraging in sugarcane and Eucalyptus (Sigrist \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). However, they differ in microhabitat preference: while \u003cem\u003eS. frontalis\u003c/em\u003e favors denser vegetation and is rarely seen, \u003cem\u003eT. caerulescens\u003c/em\u003e occupies more open areas in the understory (Sick \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e1997\u003c/span\u003e, Souza \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2004\u003c/span\u003e, Sigrist \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2013\u003c/span\u003e, Alexandrino et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2016\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe experiment was conducted from September 2014 to March 2015, which mostly corresponds to the breeding season when vocal activity is higher. Playback calls were broadcast between sunrise and 10 a.m., and from 4 p.m. until sunset. Playbacks were emitted at forest edges in sites with sinuous configurations, prompting birds to choose between a longer route inside the forest or a shortcut across the matrix to reach the source of the playback (B\u0026eacute;lisle \u0026amp; Desrochers \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). Each playback consisted of four 60-second vocalization series followed by a 30-second silence (Boscolo et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). Once the bird was sighted, another observer broadcast the playback on the opposite edge. Two additional observers, positioned with binoculars, recorded the birds' behavior while avoiding interference (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe distance between forest edges across the sinuosities ranged from 30 to 130 meters, enabling the evaluation of how far individuals were willing to cross through the matrix. Each sinuosity allowed one experiment at a distinct distance (increments of 10 meters), totaling 110 playbacks in sugarcane matrix and 110 in Eucalyptus matrix for each bird species (440 total). Experiments were randomized across the 10 landscapes, and to avoid habituation, no playback was performed at nearby points on the same day. The forest-edge route was always at least 1.5 times longer than the matrix route (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eWe also conducted control trials within the forest fragments to assess birds\u0026rsquo; response distances to playback inside the habitat. These were carried out similarly to the matrix trials, using a 60-second vocalization followed by 30 seconds of silence. After detecting a response and visually locating the bird, the next playback was emitted 10 meters farther into the forest interior. Control trials ended when individuals stopped following the stimuli. We conducted five control experiments per fragment per species, totaling 50 trials. These were performed only once per day at each fragment to ensure individuals were not sampled more than once.\u003c/p\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003eData analysis\u003c/h2\u003e \u003cp\u003eCrossing of individuals through the matrix or the forest-edge route from one side to the opposite edge of the sinuosity (1\u0026thinsp;=\u0026thinsp;crossed matrix; 0\u0026thinsp;=\u0026thinsp;forest-edge) was used as a response variable. The explanatory variables are: a) type of matrix (sugarcane or \u003cem\u003eEucalyptus\u003c/em\u003e); b) distance in meters between the opposite sides of forest edges; the response variable: preference (\u003cem\u003ei.e.\u003c/em\u003e within the matrix vs within the fragment). We performed analyses of variance based on an F-test and used the GLM function (General Linear Model) with binomial distribution of the response variable. All analyses were made in R 3.2.0 (R Development Core Team \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2015\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cp\u003eBoth bird species exhibited similar movement responses across the landscapes. \u003cem\u003eThamnophilus caerulescens\u003c/em\u003e and \u003cem\u003eSynallaxis frontalis\u003c/em\u003e preferred to travel along the forest edge rather than cross the matrix (\u003cem\u003eT. caerulescens\u003c/em\u003e: F\u0026thinsp;=\u0026thinsp;194.7, p\u0026thinsp;\u0026lt;\u0026thinsp;0.001, df\u0026thinsp;=\u0026thinsp;1, N\u0026thinsp;=\u0026thinsp;220; \u003cem\u003eS. frontalis\u003c/em\u003e: F\u0026thinsp;=\u0026thinsp;24.4, p\u0026thinsp;\u0026lt;\u0026thinsp;0.001, df\u0026thinsp;=\u0026thinsp;1, N\u0026thinsp;=\u0026thinsp;220). Only 16% of \u003cem\u003eT. caerulescens\u003c/em\u003e individuals (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ea, left column) and 10% of \u003cem\u003eS. frontalis\u003c/em\u003e individuals (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ea, right column) chose to travel through the matrix. Birds did not opt to cross the matrix at distances greater than 100 m. The probability of matrix crossing was negatively correlated with the distance between opposite forest edges (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eb, left and right columns). When comparing matrix types, both species showed similar trends, with a higher probability of crossing sugarcane than Eucalyptus (\u003cem\u003eT. caerulescens\u003c/em\u003e: F\u0026thinsp;=\u0026thinsp;34.3, p\u0026thinsp;\u0026lt;\u0026thinsp;0.001, df\u0026thinsp;=\u0026thinsp;1, N\u0026thinsp;=\u0026thinsp;220; \u003cem\u003eS. frontalis\u003c/em\u003e: F\u0026thinsp;=\u0026thinsp;22.8, p\u0026thinsp;\u0026lt;\u0026thinsp;0.001, df\u0026thinsp;=\u0026thinsp;1, N\u0026thinsp;=\u0026thinsp;220; Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eb).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eIn control trials, the maximum distance traveled by \u003cem\u003eT. caerulescens\u003c/em\u003e within forest fragments was 210 m\u0026mdash;more than double the maximum distance traveled through the matrix (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ec). The average distance following playback inside the forest was 145 m (SD\u0026thinsp;=\u0026thinsp;1.9), compared to 60 m (SD\u0026thinsp;=\u0026thinsp;2.3) in the matrix. For \u003cem\u003eS. frontalis\u003c/em\u003e, the maximum distance in the forest was 200 m and 100 m in the matrix, with an average of 125 m (SD\u0026thinsp;=\u0026thinsp;1.4) in forest control trials, and 60 m (SD\u0026thinsp;=\u0026thinsp;1.3) in the matrix. When comparing gap-crossing probabilities across matrix types and control trials (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ec), we observed a negative relationship with distance in the matrix, in contrast to control experiments, where birds crossed longer distances (\u003cem\u003eT. caerulescens\u003c/em\u003e: F\u0026thinsp;=\u0026thinsp;19.645, p\u0026thinsp;\u0026lt;\u0026thinsp;0.01, df\u0026thinsp;=\u0026thinsp;1, N\u0026thinsp;=\u0026thinsp;270; \u003cem\u003eS. frontalis\u003c/em\u003e: F\u0026thinsp;=\u0026thinsp;21.686, p\u0026thinsp;\u0026lt;\u0026thinsp;0.01, df\u0026thinsp;=\u0026thinsp;1, N\u0026thinsp;=\u0026thinsp;270).\u003c/p\u003e \u003cp\u003eAlthough both species crossed sugarcane more frequently than Eucalyptus, they differed in their overall route choices. \u003cem\u003eT. caerulescens\u003c/em\u003e crossed the matrix more frequently than \u003cem\u003eS. frontalis\u003c/em\u003e (F\u0026thinsp;=\u0026thinsp;97.6, p\u0026thinsp;\u0026lt;\u0026thinsp;\u0026lt;\u0026thinsp;0.001, N\u0026thinsp;=\u0026thinsp;440; Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). Additionally, \u003cem\u003eT. caerulescens\u003c/em\u003e was more likely to move through gaps up to 100 m than \u003cem\u003eS. frontalis\u003c/em\u003e. For distances greater than 100 m, both species equally preferred the forest edge route over the matrix.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eOur results indicate that both matrices formed by young Eucalyptus and sugarcane plantations exhibit low permeability to both \u003cem\u003eThamnophilus caerulescens\u003c/em\u003e and \u003cem\u003eSynallaxis frontalis\u003c/em\u003e. Birds preferred to move along forest edges rather than crossing shorter paths through sugarcane or Eucalyptus matrices, supporting our hypothesis that individuals favor routes within forest edges and tend to avoid crossing anthropogenic matrices. This behavior is likely related to trade-offs associated with the configuration of remaining U-shaped edges. Such trade-offs may include higher energy expenditure, increased time spent traveling through the matrix (Hinsley \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2000\u003c/span\u003e), and potential territorial defense triggered by the playback stimulus. In the case of \u003cem\u003eS. frontalis\u003c/em\u003e, this preference may also be linked to its habit of inhabiting dark and dense vegetation, likely due to thermal and light niche preferences (Jirinec et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eComparing the two matrix types, sugarcane was more permeable than Eucalyptus, contradicting our initial hypothesis that forestry matrices are generally more permeable (Watling et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2011\u003c/span\u003e, Eycott et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2012\u003c/span\u003e, Biz et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). We believe this result is due to the specific characteristics of the Eucalyptus plantations studied, which lacked any understory vegetation. The presence of scattered native trees or even early-stage understory vegetation composed of herbaceous plants and shrubs can increase bird occupancy in Eucalyptus plantations (Millan et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Thus, we propose that birds were more likely to move through sugarcane (1.5 m high) because, compared to understory-free Eucalyptus plantations, sugarcane may offer a more suitable structure for understory bird movement. Furthermore, sugarcane might pose lower predation risk, as birds can hide between plants (similar to cornfields; Biz et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), increasing its permeability relative to Eucalyptus plantations.\u003c/p\u003e \u003cp\u003eTo better understand this difference, we observed individual behaviors while crossing both matrices. In Eucalyptus plantations, individuals moved from the forest edge understory to the canopy (approximately 5 m high), likely due to the absence of lower vegetation layers. We hypothesize that this upward movement may require more energy. In contrast, movements through sugarcane were shorter and more direct, with birds either flying between plants or making a single flight within the understory from one forest edge to the opposite. These behavioral differences suggest that sugarcane requires less energetic effort, making it more permeable than Eucalyptus.\u003c/p\u003e \u003cp\u003eThe configuration of the landscape plays a key role in determining movement success. More sinuous forest edges can facilitate movement of forest-edge specialists across the landscape (Zollner and Lima 1999), while larger interpatch distances reduce dispersal success (Awade et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2017\u003c/span\u003e, Cornelius et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). The playback technique reflects home-range movement patterns (Norris and Stutchbury 2001), which typically occur over short distances related to daily activities (Awade and Metzger \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Our findings suggest that forest birds may follow remaining edges to travel distances of up to 100 m between fragments. This threshold suggests a break in habitat continuity beyond this distance, limiting functional connectivity. It may represent a critical limit for connectivity, particularly given that the average distance between fragments in the study region is ~\u0026thinsp;190 m (Valente and Vetorazzi 2003), and the average in the Atlantic Forest is even higher (~\u0026thinsp;1,440 m; Ribeiro et al. 2011), far exceeding the observed movement capacity of the focal species.\u003c/p\u003e \u003cp\u003eIn this study, we considered functional connectivity as the probability of crossing gaps between edges through different matrix types. However, many other landscape and species-specific factors can influence connectivity, such as patch size and shape (Prugh 2009), habitat quality, juvenile dispersal, and sex-biased dispersal (Awade et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eAwade and Metzger (\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2008\u003c/span\u003e) found similar results for \u003cem\u003eT. caerulescens\u003c/em\u003e in a pasture matrix, with a maximum travel distance of 80 m between fragments\u0026mdash;comparable to our findings in sugarcane (100 m) and Eucalyptus (90 m). In Mexico, forest birds also showed a movement threshold of 100 m (Ibarra-Macias et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2011\u003c/span\u003e), and in New Zealand, juvenile forest birds (\u003cem\u003ePetroica longipes\u003c/em\u003e) crossed gaps up to 110 m (Richard and Armstrong \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). Using a similar design in temperate Canadian forests, B\u0026eacute;lisle and Desrochers (\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2002\u003c/span\u003e) found that birds rarely crossed sinuosities longer than 25 m, suggesting a general limitation among forest species in crossing large gaps.\u003c/p\u003e \u003cp\u003eComparing the two species, both \u003cem\u003eT. caerulescens\u003c/em\u003e and \u003cem\u003eS. frontalis\u003c/em\u003e showed a very low likelihood of crossing matrices at distances beyond 100 m, although \u003cem\u003eT. caerulescens\u003c/em\u003e did so more frequently. This difference may reflect the distinct thermal and light niche preferences of \u003cem\u003eS. frontalis\u003c/em\u003e, which may make it more sensitive to open matrix environments compared to \u003cem\u003eT. caerulescens\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eOur findings contribute to the understanding of bird movement by offering new insights into the role of matrix type in shaping daily home-range movements of forest-dependent birds. The results indicate that such movements are constrained by the presence and structure of the surrounding matrix. Notably, young Eucalyptus plantations were less permeable than sugarcane, contrary to prevailing assumptions (Watling et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). In fact, both matrix types limited bird movement, reducing functional connectivity between fragments and potentially restricting gene flow in fragmented landscapes.\u003c/p\u003e \u003cp\u003eGiven that both \u003cem\u003eT. caerulescens\u003c/em\u003e and \u003cem\u003eS. frontalis\u003c/em\u003e exhibited limited willingness to cross structurally simplified matrices\u0026mdash;such as sugarcane fields and young Eucalyptus plantations\u0026mdash;our findings highlight the importance of maintaining short distances between forest fragments (under 100 meters) to preserve functional connectivity for understory birds. In landscapes where native habitats are embedded within matrices composed of homogeneous, low-complexity vegetation, similar to intensive agriculture or silviculture, conservation planning should consider spatial arrangements that reduce gap distances to facilitate movement. Although both matrix types restricted bird movement, we observed a slightly higher crossing frequency in sugarcane. This may reflect differences in vegetation structure at ground level, suggesting that the physical configuration of matrix habitats can influence short-distance movement behavior. Therefore, conservation efforts in fragmented landscapes should prioritize spatial configurations that minimize gap distances and reduce structural resistance to movement, even within human-modified matrices, through measures such as planting live fences or small forest patches that function as stepping stones, effectively shortening distances between fragments.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis work was supported by Coordena\u0026ccedil;\u0026atilde;o de Aperfei\u0026ccedil;oamento de Pessoal de N\u0026iacute;vel Superior (CAPES) under grant number 1238144, and by Funda\u0026ccedil;\u0026atilde;o de Amparo \u0026agrave; Pesquisa do Estado de S\u0026atilde;o Paulo (FAPESP) under grant number 2013/50421-2.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eDisclosure of potential conflicts of interest\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare that they have no conflict of interest.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eResearch involving Human Participants and/or Animals\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis article does not contain any studies with human participants or animals that require ethical approval.\u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eAll authors have read and approved the content of the manuscript. MFG conducted data collection, analysis, prepared the figures and wrote the main manucript text. All authors reviewed the manuscript.\u003c/p\u003e\u003ch2\u003eAcknowledgement\u003c/h2\u003e\u003cp\u003eWe would like to thank the LEEC team for providing suggestions on the field study and the Department of Biodiversity technical personnel, Cau\u0026ecirc; Tomaz, Harold Fowler, Felipe B\u0026uacute;falo, Vitor Marin, Felipe Tassi, Thiago de Castro Ribeiro, Georg Beckman, Kaizer Alves and Fernando Andrioli for their support during the realization of this study. Megan King, a Canadian native English speaker proof reading this manuscript.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAlexandrino ER, Buechley ER, Piratelli AJ, Ferraz KMPMB, Moral MA, Sekercioglu CH, Silva WR, Couto HTZ (2016) Bird sensitivity to disturbance as an indicator of forest patch conditions: An issue in environmental assessments. 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For Ecol Manage 336:174\u0026ndash;172.\u003c/li\u003e\n\u003cli\u003eMortelliti A, Westgate MJ, Lindenmayer DB (2014) Experimental evaluation shows limited influence of pine plantations on the connectivity of highly fragmented bird populations. J Appl Ecol 51:1179\u0026ndash;1187.\u003c/li\u003e\n\u003cli\u003eR Development Core Team (2015) R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna.\u003c/li\u003e\n\u003cli\u003eRenjifo LM (2001) Effect of natural and anthropogenic landscape matrices on the abundance of subandean bird species. Ecol Appl 11:14\u0026ndash;31.\u003c/li\u003e\n\u003cli\u003eRibeiro MC, Metzger JP, Martensen AC, Ponzoni FJ, Hirota MM (2009) The Brazilian Atlantic Forest: how much is left, and how is the remaining forest distributed? Implications for conservation. 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Avian Conserv Ecol 12:Article-16.\u003c/li\u003e\n\u003cli\u003eSigrist T (2013) Avifauna Brasileira: The Avis Brasilis Field Guide to the Birds of Brazil, 1st edn. Avis Brasilis, S\u0026atilde;o Paulo.\u003c/li\u003e\n\u003cli\u003eSilva JMC, Casteleti CHMC (2003) Status of the biodiversity of the Atlantic Forest of Brazil. In: Galindo-Leal C, C\u0026acirc;mara IG (eds) The Atlantic Forest of South America: Biodiversity Status, Threats, and Outlooks. Island Press, Washington, DC, pp 43\u0026ndash;59.\u003c/li\u003e\n\u003cli\u003eSick H (1997) Ornitologia Brasileira. Nova Fronteira, Rio de Janeiro.\u003c/li\u003e\n\u003cli\u003eSouza D (2004) Todas as Aves do Brasil: Guia de Campo para Identifica\u0026ccedil;\u0026atilde;o. DALL, Gr\u0026aacute;fica Liceu, Bahia.\u003c/li\u003e\n\u003cli\u003eValente ROA, Vettorazzi CA (2005) Avalia\u0026ccedil;\u0026atilde;o da estrutura florestal na bacia hidrogr\u0026aacute;fica do Rio Corumbata\u0026iacute;, SP. Sci For 68:45\u0026ndash;57.\u003c/li\u003e\n\u003cli\u003eWatling JI, Nowakowski AJ, Donnelly MA, Orrock JL (2011) Meta-analysis reveals the importance of matrix composition for animals in fragmented habitat. Glob Ecol Biogeogr 20:209\u0026ndash;217.\u003c/li\u003e\n\u003cli\u003eWhite A, Molnar A, Khare A, Sunderlin W (2008) Seeing people through the trees: scaling up efforts to advance rights and address poverty, conflict and climate change. Rights and Resources Initiative, Washington, DC.\u003c/li\u003e\n\u003cli\u003eYoder JM, Marshall EA, Swanson DA (2004) The cost of dispersal: predation as a function of movement and site familiarity in ruffed grouse. Behav Ecol 15:469\u0026ndash;476.\u003c/li\u003e\n\u003cli\u003eZollner PA, Lima SL (2005) Behavioral trade-offs when dispersing across a patchy landscape. Oikos 108:219\u0026ndash;230.\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"ornithology-research","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"orni","sideBox":"Learn more about [Ornithology Research](https://link.springer.com/journal/43388)","snPcode":"43388","submissionUrl":"https://submission.nature.com/new-submission/43388/3","title":"Ornithology Research","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Atlantic Forest, Thamnophilus caerulescens, Synallaxis frontalis, matrix permeability, landscape connectivity, habitat fragmentation","lastPublishedDoi":"10.21203/rs.3.rs-6463312/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6463312/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eHabitat fragmentation in forest landscapes, often resulting from agricultural expansion, poses significant challenges to wildlife movement and persistence. The ability of forest-dependent species to traverse non-forested areas, such as sugarcane and Eucalyptus plantations (i.e. matrix), is crucial for maintaining population connectivity. This study investigates how matrix composition and gap distance influence the gap-crossing behavior of two understory bird species: the Variable Antshrike (\u003cem\u003eThamnophilus caerulescens\u003c/em\u003e) and the Sooty-fronted Spinetail (\u003cem\u003eSynallaxis frontalis\u003c/em\u003e). We conducted playback experiments in ten sinuous forest fragments within the Brazilian Atlantic Forest, each surrounded by either sugarcane or Eucalyptus plantations. Birds were presented with two route options to reach the opposite edge: (1) a shorter path crossing the matrix or (2) a longer path along the forest edge. We hypothesized that: (a) birds would prefer routes within the forest edge over crossing the matrix; (b) the likelihood of gap crossing would decrease with increasing distance; and (c) Eucalyptus plantations would be more permeable than sugarcane. Edge-to-edge distances ranged from 30 to 130 meters. Control trials within continuous forest were also conducted for comparison. Results indicated a strong preference for forested routes (\u003cem\u003eT. caerulescens\u003c/em\u003e: 16% matrix crossings; \u003cem\u003eS. frontalis\u003c/em\u003e: 10%). Additionally, a negative correlation was observed between gap distance and crossing probability, with longer distances deterring matrix crossings. Contrary to our expectations, sugarcane plantations exhibited higher permeability than Eucalyptus plantations. These findings highlight the importance of matrix composition in facilitating or impeding avian movement and emphasize its relevance for landscape-level conservation strategies.​\u003c/p\u003e","manuscriptTitle":"Agricultural fields structure affects landscape perception and movement decisions of understory birds","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-06-20 17:06:27","doi":"10.21203/rs.3.rs-6463312/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-11-03T21:54:43+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-10-07T17:46:58+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"191410758196924516319040138968872765000","date":"2025-09-05T17:39:57+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-08-08T13:04:40+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"104244598020883153194098167125813317861","date":"2025-07-18T13:50:21+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-06-18T18:02:39+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-05-02T08:34:47+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-05-02T08:31:52+00:00","index":"","fulltext":""},{"type":"submitted","content":"Ornithology Research","date":"2025-04-16T11:54:19+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"ornithology-research","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"orni","sideBox":"Learn more about [Ornithology Research](https://link.springer.com/journal/43388)","snPcode":"43388","submissionUrl":"https://submission.nature.com/new-submission/43388/3","title":"Ornithology Research","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"09d4ce62-4dda-44c6-9972-6b70a65a33ce","owner":[],"postedDate":"June 20th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-05-14T19:53:09+00:00","versionOfRecord":[],"versionCreatedAt":"2025-06-20 17:06:27","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-6463312","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6463312","identity":"rs-6463312","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}