Foraging behavior and diet of lined seedeaters (Sporophila lineola) in the breeding and wintering grounds

Foraging behavior and diet of lined seedeaters (Sporophila lineola) in the breeding and wintering grounds | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Foraging behavior and diet of lined seedeaters (Sporophila lineola) in the breeding and wintering grounds Alex de Matos Latorres, Tarso Natividade Ciolete, Pedro Lage Viana, and 2 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8225100/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 7 You are reading this latest preprint version Abstract Migratory animals face seasonal variations in energetic and nutritional demands, especially between breeding and wintering grounds. In this study, we investigated the foraging behavior and diet of lined seedeaters ( Sporophila lineola ) between their breeding grounds in southeastern Brazil and wintering grounds in the Amazonia. The habitat used is a disturbed dry grassland in the breeding grounds and a natural wetland in the wintering grounds. Thus, we hypothesized that attack maneuvers would vary between habitats and that food items taken would also shift seasonally, with higher arthropod intake when breeding and higher seed intake when wintering. For that we recorded 135 and 147 foraging events in the breeding and wintering grounds, respectively, also analyzing the contents of 11 stomachs of museum specimens. Lined seedeaters were more likely to use Glean and Hang-up attack maneuvers in both grounds studied, but the proportion between them differed, likely reflecting differences in grass morphology between areas. Food category composition did not differ between grounds, with seeds being the dominant item in both areas. The diet included mainly seeds of Urochloa brizantha (an introduced African grass) in the breeding grounds, and seeds of Echinochloa polystachya and Paspalum repens (two native grasses from the Amazonia wetlands) in the wintering grounds. The stomach content analysis also confirmed the predominance of grass seeds in the diet of the species. We concluded that the lined seedeater is a generalist granivore capable of exploiting both native and invasive grasses. This flexibility may offer advantages over more specialized species in a changing world. disturbed habitats granivorous birds invasive grasses migration Neotropics Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction Temporal variations in foraging behavior and diet are commonly observed in migratory animals. This is because in addition to phenological variations in their nutritional and energy demands (e.g. territorial defense, gamete production, and parental provisioning), those animals also need to cope with the challenges faced during migration, such as adverse environmental conditions, structurally distinct habitats, and different biotic interactions (Dingle 2014 ; Hurlbert and Haskell 2003 ). For example, female green sea turtles ( Chelonia mydas ) accumulate large fat reserves before migrating to the breeding grounds, where they face high energy demands associated with reproduction (Seminoff and Nichols 2002; Taquet 2006). Humpback whale ( Megaptera novaeangliae ) populations from the Southern Hemisphere seasonally migrate between wintering grounds in polar regions and breeding grounds in tropical waters. In the wintering grounds, they primarily feed on krill (Euphausiidae), accumulating energy reserves for migrating and reproducing, while in the breeding grounds they feed little or not at all (Clapham et al. 1999 ; Danilewicz et al. 2009 ). Some Nearctic migratory birds, such as the Swainson's thrush ( Catharus ustulatus ) and the Tennessee warbler ( Leiothlypis peregrina ), also exhibit seasonal changes in diet. In the breeding grounds, both species primarily feed on arthropods, whereas in the wintering grounds, the Swainson's thrush becomes more frugivorous, while the Tennessee warbler adopts a mixed diet of fruits and arthropods (Blanc-Benigeri et al. 2024 ; Salewski et al. 2002 ; Baker 1973 ; Lovette and Holmes 1995 ). Numerous studies have demonstrated marked differences in foraging behavior and diet in animals between their breeding and wintering grounds, providing valuable insights into the evolution of different foraging strategies in migratory species (Rabol 1987 ). However, such studies are rarely conducted using standardized methodologies, as the geographic distance between breeding and wintering grounds imposes logistical and financial challenges that are difficult to overcome. This is especially true in tropical regions, which chronically lack financial resources (Tydecks et al. 2018 ), creating an important knowledge gap about the life history strategies of tropical migratory species. Within this context, the lined seedeater ( Sporophila lineola ) represents an interesting model for studying potential variations in foraging behavior and diet between breeding and wintering grounds. This granivorous songbird is widely distributed in South America, occupying various types of open natural and, more often, disturbed landscapes (Ridgely and Tudor 2009 ; Ferreira et al. 2025 ). The lined seedeater is an intra-tropical migrant bird, with populations in southeastern Brazil breeding in human-made habitats from December to April (Cunha and Lopes 2022). After breeding, individuals migrate to northern South America, where they winter in open natural habitats along the Amazon floodplains, savannas, and seasonally dry shrublands (Cunha and Lopes 2022). The main goal of this study was to investigate the foraging behavior and diet of lined seedeaters in the breeding and wintering grounds. Given the likely differences in vegetation structure and food resources availability between their wintering and breeding ground, we hypothesized that the lined seedeater attack maneuvers and food items taken would differ between the breeding and the wintering grounds, a pattern already described for other bird species (Lovette and Holmes 1995 ; Gómez et al. 2018 ). Given the distinct nutritional demands exhibited during the breeding season (e.g. mate guarding, egg production, nest provisioning), we hypothesized that there would be a higher consumption of arthropods (i.e., protein-rich items items), in the breeding grounds when compared to the wintering grounds (Díaz 1996 ). Finally, we characterized the species’ diet based on data from field observations, stomach content analysis, and literature. Methods Study Areas In the breeding grounds, we studied lined seedeaters between December 2023 and April 2024 at the Universidade Federal de Viçosa – Campus Florestal (19°53'S, 44°25'W), in the municipality of Florestal, state of Minas Gerais, southeastern Brazil. This area lies in the transitional zone between the Cerrado savannas and the Atlantic Forest, consisting of fragments of semideciduous forest embedded in an open matrix of artificial pastures, active and abandoned crops, and gardens around the university’s buildings (Lopes and Marçal 2016 ). Local vegetation used by lined seedeaters is completely anthropogenic and is dominated by planted pastures with scattered shrubs, palms, exotic trees, and small forest groves. The species usually forage on disturbed areas, such as wastelands, and roadsides dominated by exotic grasses from the genus Urochloa , and exotic and native species of Echinochloa . The height and density of the herbaceous layer is highly dynamic during the breeding season due to frequent grazing, mowing, and herbicide application. In the wintering grounds, we studied lined seedeaters in August 2024 at Marchantaria Island (03°13’S, 59°54’W), municipality of Iranduba, state of Amazonas, northern Brazil. This river island, located in the Amazon floodplain, is influenced by the seasonal flood pulse of the Solimões and Negro rivers (Remsen and Parker 1983 ). The habitat used by lined seedeaters is dominated by “floating meadows” — herbaceous mats dominated by flood-adapted grasses of the genera Paspalum and Echinochloa (Junk 1970 ; Piedade 1991). There are few woody elements and almost no vertical stratification. When we visited the region, during the dry season, large expanses of floating meadows were available, providing an abundant seed supply that was easily accessible to granivorous birds. However, during the flooding season, most of this meadow is submerged, radically reducing the amount of food available to granivorous birds. The two study areas are separated by approximately 2,500 km and are both subjected to recurrent disturbances (Fig. 1 ). However, while the breeding grounds are shaped by persistent anthropogenic disturbances, the wintering grounds are shaped by the natural dynamics of the floodplain, resulting in distinct temporal patterns of habitat and food availability. Data Collection We concentrated our observations in the morning period, from 06:00 to 09:00 am, with occasional observations made in the afternoon, from 03:00 to 05:00 pm. We made observations of the foraging behavior of lined seedeaters using binoculars, thus ensuring minimal interference with the birds' natural behaviors. To describe the foraging behavior of lined seedeaters, we followed the classification scheme of Remsen and Robinson ( 1990 ), which consists of sequential components of foraging behavior. It is important to note that a single foraging event could include more than one attack maneuver. Individuals often adjusted their position on the same grass panicle as it bent under their body weight. Thus, each maneuver was recorded independently, even when performed within the same event. To investigate our hypotheses, we quantified two of these sequential components, namely the attack maneuvers used (Fig. 2 ) and the type of food items taken. Whenever possible, the arthropods caught were identified in the field with the help of binoculars to the taxonomic level of Orders. Plants consumed were photographed in the field and, whenever necessary, collected in plastic bags with proper labeling for later preparation and identification by a plant taxonomist (PLV). Plant specimens (stems with attached leaves, flowers and/or fruits) were then pressed and prepared as herbarium vouchers. After identification, these vouchers were deposited in the herbarium of the National Institute of the Atlantic Forest (INMA), in Santa Teresa, Brazil (See supplementary material). In addition to field observations, we conducted stomach content analysis of museum specimens to characterize the species’ diet. The stomach contents from 11 birds were removed from carcasses deposited in the ornithological collection at the Universidade Federal de Viçosa – Campus Florestal (UFV-CAF). These carcasses were saved during the taxidermy of birds found dead or that have been collected for other purposes. Carcasses were fixed in 10% formalin and later preserved in 70% ethanol. The stomach contents (including also crop contents, whenever available) were removed from carcasses and stored in labeled vials with 70% ethanol. Stomach content analysis was conducted under a stereomicroscope, following the procedures described by Lopes et al. ( 2005 ). Finally, to characterize the species diet, we performed a literature review with the help of three searching engines (Google Scholar, Scopus, and Web of Science), using the following keywords and their combinations: “ Sporophila lineola ”, “diet”, “foraging behavior”, “feeding ecology”, and “granivorous”. We also searched for the equivalent keywords in Portuguese and Spanish (search performed in December 2024). Data Analysis To test whether proportions of attack maneuver differed between sites (breeding vs. wintering grounds), we used a Pearson's Chi-squared test performed in R version 4.4.1 (R Core Team 2024 ), using the RStudio environment (Posit Team, 2025 ). To classify the diet type of lined seedeaters, we followed the scheme proposed by Lopes et al. ( 2016 ), which is based on the proportion of each food category consumed, as verified after stomach content analysis. This scheme considers the volumetric proportion of each food category observed in the diet, such as fruits, seeds, and insects, allowing for the determination of the species’ diet type. Results We recorded 135 attack maneuvers at the breeding grounds and 147 at the wintering grounds. Our findings confirmed that individuals employed different attack maneuvers depending on the seasonal grounds (χ² = 11.748, df = 3, p-value = 0.008). The Glean was the predominant attach maneuver in both the breeding (55.1%) and wintering (49.7%) grounds. In contrast, Hang-up was more common at the wintering grounds (38.2%) compared to the breeding site (25.3%), whereas Hang-down was relatively rare in the wintering grounds (1.3%) when compared to the breeding grounds (7.9%) (Fig. 3 ). At the breeding grounds, the most frequently consumed food item was seeds of Urochloa brizantha (93 occurrences). At the wintering grounds, the most consumed food items were seeds of Echinochloa polystachya (85 occurrences) and Paspalum repens (61 occurrences) (Fig. 4 ). Field observations combined with stomach content analyses (Appendix 1), and literature data (Table 1) revealed a predominant consumption of grass seeds by lined seedeaters in both breeding and wintering grounds. Field observations showed that 99.4% of the food items consumed by lined seedeaters consisted of seeds, while only 0.6% were arthropods. Similarly, the analysis of 22 stomach contents (Appendix 1) revealed an average volumetric composition per stomach of 98.6 ± 5.5% seeds and 1.4 ± 5.5% arthropods. Specifically, 20 individuals from different regions and periods showed diets exclusively composed of seeds, mostly Poaceae. Arthropods were only found in two out of 22 stomach contents analyzed. These findings further confirmed the species’ granivorous diet across its range, according to the scheme proposed by Lopes et al. ( 2016 ). Discussion In this study, we investigated potential differences in the foraging behavior and diet of lined seedeaters between the breeding and wintering grounds. We found statistically significant differences between the frequency of Glean and Hang-down attack maneuvers between the two study areas. Additionally, the consumption of arthropods was low or nonexistent on both sites, with no important differences between them. Finally, the species of grass seeds consumed differed notably between the two study areas. In both study areas, the Glean and Hang-up attack maneuvers predominated, with lined seedeaters capturing seeds directly from the grass panicles while perched on the stalks, never foraging on the ground. The frequency of Glean, however, was lower in the wintering grounds than in the breeding grounds. This can be attributed to significant differences in plant species composition and structure between the two sites. The greater robustness of the grasses most frequently consumed in the wintering grounds, such as Echinochloa polystachya (stalk diameter ~ 2 mm) and Paspalum repens (~ 1.7 mm), may hinder certain attack maneuvers. The stalk of those grasses is, for example, nearly three times thicker than the stalk of U. brizantha , the grass predominantly consumed in the breeding grounds, suggesting that structural features of the vegetation may constrain maneuverability and influence foraging strategies. The greater robustness of the grasses consumed in the wintering grounds hindered the bending of the stalk when the bird perched, preventing it from adopting the horizontal position necessary for executing the Glean maneuver. Differences in stalk robustness also explain why the Hang-down maneuver was more frequent in the breeding grounds, as in some cases, the stalk bent in such a way that the lined seedeaters were forced to adopt an upside-down position to forage. Lined seedeaters took seeds directly from the grass panicle, a foraging strategy that is recurrent among members of the genus Sporophila (Remsen and Hunn 1979 ; Schwartz 1975 ; Silva 1999 ), reflecting a likely adaptation to the relatively homogeneous environment and the type of resource exploited, which is abundant, static, and readily accessible. Thus, the birds adopt simple, low-energy attack maneuvers (Remsen and Robinson 1990 ), which maximize energy gain by reducing the physical effort required to obtain food (Moreno and Carrascal 1993 ). Lined seedeaters are granivorous birds, with a diet almost exclusively composed of grass seeds at both breeding and wintering grounds. This characteristic is already highlighted by the English name of the genus representatives, which are collectively known as “seedeaters" (in Brazil, several species are known as "papa-capim", which means “grass seed eaters”). Lined seedeaters do not consume arthropods in an important way during reproduction, unlike other bird species that undergo significant seasonal dietary changes (Savory 1989 ; Carey 1996 ). For instance, studies on other passerine species, such as blue tits ( Cyanistes caeruleus ) and great tits ( Parus major ), revealed important seasonal dietary changes, alternating between a mixed diet of plant and animal food items during winter and a diet predominantly composed of arthropods during the breeding season (Pollock et al. 2017 ; Serrano-Davies and Sanz 2017 ). It is worth noting that the occasional observation of lined seedeaters capturing arthropods in the breeding grounds does not necessarily imply their consumption by adults. They could have been merely carrying these arthropods to the nest to meet the nutritional needs of growing nestlings, which require higher protein intake (Siriwardena et al. 2000 ). Therefore, the actual extent of arthropod consumption by adult lined seedeaters may be even lower than reported here. In terms of the number of species consumed, grasses predominated in both the breeding and wintering grounds. Among the most consumed species, those from the genera Paspalum , Echinochloa , and Panicum stand out, not only for our study species, but also for other Neotropical seedeaters (Ilha and Ragusa-Netto 2023 ; Sales Jr. and Major 2001 ; Bencke,2010; Olifiers et al. 2001 ). The genus Paspalum seems to be particularly important due to its abundance in various habitat types and the high number of seeds produced (Maciel et al. 2009 ). Other members of Sporophila , such as the double-collared seedeater ( S. caerulescens ), the rusty-collared seedeater ( S. collaris ), and the tawny-bellied seedeater ( S. hypoxantha ) have also been reported consuming Paspalum seeds, highlighting the importance of this food resource (Ilha and Ragusa-Netto 2023 ; Areta and Almirón 2009 ; Rosoni et al. 2019 ). Seeds from Urochloa predominated in the diet of lined seedeaters in the breeding grounds, while seeds from Echinochloa and Paspalum predominated in the wintering grounds. Thus, the species of grass consumed seems to reflect the ecological conditions and resources availability at each study area. Grasses such as Echinochloa and Paspalum are widely found in both floodplain areas as well as on non-flooded areas. However, some species from these genera have specific preferences for flooded habitats, such as Echinochloa polystachya and Paspalum repens , which are abundant in floodplain habitats across much of Amazonia (Lopes and Piedade 2009 ; Piedade 1993 ). On the other hand, species like Megathyrsus maximus , Panicum fasciculatum , and U. brizantha are more frequently found in non-flooded disturbed habitats (Viciedo et al. 2019 ; Baptistella et al. 2020 ; Sales Jr. and Major 2001 ). This ecological differentiation of the consumed grass species is good indication of the feeding plasticity of lined seedeaters, which use different habitat types and food resources throughout its life cycle. The diet of lined seedeaters includes both native and invasive African grasses. Genera such as Paspalum , Setaria , and Echinochloa are widely distributed across tropical and temperate habitats worldwide, with many native species found across the distinct habitat types (Conserva and Piedade 2001 ; Viana and Filgueiras 2008 ; Filgueiras 2021 ) used by lined seedeaters across their migratory journey in South America. Thus, the migration of lined seedeater is likely associated to the distinct phenology of grasses at the breeding and wintering grounds (Areta et al. 2012; Cunha et al. 2022 ). This is because the lined seedeater, like other Sporophila species, generally does not exploit seed stocks accumulated on the ground (Remsen and Hunn 1979 ). Thus, lined seedeaters would leave the breeding grounds in search for abundant food in the floating meadows along the Amazonian floodplains, which provide abundant resources during the species' wintering period. Invasive African grasses, such as U. brizantha and U. plantaginea , are widely consumed by lined seedeaters and other farmland birds in disturbed habitats such as man-made pastures, wastelands, vacant lots, and roadsides (Baptistella et al. 2020 ; Ferreira et al. 2025 ; Sales Jr. and Major 2001 ). This preference for invasive grasses in man-made habitats aligns with the theory of biotic homogenization, where species like the lined seedeater benefit from changes induced by habitat disturbance, while species specialized in native, undisturbed habitats tend to decline (McKinney 2002 ; Smart et al. 2006 ; Callaghan et al. 2019 ; Santos et al. 2024 ). Therefore, the high consumption of invasive grasses associated with man-made habitats is a key component explaining why the lined seedeater have expanded its range so dramatically in southeastern and southern Brazil during the last century (Ferreira et al. 2025 ). The high dietary adaptability of lined seedeaters seems to contribute to its wide distribution and large population size, factors that justify its classification as "Least Concern" by the IUCN ( 2025 ). In contrast, Sporophila species more specialized in native grasses have faced significant population declines during the last decades due to habitat degradation (Franz 2012 ; IUCN 2025 ). This scenario highlights the resilience of lined seedeaters and the vulnerability of its congeners less adapted to environmental changes. Declarations Acknowledgments We thank the Lined Seedeater Project team for their help during data collection. Ismael Franz and Jonas Rosoni provided important criticism on an earlier version of this manuscript. Author contributions AML: Conceptualization, Methodology Data collection, Formal analysis, Writing-original draft (lead); TNC: Methodology, Formal analysis, Writing-review and editing; PLV: Botanic identification, Writing-review and editing; FCRC: Supervision, Formal analysis, Writing-review and editing; LEL: Conceptualization, Methodology, Supervision, Writing-review and editing. Funding Financial support to this study was provided by Coordenação de Aperfeiçoamento de Pessoal de Nível Superior-Brasil – Finance Code 001 (scholarships to AML and TNC) and Conselho Nacional de Desenvolvimento Científico e Tecnológico (research fellowship to LEL, 316960/2021-7). Data availability All data supporting the findings of this study are available within the paper and its Supplementary Information. Ethical approval Permits were granted by ICMBio (61078-1) and CEUA-UFV (03/2018). Competing interests The authors declare no competing interest References Areta JI, Almirón D (2009) Comentarios sobre la presencia, voces y alimentación del Corbatita Overo Sporophila lineola a orillas de los ríos Paraná e Iguazú en Misiones, Argentina. Cotinga 31:59–62. Areta JI (2012) Winter songs reveal geographic origin of three migratory seedeaters ( Sporophila spp.) in southern Neotropical grasslands. Wilson J Ornithol 124:688–697. https://doi.org/10.1676/1559-4491-124.4.688 Baker MC, Baker AEM (1973) Niche relationships among six species of shorebirds on their wintering and breeding ranges. Ecol Monogr 43:193–212. https://doi.org/10.2307/1942194 Baptistella JLC, Andrade SAL, Favarin JL, Mazzafera P (2020) Urochloa in tropical agroecosystems. Front Sustain Food Syst 4:119. https://doi.org/10.3389/fsufs.2020.00119 Bencke, G. A. (2010). New and significant bird records from Rio Grande do Sul, with comments on biogeography and conservation of the southern Brazilian avifauna. Iheringia. Série Zoologia , 100, 391–402. https://doi.org/10.1590/s0073-47212010000400014 Blanc-Benigeri A, Poirier V, Narango D, Elliott KH, Frei B (2024) Diet of moulting Swainson's thrushes ( Catharus ustulatus ) and Tennessee warblers ( Leiothlypis peregrina ) at a stopover site during fall migration measured with fecal DNA metabarcoding. Sci Rep 14:9913. https://doi.org/10.1038/s41598-024-59462-0 Callaghan CT, Major RE, Wilshire JH, Martin JM, Kingsford RT, Cornwell WK (2019) Generalists are the most urban-tolerant of birds: a phylogenetically controlled analysis of ecological and life history traits using a novel continuous measure of bird responses to urbanization. Oikos 128:845–858. https://doi.org/10.1111/oik.06158 Carey C (1996) Avian energetics and nutritional ecology. Chapman & Hall, London. Clapham PJ, Wetmore SE, Smith TD, Mead JG (1999) Length at birth and at independence in humpback whales. J. Cetacean Res. Manage 1:141–146. Conserva AS, Piedade MTF (2001) Ciclo de vida e ecologia de Paspalum fasciculatum Willd. ex Fluegge (Poaceae), na várzea da Amazônia central. Acta Amazonica 31:205–220. https://doi.org/10.1590/1809-43922001312220 Cunha FCR, Lopes LE, Selezneva A (2022) Revealing migration schedule and potential breeding grounds of lined seedeaters using citizen science data. Emu 122:167–175. https://doi.org/10.1080/01584197.2022.2105235 Danilewicz D, Tavares M, Moreno IB, Ott PH, Trigo CC (2009) Evidence of feeding by the humpback whale ( Megaptera novaeangliae ) in mid-latitude waters of the western South Atlantic. Mar Biodivers Rec, 2:e88. https://doi.org/10.1017/S1755267209000943 Díaz, M. (1996). Food choice by seed-eating birds in relation to seed chemistry. Comparative Biochemistry and Physiology Part A: Physiology , 113, 239–246. https://doi.org/10.1016/0300-9629(95)02093-4 Di Giacomo AG (2005) Aves de la Reserva El Bagual. In: Di Giacomo AG and Krapovickas SF (eds), História natural y paisaje de la Reserva El Bagual, Provincia de Formosa, Argentina. Asociación Ornitológica del Plata, Buenos Aires, pp. 201-465. Dingle H (2014) Migration: The biology of life on the move, 2 nd ed. Oxford University Press, Oxford. https://doi.org/10.1093/acprof:oso/9780199640386.001.0001 Ferreira LJ, Tavares PA, Cunha FCR, Solar R, Lopes LE (2025) Changes in land cover drove the recent range expansion of a tropical farmland bird. Oikos 2025:e11171. https://doi.org/10.1002/oik.11171 Filgueiras TS (2021) Gramíneas do Cerrado. Instituto Brasileiro de Geografia e Estatística, Rio de Janeiro. Franz I (2012) História natural de Sporophila hypoxantha Cabanis, 1851 (Aves: Emberizidae) em campos de altitude no sul do brasil. MSc Dissertation, Pontifícia Universidade Católica do Rio Grande do Sul, Porto Alegre. Gómez C, Larsen T, Popp B, Hobson KA, Cadena CD (2018). Assessing seasonal changes in animal diets with stable-isotope analysis of amino acids: a migratory boreal songbird switches diet over its annual cycle. Oecologia 187:1–13. https://doi.org/10.1007/s00442-018-4113-7 Hurlbert AH, Haskell JP (2003) The effect of energy and seasonality on avian species richness and community composition. Am Nat 161:83–97. https://doi.org/10.1086/345459 Ilha IMN, Ragusa-Netto J (2023) Seedeaters and seeds at a Tecoma savanna in the southern Pantanal, Brazil. Ornitol Neotrop 34:128–137. https://doi.org/10.58843/ornneo.v34i2.1003 IUCN (2025) The IUCN Red List of threatened species. International Union for Conservation of Nature. https://www.iucnredlist.org. Accessed October 2025. Junk W (1970) Investigations on the ecology and production-biology of the "floating meadows" ( Paspalo-Echinochloetum ) on the middle Amazon: Part 1: The floating vegetation and its ecology. Amazonia 2:449–495. Lopes A, Piedade MTF (2009) Estabelecimento de Echinochloa polystachya (H.B.K.) Hitchcock (Poaceae) em solo de várzea contaminado com petróleo de Urucu. Acta Amazonica 39:583–590. https://doi.org/10.1590/s0044-59672009000300013 Lopes LE, Marçal BF (2016) Avifauna do Campus Florestal da Universidade Federal de Viçosa, Minas Gerais, Brasil. Atual Ornitol 193:41–56. Lopes LE, Fernandes AM, Marini MÂ (2005) Diet of some Atlantic Forest birds. Ararajuba 13:95–103. Lopes LE, Fernandes AM, Medeiros MCI, Marini MÂ (2016) A classification scheme for avian diet types. J Field Ornithol 87:309–322. https://doi.org/10.1111/jofo.12158 Lovette IJ, Holmes RT (1995) Foraging behavior of American redstarts in breeding and wintering habitats: Implications for relative food availability. Condor 97:782–791. https://doi.org/10.2307/1369186 Maciel JR, Oliveira RC, Alves M (2009) Padrões de distribuição das espécies de Paspalum L. (Poaceae: Panicoideae: Paniceae) ocorrentes em Pernambuco, Brasil. Rev Bras Bot 32:597–605. https://doi.org/10.1590/s0100-84042009000300017 McKinney ML (2002) Urbanization, biodiversity, and conservation: The impacts of urbanization on native species are poorly studied, but educating a highly urbanized human population about these impacts can greatly improve species conservation in all ecosystems. BioScience 52:883–890. https://doi.org/10.1641/0006-3568(2002)052[0883:UBAC]2.0.CO;2 Moreno E, Carrascal LM (1993) Leg morphology and feeding postures in four Parus species: An experimental ecomorphological approach. Ecology 74:2037–2044. https://doi.org/10.2307/1940849 Olifiers N, McCain C, Prieto E, Oliveira FB, Darrigo MR (2001) Estudo da dieta de duas espécies de aves granívoras na várzea da Ilha da Marchantaria, AM. In: INPA (ed.) Curso de campo: Ecologia da Floresta Amazônica, Instituto Nacional de Pesquisas da Amazônia, Manaus, pp. 35–36. Piedade MTF (1993) Biologia e ecologia de Echinochloa polystachya (H.B.K) Hitchcock (Gramineae=Poaceae), capim semi-aquático da várzea amazônica. Acta Limnol Bras 6:173–185. Piedade MTF, Junk WJ, Long SP (1991) The productivity of the C₄ grass Echinochloa polystachya on the Amazon floodplain. Ecology 72:1456–1463. https://doi.org/10.2307/1941118 Pollock CJ, Capilla-Lasheras P, McGill RAR, Helm B, Dominoni DM (2017) Integrated behavioral and stable isotope data reveal altered diet linked to low breeding success in urban-dwelling blue tits ( Cyanistes caeruleus ). Sci Rep 7:5014. https://doi.org/10.1038/s41598-017-04575-y Posit Team (2025) RStudio: Integrated development environment for R. Posit Software. https://posit.co. Accessed September 2025. Poulin B, Lefebvre G, McNeil R (1994) Diets of land birds from northeastern Venezuela. Condor 96:354–367. https://doi.org/10.2307/1369320 R Core Team (2024) R: A language and environment for statistical computing, version 4.4.1. R Foundation for Statistical Computing. https://www.r-project.org. Accessed September 2025. Rabol J (1987) Coexistence and competition between overwintering willow warblers ( Phylloscopus trochilus ) and local warblers at Lake Naivasha, Kenya. Ornis Scandinavica , 18, 101–121. https://doi.org/10.2307/3676845 Remsen JV, Hunn EA (1979) First records of Sporophila caerulescens from Colombia; a probable long-distance migrant from southern South America. Wilson Bull 91:287–290. https://doi.org/10.5281/zenodo.16002554 Remsen JV Jr, Parker TA III (1983) Contribution of river-created habitats to bird species richness in Amazonia. Biotropica 15:223–231. http://dx.doi.org/10.2307/2387833 Remsen JV Jr, Robinson SK (1990) A classification scheme for foraging behavior of birds in terrestrial habitats. Stud Avian Biol 13:144–160. Ridgely RS, Tudor G (2009) Field guide to the songbirds of South America: the passerines. University of Texas Press, Austin. Rosoni JRR, Krügel MM, Fontana CS, Behr ER (2019) Território reprodutivo do coleiro-do-brejo ( Sporophila collaris ) no sul do Brasil: seleção e descrição de áreas de nidificação e alimentação. Ornitol Neotrop 30:33–43. https://doi.org/10.58843/ornneo.v30i0.377 Salewski V, Bairlein F, Leisler B (2002) Different wintering strategies of two Palearctic migrants in West Africa – A consequence of foraging strategies? Ibis 144:85–93. https://doi.org/10.1046/j.0019-1019.2001.00007.x Sales LG Jr, Major I (2001) Dados bio-etológicos do Sporophila lineola lineola (Emberizidae) no estado do Ceará de 1987 a 1999. In: Straube FC (ed.), Ornitologia sem fronteiras, incluindo os resumos do IX Congresso Brasileiro de Ornitologia. Pontifícia Universidade Católica do Paraná, Curitiba, pp. 341-342. Santos BA, Alvarado F, Morante-Filho JC (2024) Impacts of urbanization on multiple dimensions of bird diversity in Atlantic Forest landscapes. Global Ecol Conserv 54:e03078. https://doi.org/10.1016/j.gecco.2024.e03078 Savory CJ (1989) The importance of invertebrate food to chicks of gallinaceous species. Proc Nutr Soc: 48:113–133. https://doi.org/10.1079/pns19890015 Schubart O, Aguirre ÁC, Sick H (1965) Contribuição para o conhecimento da alimentação das aves brasileiras. Arq Zool 12:95–249. https://doi.org/10.11606/issn.2176-7793.19651295-249 Schwartz P (1975) Solved and unsolved problems in the Sporophila lineola/bouvronides complex (Aves: Emberizidae). Ann Carnegie Mus 45:277–285. https://doi.org/10.5962/p.330514 Seminoff JA, Resendiz A, Nichols WJ (2002) Diet of east Pacific green turtles ( Chelonia mydas ) in the central Gulf of California, Mexico. J Herpetol 36:447–453. https://doi.org/10.1670/0022-1511(2002)036[0447:DOEPGT]2.0.CO;2 Serrano-Davies E, Sanz JJ (2017) Habitat structure modulates nestling diet composition and fitness of blue tits Cyanistes caeruleus in the Mediterranean region. Bird Study 64:295–305. https://doi.org/10.1080/00063657.2017.1357678 Silva JMC (1999) Seasonal movements and conservation of seedeaters of the genus Sporophila in South America. Stud Avian Biol 19:272–280. Siriwardena GM, Baillie SR, Crick HQP, Wilson JD (2000) The importance of variation in the breeding performance of seed-eating birds in determining their population trends on farmland. J Appl Ecol 37:128–148. https://doi.org/10.1046/j.1365-2664.2000.00484.x Smart SM, Thompson K, Marrs RH, Le Duc MG, Maskell LC, Firbank LG (2006) Biotic homogenization and changes in species diversity across human-modified ecosystems. Proc R Soc Lond B Biol Sci 273:2659–2665. https://doi.org/10.1098/rspb.2006.3630 Taquet C, Taquet M, Dempster T, Soria M, Ciccione S, Roos D, Dagorn L (2006) Foraging of the green sea turtle Chelonia mydas on seagrass beds at Mayotte Island (Indian Ocean), determined by acoustic transmitters. Mar. Ecol. Prog. Ser 306:295–302. https://doi.org/10.3354/meps306295 Tydecks L, Jeschke JM, Wolf M, Singer G, Tockner K (2018) Spatial and topical imbalances in biodiversity research. PLOS One 13:e0199327. https://doi.org/10.1371/journal.pone.0199327 Viana PL, Filgueiras TS (2008) Inventário e distribuição geográfica das gramíneas (Poaceae) na Cadeia do Espinhaço, Brasil. Megadiversidade 4:71–88. Viciedo DO, Prado RM, Martínez CA, Habermann E, Piccolo MC (2019) Short-term warming and water stress affect Panicum maximum Jacq. stoichiometric homeostasis and biomass production. Sci Total Environ 681:267–274. https://doi.org/10.1016/j.scitotenv.2019.05.108 Table 1 Table I. Food items consumed by lined seedeaters ( Sporophila lineola ) based on literature data and field observations. The food items were marked as having been consumed at the breeding or wintering grounds, which also included individuals observed foraging along migratory pathways (i.e., transient birds). FOOD ITEM BREEDI NG WINTE RING SOURCES VEGETABLE MATTER Poaceae Seeds Andropogon bicornis X Ilha and Ragusa-Neto (2023) Axonopus argentinus X Ilha and Ragusa-Neto (2023) Digitaria horizontalis X This study Digitaria insularis X Ilha and Ragusa-Neto (2023) Echinochloa colona X This study; Sales Jr. and Major (2001) Echinochloa crus-pavonis X Bencke (2010) Echinochloa polystachya X X This study; Olifiers et al. (2001); Areta and Almirón (2009); Bencke (2010) Eleusine indica X This study Hymenachne amplexicaulis X X This study Megathyrsus maximus X This study Panicum cf. aquaticum X This study Panicum fasciculatum X Sales Jr. and Major (2001) Paspalum hydrophilum X Ilha and Ragusa-Neto (2023) Paspalum intermedium X Di Giacomo (2005) Paspalum notatum X This study; Areta and Almirón (2009) Paspalum plicatulum X This study Paspalum repens X This study; Olifiers et al. (2001) Paspalum urvillei X Di Giacomo (2005); Areta and Almirón (2009) Setaria parviflora X Ilha and Ragusa-Neto (2023) Setaria verticillata X Sales Jr. and Major (2001) Steinchisma laxum X Ilha and Ragusa-Neto (2023) Urochloa arrecta X This study Urochloa brizantha X This study Urochloa plantaginea X Sales Jr. and Major (2001) Amaranthaceae Seeds Amaranthus viridis X This study Astereacae Seeds Unidentified species X This study Cyperaceae Seeds Cyperus sp. X Sales Jr. and Major (2001) ANIMAL MATTER Unidentified spiders X This study Unidentified arthropods X Ilha and Ragusa-Neto (2023) Additional Declarations No competing interests reported. Supplementary Files LaTex.rar APPENDIX.docx Cite Share Download PDF Status: Under Review Version 1 posted Reviewers agreed at journal 25 Mar, 2026 Reviews received at journal 23 Mar, 2026 Reviewers agreed at journal 05 Mar, 2026 Reviewers invited by journal 05 Mar, 2026 Editor assigned by journal 02 Dec, 2025 Submission checks completed at journal 02 Dec, 2025 First submitted to journal 27 Nov, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. 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-8225100","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":603109239,"identity":"2a904dcd-4c12-4f5e-8d08-b1181a154476","order_by":0,"name":"Alex de Matos Latorres","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA9ElEQVRIiWNgGAWjYDACdhBRAONVADEzcwN+LcwgwoBBAsI7AxJhJEULYxuYxK+Fv5n5mNQNA7s6fumzDx/+nFcbzd8O1PKjYhtOLRKH2ZKNcwySJST70o2Nebcdz51xmLGBsefMbdzWHOYxfJxjwCxhcIaNTZpx27HcBqAWZsY23FrkD/N/OJxjUA/Swv7z55xjufMJaTE4zMMItOUw2BYG3oaa3A2EtBgeZjMG+uW45MweNmZpnmMHcjcCtRzE5xe5483PpHMqqvn5edgYP/6oqcudd/7wwQc/KvB4Hw0cBpMHiFYPBHWkKB4Fo2AUjIIRAgATz1Jo8LWFHAAAAABJRU5ErkJggg==","orcid":"","institution":"Universidade Federal de Minas Gerais","correspondingAuthor":true,"prefix":"","firstName":"Alex","middleName":"de Matos","lastName":"Latorres","suffix":""},{"id":603109240,"identity":"0c884aa2-100c-4a2e-8009-bf4d8255d86a","order_by":1,"name":"Tarso Natividade Ciolete","email":"","orcid":"","institution":"Universidade Federal de Minas Gerais","correspondingAuthor":false,"prefix":"","firstName":"Tarso","middleName":"Natividade","lastName":"Ciolete","suffix":""},{"id":603109241,"identity":"2b6940ea-8b8a-40b7-8a49-abe308c71521","order_by":2,"name":"Pedro Lage Viana","email":"","orcid":"","institution":"Instituto Nacional da Mata Atlântica","correspondingAuthor":false,"prefix":"","firstName":"Pedro","middleName":"Lage","lastName":"Viana","suffix":""},{"id":603109242,"identity":"ca7bba39-b8a1-4deb-b358-de815b8e28a1","order_by":3,"name":"Filipe C. R. Cunha","email":"","orcid":"","institution":"Wageningen University \u0026 Research","correspondingAuthor":false,"prefix":"","firstName":"Filipe","middleName":"C. R.","lastName":"Cunha","suffix":""},{"id":603109243,"identity":"bbcf2d35-46be-452c-b90e-af338b67e133","order_by":4,"name":"Leonardo Esteves Lopes","email":"","orcid":"","institution":"Universidade Federal de Viçosa – Campus Florestal","correspondingAuthor":false,"prefix":"","firstName":"Leonardo","middleName":"Esteves","lastName":"Lopes","suffix":""}],"badges":[],"createdAt":"2025-11-27 21:38:16","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-8225100/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-8225100/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":104377545,"identity":"b51c20a5-e5dc-406b-a372-0cc764fbe0e9","added_by":"auto","created_at":"2026-03-11 06:49:38","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":136930,"visible":true,"origin":"","legend":"\u003cp\u003ea) Map of South America showing the approximate location of the wintering (blue line) and breeding grounds (red lines) of lined seedeaters (\u003cem\u003eSporophila lineola\u003c/em\u003e), according to Cunha et al. (2022). The study areas are indicated by the blue star (Ilha da Marchantaria, Iranbuba, Amazonas) and the red star (Campus Florestal – Universidade Federal de Viçosa, Florestal, Minas Gerais). Satellite images of the study areas are presented for the wintering grounds (b) and for the breeding grounds (c). Satellite images obtained from Google Earth™.\u003c/p\u003e","description":"","filename":"Fig1.jpg","url":"https://assets-eu.researchsquare.com/files/rs-8225100/v1/ff0cacfa43a141ccbf1ac3de.jpg"},{"id":104780052,"identity":"f2f543d7-f32e-4323-9d39-250b343e3481","added_by":"auto","created_at":"2026-03-17 07:49:52","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":1861212,"visible":true,"origin":"","legend":"\u003cp\u003eSurface attack maneuvers used by the lined seedeater (\u003cem\u003eSporophila lineola\u003c/em\u003e), according to the classification scheme for foraging behavior developed by Remsen and Robinson (1990). Glean consists of picking up food items from a nearby substrate, without involving acrobatic movements; Hang-up consists of using the legs and toes to suspend the body, which remains hanging, with the head upwards, and can be performed on vertical (a1) or horizontal (a2) perches; Hang down consists of hanging the body head-down, and can be performed clinging on vertical (b1) or horizontal (b2) perches; Hang-sideways consists of hanging on the side of a substrate, with the body axis parallel to the ground and the bird’s side oriented upwards; Hang-upside down consists of hanging, belly-up, on underside of horizontal or diagonal perches. In addition to the attack maneuvers illustrated, the “Reach” is also recognized by Remsen and Robinson (1990). This attack maneuver is similar to Glean but involves extending completely the legs or neck to reach food. Due to the strong similarity between the two maneuvers and the difficulty of separating them in the field, both were considered synonymous in this study. Artwork by Leonardo Lopes.\u003c/p\u003e","description":"","filename":"Fig2.png","url":"https://assets-eu.researchsquare.com/files/rs-8225100/v1/00d3ca0c89cb402bf52ed14d.png"},{"id":104406136,"identity":"cc5905d8-3b95-4d33-a541-5f7168bcda92","added_by":"auto","created_at":"2026-03-11 12:24:54","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":1637022,"visible":true,"origin":"","legend":"\u003cp\u003eDifferent attack maneuvers exhibited by lined seedeaters (\u003cem\u003eSporophila lineola\u003c/em\u003e) at the breeding (above) and wintering grounds (below).\u003c/p\u003e","description":"","filename":"Fig3.png","url":"https://assets-eu.researchsquare.com/files/rs-8225100/v1/82fa0b07b47c91e38dad7af1.png"},{"id":104377546,"identity":"27ef09db-2153-4662-a84d-e4d2c0c67a90","added_by":"auto","created_at":"2026-03-11 06:49:39","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":2168203,"visible":true,"origin":"","legend":"\u003cp\u003eComparison between the types of food items consumed by lined seedeaters (\u003cem\u003eSporophila lineola\u003c/em\u003e) at the breeding grounds (top) and the wintering grounds (bottom).\u003c/p\u003e","description":"","filename":"Fig4.png","url":"https://assets-eu.researchsquare.com/files/rs-8225100/v1/15c8b18d9a96585ccf50fbd1.png"},{"id":104785735,"identity":"5ba739c3-8c72-4981-98af-3e04b395f489","added_by":"auto","created_at":"2026-03-17 08:12:54","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":5974197,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8225100/v1/2c9815f9-3018-4ea7-b9d3-e56d3d3acdc2.pdf"},{"id":104377542,"identity":"a7c8c38a-8a51-443d-b140-d6ce19c75e46","added_by":"auto","created_at":"2026-03-11 06:49:38","extension":"rar","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":945771,"visible":true,"origin":"","legend":"","description":"","filename":"LaTex.rar","url":"https://assets-eu.researchsquare.com/files/rs-8225100/v1/34ddb795d95165e7a9b2af51.rar"},{"id":104406389,"identity":"0546b146-d87b-4df4-a99c-1119f2c202a2","added_by":"auto","created_at":"2026-03-11 12:25:38","extension":"docx","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":17593,"visible":true,"origin":"","legend":"","description":"","filename":"APPENDIX.docx","url":"https://assets-eu.researchsquare.com/files/rs-8225100/v1/609cdd76c56868fca4986446.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Foraging behavior and diet of lined seedeaters (Sporophila lineola) in the breeding and wintering grounds","fulltext":[{"header":"Introduction","content":"\u003cp\u003eTemporal variations in foraging behavior and diet are commonly observed in migratory animals. This is because in addition to phenological variations in their nutritional and energy demands (e.g. territorial defense, gamete production, and parental provisioning), those animals also need to cope with the challenges faced during migration, such as adverse environmental conditions, structurally distinct habitats, and different biotic interactions (Dingle \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Hurlbert and Haskell \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2003\u003c/span\u003e). For example, female green sea turtles (\u003cem\u003eChelonia mydas\u003c/em\u003e) accumulate large fat reserves before migrating to the breeding grounds, where they face high energy demands associated with reproduction (Seminoff and Nichols 2002; Taquet 2006). Humpback whale (\u003cem\u003eMegaptera novaeangliae\u003c/em\u003e) populations from the Southern Hemisphere seasonally migrate between wintering grounds in polar regions and breeding grounds in tropical waters. In the wintering grounds, they primarily feed on krill (Euphausiidae), accumulating energy reserves for migrating and reproducing, while in the breeding grounds they feed little or not at all (Clapham et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Danilewicz et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). Some Nearctic migratory birds, such as the Swainson's thrush (\u003cem\u003eCatharus ustulatus\u003c/em\u003e) and the Tennessee warbler (\u003cem\u003eLeiothlypis peregrina\u003c/em\u003e), also exhibit seasonal changes in diet. In the breeding grounds, both species primarily feed on arthropods, whereas in the wintering grounds, the Swainson's thrush becomes more frugivorous, while the Tennessee warbler adopts a mixed diet of fruits and arthropods (Blanc-Benigeri et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Salewski et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Baker \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1973\u003c/span\u003e; Lovette and Holmes \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e1995\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eNumerous studies have demonstrated marked differences in foraging behavior and diet in animals between their breeding and wintering grounds, providing valuable insights into the evolution of different foraging strategies in migratory species (Rabol \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e1987\u003c/span\u003e). However, such studies are rarely conducted using standardized methodologies, as the geographic distance between breeding and wintering grounds imposes logistical and financial challenges that are difficult to overcome. This is especially true in tropical regions, which chronically lack financial resources (Tydecks et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), creating an important knowledge gap about the life history strategies of tropical migratory species.\u003c/p\u003e \u003cp\u003eWithin this context, the lined seedeater (\u003cem\u003eSporophila lineola\u003c/em\u003e) represents an interesting model for studying potential variations in foraging behavior and diet between breeding and wintering grounds. This granivorous songbird is widely distributed in South America, occupying various types of open natural and, more often, disturbed landscapes (Ridgely and Tudor \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Ferreira et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). The lined seedeater is an intra-tropical migrant bird, with populations in southeastern Brazil breeding in human-made habitats from December to April (Cunha and Lopes 2022). After breeding, individuals migrate to northern South America, where they winter in open natural habitats along the Amazon floodplains, savannas, and seasonally dry shrublands (Cunha and Lopes 2022). The main goal of this study was to investigate the foraging behavior and diet of lined seedeaters in the breeding and wintering grounds.\u003c/p\u003e \u003cp\u003eGiven the likely differences in vegetation structure and food resources availability between their wintering and breeding ground, we hypothesized that the lined seedeater attack maneuvers and food items taken would differ between the breeding and the wintering grounds, a pattern already described for other bird species (Lovette and Holmes \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; G\u0026oacute;mez et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Given the distinct nutritional demands exhibited during the breeding season (e.g. mate guarding, egg production, nest provisioning), we hypothesized that there would be a higher consumption of arthropods (i.e., protein-rich items items), in the breeding grounds when compared to the wintering grounds (D\u0026iacute;az \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e1996\u003c/span\u003e). Finally, we characterized the species\u0026rsquo; diet based on data from field observations, stomach content analysis, and literature.\u003c/p\u003e"},{"header":"Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eStudy Areas\u003c/h2\u003e \u003cp\u003eIn the breeding grounds, we studied lined seedeaters between December 2023 and April 2024 at the Universidade Federal de Vi\u0026ccedil;osa \u0026ndash; Campus Florestal (19\u0026deg;53'S, 44\u0026deg;25'W), in the municipality of Florestal, state of Minas Gerais, southeastern Brazil. This area lies in the transitional zone between the Cerrado savannas and the Atlantic Forest, consisting of fragments of semideciduous forest embedded in an open matrix of artificial pastures, active and abandoned crops, and gardens around the university\u0026rsquo;s buildings (Lopes and Mar\u0026ccedil;al \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Local vegetation used by lined seedeaters is completely anthropogenic and is dominated by planted pastures with scattered shrubs, palms, exotic trees, and small forest groves. The species usually forage on disturbed areas, such as wastelands, and roadsides dominated by exotic grasses from the genus \u003cem\u003eUrochloa\u003c/em\u003e, and exotic and native species of \u003cem\u003eEchinochloa\u003c/em\u003e. The height and density of the herbaceous layer is highly dynamic during the breeding season due to frequent grazing, mowing, and herbicide application.\u003c/p\u003e \u003cp\u003eIn the wintering grounds, we studied lined seedeaters in August 2024 at Marchantaria Island (03\u0026deg;13\u0026rsquo;S, 59\u0026deg;54\u0026rsquo;W), municipality of Iranduba, state of Amazonas, northern Brazil. This river island, located in the Amazon floodplain, is influenced by the seasonal flood pulse of the Solim\u0026otilde;es and Negro rivers (Remsen and Parker \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e1983\u003c/span\u003e). The habitat used by lined seedeaters is dominated by \u0026ldquo;floating meadows\u0026rdquo; \u0026mdash; herbaceous mats dominated by flood-adapted grasses of the genera \u003cem\u003ePaspalum\u003c/em\u003e and \u003cem\u003eEchinochloa\u003c/em\u003e (Junk \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e1970\u003c/span\u003e; Piedade 1991). There are few woody elements and almost no vertical stratification. When we visited the region, during the dry season, large expanses of floating meadows were available, providing an abundant seed supply that was easily accessible to granivorous birds. However, during the flooding season, most of this meadow is submerged, radically reducing the amount of food available to granivorous birds.\u003c/p\u003e \u003cp\u003eThe two study areas are separated by approximately 2,500 km and are both subjected to recurrent disturbances (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). However, while the breeding grounds are shaped by persistent anthropogenic disturbances, the wintering grounds are shaped by the natural dynamics of the floodplain, resulting in distinct temporal patterns of habitat and food availability.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eData Collection\u003c/h3\u003e\n\u003cp\u003eWe concentrated our observations in the morning period, from 06:00 to 09:00 am, with occasional observations made in the afternoon, from 03:00 to 05:00 pm. We made observations of the foraging behavior of lined seedeaters using binoculars, thus ensuring minimal interference with the birds' natural behaviors. To describe the foraging behavior of lined seedeaters, we followed the classification scheme of Remsen and Robinson (\u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e1990\u003c/span\u003e), which consists of sequential components of foraging behavior. It is important to note that a single foraging event could include more than one attack maneuver. Individuals often adjusted their position on the same grass panicle as it bent under their body weight. Thus, each maneuver was recorded independently, even when performed within the same event.\u003c/p\u003e \u003cp\u003eTo investigate our hypotheses, we quantified two of these sequential components, namely the attack maneuvers used (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e) and the type of food items taken. Whenever possible, the arthropods caught were identified in the field with the help of binoculars to the taxonomic level of Orders. Plants consumed were photographed in the field and, whenever necessary, collected in plastic bags with proper labeling for later preparation and identification by a plant taxonomist (PLV). Plant specimens (stems with attached leaves, flowers and/or fruits) were then pressed and prepared as herbarium vouchers. After identification, these vouchers were deposited in the herbarium of the National Institute of the Atlantic Forest (INMA), in Santa Teresa, Brazil (See supplementary material).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eIn addition to field observations, we conducted stomach content analysis of museum specimens to characterize the species\u0026rsquo; diet. The stomach contents from 11 birds were removed from carcasses deposited in the ornithological collection at the Universidade Federal de Vi\u0026ccedil;osa \u0026ndash; Campus Florestal (UFV-CAF). These carcasses were saved during the taxidermy of birds found dead or that have been collected for other purposes. Carcasses were fixed in 10% formalin and later preserved in 70% ethanol. The stomach contents (including also crop contents, whenever available) were removed from carcasses and stored in labeled vials with 70% ethanol. Stomach content analysis was conducted under a stereomicroscope, following the procedures described by Lopes et al. (\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). Finally, to characterize the species diet, we performed a literature review with the help of three searching engines (Google Scholar, Scopus, and Web of Science), using the following keywords and their combinations: \u0026ldquo;\u003cem\u003eSporophila lineola\u003c/em\u003e\u0026rdquo;, \u0026ldquo;diet\u0026rdquo;, \u0026ldquo;foraging behavior\u0026rdquo;, \u0026ldquo;feeding ecology\u0026rdquo;, and \u0026ldquo;granivorous\u0026rdquo;. We also searched for the equivalent keywords in Portuguese and Spanish (search performed in December 2024).\u003c/p\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003eData Analysis\u003c/h2\u003e \u003cp\u003eTo test whether proportions of attack maneuver differed between sites (breeding vs. wintering grounds), we used a Pearson's Chi-squared test performed in R version 4.4.1 (R Core Team \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), using the RStudio environment (Posit Team, \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). To classify the diet type of lined seedeaters, we followed the scheme proposed by Lopes et al. (\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), which is based on the proportion of each food category consumed, as verified after stomach content analysis. This scheme considers the volumetric proportion of each food category observed in the diet, such as fruits, seeds, and insects, allowing for the determination of the species\u0026rsquo; diet type.\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cp\u003eWe recorded 135 attack maneuvers at the breeding grounds and 147 at the wintering grounds. Our findings confirmed that individuals employed different attack maneuvers depending on the seasonal grounds (χ\u0026sup2; = 11.748, df\u0026thinsp;=\u0026thinsp;3, p-value\u0026thinsp;=\u0026thinsp;0.008). The Glean was the predominant attach maneuver in both the breeding (55.1%) and wintering (49.7%) grounds. In contrast, Hang-up was more common at the wintering grounds (38.2%) compared to the breeding site (25.3%), whereas Hang-down was relatively rare in the wintering grounds (1.3%) when compared to the breeding grounds (7.9%) (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eAt the breeding grounds, the most frequently consumed food item was seeds of \u003cem\u003eUrochloa brizantha\u003c/em\u003e (93 occurrences). At the wintering grounds, the most consumed food items were seeds of \u003cem\u003eEchinochloa polystachya\u003c/em\u003e (85 occurrences) and \u003cem\u003ePaspalum repens\u003c/em\u003e (61 occurrences) (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eField observations combined with stomach content analyses (Appendix 1), and literature data (Table\u0026nbsp;1) revealed a predominant consumption of grass seeds by lined seedeaters in both breeding and wintering grounds. Field observations showed that 99.4% of the food items consumed by lined seedeaters consisted of seeds, while only 0.6% were arthropods. Similarly, the analysis of 22 stomach contents (Appendix 1) revealed an average volumetric composition per stomach of 98.6\u0026thinsp;\u0026plusmn;\u0026thinsp;5.5% seeds and 1.4\u0026thinsp;\u0026plusmn;\u0026thinsp;5.5% arthropods. Specifically, 20 individuals from different regions and periods showed diets exclusively composed of seeds, mostly Poaceae. Arthropods were only found in two out of 22 stomach contents analyzed. These findings further confirmed the species\u0026rsquo; granivorous diet across its range, according to the scheme proposed by Lopes et al. (\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2016\u003c/span\u003e).\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eIn this study, we investigated potential differences in the foraging behavior and diet of lined seedeaters between the breeding and wintering grounds. We found statistically significant differences between the frequency of Glean and Hang-down attack maneuvers between the two study areas. Additionally, the consumption of arthropods was low or nonexistent on both sites, with no important differences between them. Finally, the species of grass seeds consumed differed notably between the two study areas.\u003c/p\u003e \u003cp\u003eIn both study areas, the Glean and Hang-up attack maneuvers predominated, with lined seedeaters capturing seeds directly from the grass panicles while perched on the stalks, never foraging on the ground. The frequency of Glean, however, was lower in the wintering grounds than in the breeding grounds. This can be attributed to significant differences in plant species composition and structure between the two sites. The greater robustness of the grasses most frequently consumed in the wintering grounds, such as \u003cem\u003eEchinochloa polystachya\u003c/em\u003e (stalk diameter\u0026thinsp;~\u0026thinsp;2 mm) and \u003cem\u003ePaspalum repens\u003c/em\u003e (~\u0026thinsp;1.7 mm), may hinder certain attack maneuvers. The stalk of those grasses is, for example, nearly three times thicker than the stalk of \u003cem\u003eU. brizantha\u003c/em\u003e, the grass predominantly consumed in the breeding grounds, suggesting that structural features of the vegetation may constrain maneuverability and influence foraging strategies. The greater robustness of the grasses consumed in the wintering grounds hindered the bending of the stalk when the bird perched, preventing it from adopting the horizontal position necessary for executing the Glean maneuver. Differences in stalk robustness also explain why the Hang-down maneuver was more frequent in the breeding grounds, as in some cases, the stalk bent in such a way that the lined seedeaters were forced to adopt an upside-down position to forage.\u003c/p\u003e \u003cp\u003eLined seedeaters took seeds directly from the grass panicle, a foraging strategy that is recurrent among members of the genus \u003cem\u003eSporophila\u003c/em\u003e (Remsen and Hunn \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e1979\u003c/span\u003e; Schwartz \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e1975\u003c/span\u003e; Silva \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e1999\u003c/span\u003e), reflecting a likely adaptation to the relatively homogeneous environment and the type of resource exploited, which is abundant, static, and readily accessible. Thus, the birds adopt simple, low-energy attack maneuvers (Remsen and Robinson \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e1990\u003c/span\u003e), which maximize energy gain by reducing the physical effort required to obtain food (Moreno and Carrascal \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1993\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eLined seedeaters are granivorous birds, with a diet almost exclusively composed of grass seeds at both breeding and wintering grounds. This characteristic is already highlighted by the English name of the genus representatives, which are collectively known as \u0026ldquo;seedeaters\" (in Brazil, several species are known as \"papa-capim\", which means \u0026ldquo;grass seed eaters\u0026rdquo;). Lined seedeaters do not consume arthropods in an important way during reproduction, unlike other bird species that undergo significant seasonal dietary changes (Savory \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e1989\u003c/span\u003e; Carey \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e1996\u003c/span\u003e). For instance, studies on other passerine species, such as blue tits (\u003cem\u003eCyanistes caeruleus\u003c/em\u003e) and great tits (\u003cem\u003eParus major\u003c/em\u003e), revealed important seasonal dietary changes, alternating between a mixed diet of plant and animal food items during winter and a diet predominantly composed of arthropods during the breeding season (Pollock et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Serrano-Davies and Sanz \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). It is worth noting that the occasional observation of lined seedeaters capturing arthropods in the breeding grounds does not necessarily imply their consumption by adults. They could have been merely carrying these arthropods to the nest to meet the nutritional needs of growing nestlings, which require higher protein intake (Siriwardena et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). Therefore, the actual extent of arthropod consumption by adult lined seedeaters may be even lower than reported here.\u003c/p\u003e \u003cp\u003eIn terms of the number of species consumed, grasses predominated in both the breeding and wintering grounds. Among the most consumed species, those from the genera \u003cem\u003ePaspalum\u003c/em\u003e, \u003cem\u003eEchinochloa\u003c/em\u003e, and \u003cem\u003ePanicum\u003c/em\u003e stand out, not only for our study species, but also for other Neotropical seedeaters (Ilha and Ragusa-Netto \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Sales Jr. and Major \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Bencke,2010; Olifiers et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). The genus \u003cem\u003ePaspalum\u003c/em\u003e seems to be particularly important due to its abundance in various habitat types and the high number of seeds produced (Maciel et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). Other members of \u003cem\u003eSporophila\u003c/em\u003e, such as the double-collared seedeater (\u003cem\u003eS. caerulescens\u003c/em\u003e), the rusty-collared seedeater (\u003cem\u003eS. collaris\u003c/em\u003e), and the tawny-bellied seedeater (\u003cem\u003eS. hypoxantha\u003c/em\u003e) have also been reported consuming \u003cem\u003ePaspalum\u003c/em\u003e seeds, highlighting the importance of this food resource (Ilha and Ragusa-Netto \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Areta and Almir\u0026oacute;n \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Rosoni et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSeeds from \u003cem\u003eUrochloa\u003c/em\u003e predominated in the diet of lined seedeaters in the breeding grounds, while seeds from \u003cem\u003eEchinochloa\u003c/em\u003e and \u003cem\u003ePaspalum\u003c/em\u003e predominated in the wintering grounds. Thus, the species of grass consumed seems to reflect the ecological conditions and resources availability at each study area. Grasses such as \u003cem\u003eEchinochloa\u003c/em\u003e and \u003cem\u003ePaspalum\u003c/em\u003e are widely found in both floodplain areas as well as on non-flooded areas. However, some species from these genera have specific preferences for flooded habitats, such as \u003cem\u003eEchinochloa polystachya\u003c/em\u003e and \u003cem\u003ePaspalum repens\u003c/em\u003e, which are abundant in floodplain habitats across much of Amazonia (Lopes and Piedade \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Piedade \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e1993\u003c/span\u003e). On the other hand, species like \u003cem\u003eMegathyrsus maximus\u003c/em\u003e, \u003cem\u003ePanicum fasciculatum\u003c/em\u003e, and \u003cem\u003eU. brizantha\u003c/em\u003e are more frequently found in non-flooded disturbed habitats (Viciedo et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Baptistella et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Sales Jr. and Major \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). This ecological differentiation of the consumed grass species is good indication of the feeding plasticity of lined seedeaters, which use different habitat types and food resources throughout its life cycle.\u003c/p\u003e \u003cp\u003eThe diet of lined seedeaters includes both native and invasive African grasses. Genera such as \u003cem\u003ePaspalum\u003c/em\u003e, \u003cem\u003eSetaria\u003c/em\u003e, and \u003cem\u003eEchinochloa\u003c/em\u003e are widely distributed across tropical and temperate habitats worldwide, with many native species found across the distinct habitat types (Conserva and Piedade \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Viana and Filgueiras \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Filgueiras \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) used by lined seedeaters across their migratory journey in South America. Thus, the migration of lined seedeater is likely associated to the distinct phenology of grasses at the breeding and wintering grounds (Areta et al. 2012; Cunha et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). This is because the lined seedeater, like other \u003cem\u003eSporophila\u003c/em\u003e species, generally does not exploit seed stocks accumulated on the ground (Remsen and Hunn \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e1979\u003c/span\u003e). Thus, lined seedeaters would leave the breeding grounds in search for abundant food in the floating meadows along the Amazonian floodplains, which provide abundant resources during the species' wintering period.\u003c/p\u003e \u003cp\u003eInvasive African grasses, such as \u003cem\u003eU. brizantha\u003c/em\u003e and \u003cem\u003eU. plantaginea\u003c/em\u003e, are widely consumed by lined seedeaters and other farmland birds in disturbed habitats such as man-made pastures, wastelands, vacant lots, and roadsides (Baptistella et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Ferreira et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Sales Jr. and Major \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). This preference for invasive grasses in man-made habitats aligns with the theory of biotic homogenization, where species like the lined seedeater benefit from changes induced by habitat disturbance, while species specialized in native, undisturbed habitats tend to decline (McKinney \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Smart et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Callaghan et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Santos et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Therefore, the high consumption of invasive grasses associated with man-made habitats is a key component explaining why the lined seedeater have expanded its range so dramatically in southeastern and southern Brazil during the last century (Ferreira et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe high dietary adaptability of lined seedeaters seems to contribute to its wide distribution and large population size, factors that justify its classification as \"Least Concern\" by the IUCN (\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). In contrast, \u003cem\u003eSporophila\u003c/em\u003e species more specialized in native grasses have faced significant population declines during the last decades due to habitat degradation (Franz \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; IUCN \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). This scenario highlights the resilience of lined seedeaters and the vulnerability of its congeners less adapted to environmental changes.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgments\u0026nbsp;\u003c/strong\u003eWe thank the Lined Seedeater Project team for their help during data collection. Ismael Franz and Jonas Rosoni provided important criticism on an earlier version of this manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor contributions\u003c/strong\u003e AML: Conceptualization, Methodology Data collection, Formal analysis, Writing-original draft (lead); TNC: Methodology, Formal analysis, Writing-review and editing; PLV: Botanic identification, Writing-review and editing; FCRC: Supervision, Formal analysis, Writing-review and editing; LEL: Conceptualization, Methodology, Supervision, Writing-review and editing.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e Financial support to this study was provided by Coordena\u0026ccedil;\u0026atilde;o de Aperfei\u0026ccedil;oamento de Pessoal de N\u0026iacute;vel Superior-Brasil \u0026ndash; Finance Code 001 (scholarships to AML and TNC) and Conselho Nacional de Desenvolvimento Cient\u0026iacute;fico e Tecnol\u0026oacute;gico (research fellowship to LEL, 316960/2021-7).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData availability\u0026nbsp;\u003c/strong\u003eAll data supporting the findings of this study are available within the paper and its Supplementary Information.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEthical approval\u003c/strong\u003e Permits were granted by ICMBio (61078-1) and CEUA-UFV (03/2018).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting interests\u003c/strong\u003e The authors declare no competing interest\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAreta JI, Almir\u0026oacute;n D (2009) Comentarios sobre la presencia, voces y alimentaci\u0026oacute;n del Corbatita Overo \u003cem\u003eSporophila lineola\u003c/em\u003e a orillas de los r\u0026iacute;os Paran\u0026aacute; e Iguaz\u0026uacute; en Misiones, Argentina. Cotinga 31:59\u0026ndash;62.\u003c/li\u003e\n\u003cli\u003eAreta JI (2012) Winter songs reveal geographic origin of three migratory seedeaters (\u003cem\u003eSporophila\u003c/em\u003e spp.) in southern Neotropical grasslands. Wilson J Ornithol 124:688\u0026ndash;697. https://doi.org/10.1676/1559-4491-124.4.688 \u003c/li\u003e\n\u003cli\u003eBaker MC, Baker AEM (1973) Niche relationships among six species of shorebirds on their wintering and breeding ranges. Ecol Monogr 43:193\u0026ndash;212. https://doi.org/10.2307/1942194 \u003c/li\u003e\n\u003cli\u003eBaptistella JLC, Andrade SAL, Favarin JL, Mazzafera P (2020) \u003cem\u003eUrochloa\u003c/em\u003e in tropical agroecosystems. Front Sustain Food Syst 4:119. https://doi.org/10.3389/fsufs.2020.00119\u003c/li\u003e\n\u003cli\u003eBencke, G. A. (2010). New and significant bird records from Rio Grande do Sul, with comments on biogeography and conservation of the southern Brazilian avifauna. \u003cem\u003eIheringia. S\u0026eacute;rie Zoologia\u003c/em\u003e, 100, 391\u0026ndash;402. https://doi.org/10.1590/s0073-47212010000400014\u003c/li\u003e\n\u003cli\u003eBlanc-Benigeri A, Poirier V, Narango D, Elliott KH, Frei B (2024) Diet of moulting Swainson\u0026apos;s thrushes (\u003cem\u003eCatharus ustulatus\u003c/em\u003e) and Tennessee warblers (\u003cem\u003eLeiothlypis peregrina\u003c/em\u003e) at a stopover site during fall migration measured with fecal DNA metabarcoding. Sci Rep 14:9913. https://doi.org/10.1038/s41598-024-59462-0\u003c/li\u003e\n\u003cli\u003eCallaghan CT, Major RE, Wilshire JH, Martin JM, Kingsford RT, Cornwell WK (2019) Generalists are the most urban-tolerant of birds: a phylogenetically controlled analysis of ecological and life history traits using a novel continuous measure of bird responses to urbanization. Oikos 128:845\u0026ndash;858. https://doi.org/10.1111/oik.06158\u003c/li\u003e\n\u003cli\u003eCarey C (1996) Avian energetics and nutritional ecology. Chapman \u0026amp; Hall, London.\u003c/li\u003e\n\u003cli\u003eClapham PJ, Wetmore SE, Smith TD, Mead JG (1999) Length at birth and at independence in humpback whales. J. Cetacean Res. Manage\u003cem\u003e \u003c/em\u003e1:141\u0026ndash;146.\u003c/li\u003e\n\u003cli\u003eConserva AS, Piedade MTF (2001) Ciclo de vida e ecologia de \u003cem\u003ePaspalum fasciculatum\u003c/em\u003e Willd. ex Fluegge (Poaceae), na v\u0026aacute;rzea da Amaz\u0026ocirc;nia central. Acta Amazonica 31:205\u0026ndash;220. https://doi.org/10.1590/1809-43922001312220\u003c/li\u003e\n\u003cli\u003eCunha FCR, Lopes LE, Selezneva A (2022) Revealing migration schedule and potential breeding grounds of lined seedeaters using citizen science data. Emu 122:167\u0026ndash;175. https://doi.org/10.1080/01584197.2022.2105235\u003c/li\u003e\n\u003cli\u003eDanilewicz D, Tavares M, Moreno IB, Ott PH, Trigo CC (2009) Evidence of feeding by the humpback whale (\u003cem\u003eMegaptera novaeangliae\u003c/em\u003e) in mid-latitude waters of the western South Atlantic. Mar Biodivers Rec, 2:e88. https://doi.org/10.1017/S1755267209000943\u003c/li\u003e\n\u003cli\u003eD\u0026iacute;az, M. (1996). Food choice by seed-eating birds in relation to seed chemistry. \u003cem\u003eComparative Biochemistry and Physiology Part A: Physiology\u003c/em\u003e, 113, 239\u0026ndash;246. https://doi.org/10.1016/0300-9629(95)02093-4\u003c/li\u003e\n\u003cli\u003eDi Giacomo AG (2005) Aves de la Reserva El Bagual. In: Di Giacomo AG and Krapovickas SF (eds), Hist\u0026oacute;ria natural y paisaje de la Reserva El Bagual, Provincia de Formosa, Argentina. Asociaci\u0026oacute;n Ornitol\u0026oacute;gica del Plata, Buenos Aires, pp. 201-465.\u003c/li\u003e\n\u003cli\u003eDingle H (2014) Migration: The biology of life on the move, 2\u003csup\u003end\u003c/sup\u003e ed. Oxford University Press, Oxford. https://doi.org/10.1093/acprof:oso/9780199640386.001.0001\u003c/li\u003e\n\u003cli\u003eFerreira LJ, Tavares PA, Cunha FCR, Solar R, Lopes LE (2025) Changes in land cover drove the recent range expansion of a tropical farmland bird. Oikos\u003cem\u003e \u003c/em\u003e2025:e11171. https://doi.org/10.1002/oik.11171\u003c/li\u003e\n\u003cli\u003eFilgueiras TS (2021) Gram\u0026iacute;neas do Cerrado. Instituto Brasileiro de Geografia e Estat\u0026iacute;stica, Rio de Janeiro.\u003c/li\u003e\n\u003cli\u003eFranz I (2012) Hist\u0026oacute;ria natural de \u003cem\u003eSporophila hypoxantha\u003c/em\u003e Cabanis, 1851 (Aves: Emberizidae) em campos de altitude no sul do brasil. MSc Dissertation, Pontif\u0026iacute;cia Universidade Cat\u0026oacute;lica do Rio Grande do Sul, Porto Alegre.\u003c/li\u003e\n\u003cli\u003eG\u0026oacute;mez C, Larsen T, Popp B, Hobson KA, Cadena CD (2018). Assessing seasonal changes in animal diets with stable-isotope analysis of amino acids: a migratory boreal songbird switches diet over its annual cycle. Oecologia 187:1\u0026ndash;13. https://doi.org/10.1007/s00442-018-4113-7\u003c/li\u003e\n\u003cli\u003eHurlbert AH, Haskell JP (2003) The effect of energy and seasonality on avian species richness and community composition. Am Nat 161:83\u0026ndash;97. https://doi.org/10.1086/345459\u003c/li\u003e\n\u003cli\u003eIlha IMN, Ragusa-Netto J (2023) Seedeaters and seeds at a \u003cem\u003eTecoma savanna\u003c/em\u003e in the southern Pantanal, Brazil. Ornitol Neotrop 34:128\u0026ndash;137. https://doi.org/10.58843/ornneo.v34i2.1003\u003c/li\u003e\n\u003cli\u003eIUCN (2025) The IUCN Red List of threatened species. International Union for Conservation of Nature. https://www.iucnredlist.org. Accessed October 2025.\u003c/li\u003e\n\u003cli\u003eJunk W (1970) Investigations on the ecology and production-biology of the \u0026quot;floating meadows\u0026quot; (\u003cem\u003ePaspalo-Echinochloetum\u003c/em\u003e) on the middle Amazon: Part 1: The floating vegetation and its ecology. Amazonia\u003cem\u003e \u003c/em\u003e2:449\u0026ndash;495. \u003c/li\u003e\n\u003cli\u003eLopes A, Piedade MTF (2009) Estabelecimento de \u003cem\u003eEchinochloa polystachya\u003c/em\u003e (H.B.K.) Hitchcock (Poaceae) em solo de v\u0026aacute;rzea contaminado com petr\u0026oacute;leo de Urucu. Acta Amazonica 39:583\u0026ndash;590. https://doi.org/10.1590/s0044-59672009000300013\u003c/li\u003e\n\u003cli\u003eLopes LE, Mar\u0026ccedil;al BF (2016) Avifauna do Campus Florestal da Universidade Federal de Vi\u0026ccedil;osa, Minas Gerais, Brasil. Atual Ornitol 193:41\u0026ndash;56.\u003c/li\u003e\n\u003cli\u003eLopes LE, Fernandes AM, Marini M\u0026Acirc; (2005) Diet of some Atlantic Forest birds. Ararajuba 13:95\u0026ndash;103. \u003c/li\u003e\n\u003cli\u003eLopes LE, Fernandes AM, Medeiros MCI, Marini M\u0026Acirc; (2016) A classification scheme for avian diet types. J Field Ornithol 87:309\u0026ndash;322. https://doi.org/10.1111/jofo.12158\u003c/li\u003e\n\u003cli\u003eLovette IJ, Holmes RT (1995) Foraging behavior of American redstarts in breeding and wintering habitats: Implications for relative food availability. Condor 97:782\u0026ndash;791. https://doi.org/10.2307/1369186\u003c/li\u003e\n\u003cli\u003eMaciel JR, Oliveira RC, Alves M (2009) Padr\u0026otilde;es de distribui\u0026ccedil;\u0026atilde;o das esp\u0026eacute;cies de \u003cem\u003ePaspalum\u003c/em\u003e L. (Poaceae: Panicoideae: Paniceae) ocorrentes em Pernambuco, Brasil. Rev Bras Bot 32:597\u0026ndash;605. https://doi.org/10.1590/s0100-84042009000300017\u003c/li\u003e\n\u003cli\u003eMcKinney ML (2002) Urbanization, biodiversity, and conservation: The impacts of urbanization on native species are poorly studied, but educating a highly urbanized human population about these impacts can greatly improve species conservation in all ecosystems. BioScience 52:883\u0026ndash;890. https://doi.org/10.1641/0006-3568(2002)052[0883:UBAC]2.0.CO;2\u003c/li\u003e\n\u003cli\u003eMoreno E, Carrascal LM (1993) Leg morphology and feeding postures in four \u003cem\u003eParus\u003c/em\u003e species: An experimental ecomorphological approach. Ecology\u003cem\u003e \u003c/em\u003e74:2037\u0026ndash;2044. https://doi.org/10.2307/1940849\u003c/li\u003e\n\u003cli\u003eOlifiers N, McCain C, Prieto E, Oliveira FB, Darrigo MR (2001) Estudo da dieta de duas esp\u0026eacute;cies de aves gran\u0026iacute;voras na v\u0026aacute;rzea da Ilha da Marchantaria, AM. In: INPA (ed.) Curso de campo: Ecologia da Floresta Amaz\u0026ocirc;nica, Instituto Nacional de Pesquisas da Amaz\u0026ocirc;nia, Manaus, pp. 35\u0026ndash;36.\u003c/li\u003e\n\u003cli\u003ePiedade MTF (1993) Biologia e ecologia de \u003cem\u003eEchinochloa polystachya\u003c/em\u003e (H.B.K) Hitchcock (Gramineae=Poaceae), capim semi-aqu\u0026aacute;tico da v\u0026aacute;rzea amaz\u0026ocirc;nica. Acta Limnol Bras\u003cem\u003e \u003c/em\u003e6:173\u0026ndash;185.\u003c/li\u003e\n\u003cli\u003ePiedade MTF, Junk WJ, Long SP (1991) The productivity of the C₄ grass \u003cem\u003eEchinochloa polystachya\u003c/em\u003e on the Amazon floodplain. Ecology 72:1456\u0026ndash;1463. https://doi.org/10.2307/1941118\u003c/li\u003e\n\u003cli\u003ePollock CJ, Capilla-Lasheras P, McGill RAR, Helm B, Dominoni DM (2017) Integrated behavioral and stable isotope data reveal altered diet linked to low breeding success in urban-dwelling blue tits (\u003cem\u003eCyanistes caeruleus\u003c/em\u003e). Sci Rep 7:5014. https://doi.org/10.1038/s41598-017-04575-y\u003c/li\u003e\n\u003cli\u003ePosit Team (2025) RStudio: Integrated development environment for R. Posit Software. https://posit.co. Accessed September 2025.\u003c/li\u003e\n\u003cli\u003ePoulin B, Lefebvre G, McNeil R (1994) Diets of land birds from northeastern Venezuela. Condor 96:354\u0026ndash;367. https://doi.org/10.2307/1369320\u003c/li\u003e\n\u003cli\u003eR Core Team (2024) R: A language and environment for statistical computing, version 4.4.1. R Foundation for Statistical Computing. https://www.r-project.org. Accessed September 2025.\u003c/li\u003e\n\u003cli\u003eRabol J (1987) Coexistence and competition between overwintering willow warblers (\u003cem\u003ePhylloscopus trochilus\u003c/em\u003e) and local warblers at Lake Naivasha, Kenya. \u003cem\u003eOrnis Scandinavica\u003c/em\u003e, 18, 101\u0026ndash;121. https://doi.org/10.2307/3676845\u003c/li\u003e\n\u003cli\u003eRemsen JV, Hunn EA (1979) First records of \u003cem\u003eSporophila caerulescens\u003c/em\u003e from Colombia; a probable long-distance migrant from southern South America. Wilson Bull 91:287\u0026ndash;290. https://doi.org/10.5281/zenodo.16002554\u003c/li\u003e\n\u003cli\u003eRemsen JV Jr, Parker TA III (1983) Contribution of river-created habitats to bird species richness in Amazonia. Biotropica 15:223\u0026ndash;231. http://dx.doi.org/10.2307/2387833\u003c/li\u003e\n\u003cli\u003eRemsen JV Jr, Robinson SK (1990) A classification scheme for foraging behavior of birds in terrestrial habitats. Stud Avian Biol\u003cem\u003e \u003c/em\u003e13:144\u0026ndash;160.\u003c/li\u003e\n\u003cli\u003eRidgely RS, Tudor G (2009) Field guide to the songbirds of South America: the passerines. University of Texas Press, Austin.\u003c/li\u003e\n\u003cli\u003eRosoni JRR, Kr\u0026uuml;gel MM, Fontana CS, Behr ER (2019) Territ\u0026oacute;rio reprodutivo do coleiro-do-brejo (\u003cem\u003eSporophila collaris\u003c/em\u003e) no sul do Brasil: sele\u0026ccedil;\u0026atilde;o e descri\u0026ccedil;\u0026atilde;o de \u0026aacute;reas de nidifica\u0026ccedil;\u0026atilde;o e alimenta\u0026ccedil;\u0026atilde;o. Ornitol Neotrop 30:33\u0026ndash;43. https://doi.org/10.58843/ornneo.v30i0.377 \u003c/li\u003e\n\u003cli\u003eSalewski V, Bairlein F, Leisler B (2002) Different wintering strategies of two Palearctic migrants in West Africa \u0026ndash; A consequence of foraging strategies? Ibis 144:85\u0026ndash;93. https://doi.org/10.1046/j.0019-1019.2001.00007.x \u003c/li\u003e\n\u003cli\u003eSales LG Jr, Major I (2001) Dados bio-etol\u0026oacute;gicos do \u003cem\u003eSporophila lineola lineola\u003c/em\u003e (Emberizidae) no estado do Cear\u0026aacute; de 1987 a 1999. In: Straube FC (ed.), Ornitologia sem fronteiras, incluindo os resumos do IX Congresso Brasileiro de Ornitologia. Pontif\u0026iacute;cia Universidade Cat\u0026oacute;lica do Paran\u0026aacute;, Curitiba, pp. 341-342.\u003c/li\u003e\n\u003cli\u003eSantos BA, Alvarado F, Morante-Filho JC (2024) Impacts of urbanization on multiple dimensions of bird diversity in Atlantic Forest landscapes. Global Ecol Conserv 54:e03078. https://doi.org/10.1016/j.gecco.2024.e03078\u003c/li\u003e\n\u003cli\u003eSavory CJ (1989) The importance of invertebrate food to chicks of gallinaceous species. Proc Nutr Soc: 48:113\u0026ndash;133. https://doi.org/10.1079/pns19890015\u003c/li\u003e\n\u003cli\u003eSchubart O, Aguirre \u0026Aacute;C, Sick H (1965) Contribui\u0026ccedil;\u0026atilde;o para o conhecimento da alimenta\u0026ccedil;\u0026atilde;o das aves brasileiras. Arq Zool 12:95\u0026ndash;249. https://doi.org/10.11606/issn.2176-7793.19651295-249\u003c/li\u003e\n\u003cli\u003eSchwartz P (1975) Solved and unsolved problems in the \u003cem\u003eSporophila lineola/bouvronides\u003c/em\u003e complex (Aves: Emberizidae). Ann Carnegie Mus 45:277\u0026ndash;285. https://doi.org/10.5962/p.330514\u003c/li\u003e\n\u003cli\u003eSeminoff JA, Resendiz A, Nichols WJ (2002) Diet of east Pacific green turtles (\u003cem\u003eChelonia mydas\u003c/em\u003e) in the central Gulf of California, Mexico. J Herpetol 36:447\u0026ndash;453. https://doi.org/10.1670/0022-1511(2002)036[0447:DOEPGT]2.0.CO;2 \u003c/li\u003e\n\u003cli\u003eSerrano-Davies E, Sanz JJ (2017) Habitat structure modulates nestling diet composition and fitness of blue tits \u003cem\u003eCyanistes caeruleus\u003c/em\u003e in the Mediterranean region. Bird Study 64:295\u0026ndash;305. https://doi.org/10.1080/00063657.2017.1357678\u003c/li\u003e\n\u003cli\u003eSilva JMC (1999) Seasonal movements and conservation of seedeaters of the genus \u003cem\u003eSporophila\u003c/em\u003e in South America. Stud Avian Biol 19:272\u0026ndash;280.\u003c/li\u003e\n\u003cli\u003eSiriwardena GM, Baillie SR, Crick HQP, Wilson JD (2000) The importance of variation in the breeding performance of seed-eating birds in determining their population trends on farmland. J Appl Ecol\u003cem\u003e \u003c/em\u003e37:128\u0026ndash;148. https://doi.org/10.1046/j.1365-2664.2000.00484.x\u003c/li\u003e\n\u003cli\u003eSmart SM, Thompson K, Marrs RH, Le Duc MG, Maskell LC, Firbank LG (2006) Biotic homogenization and changes in species diversity across human-modified ecosystems. Proc R Soc Lond B Biol Sci 273:2659\u0026ndash;2665. https://doi.org/10.1098/rspb.2006.3630\u003c/li\u003e\n\u003cli\u003eTaquet C, Taquet M, Dempster T, Soria M, Ciccione S, Roos D, Dagorn L (2006) Foraging of the green sea turtle \u003cem\u003eChelonia mydas\u003c/em\u003e on seagrass beds at Mayotte Island (Indian Ocean), determined by acoustic transmitters. Mar. Ecol. Prog. Ser\u003cem\u003e \u003c/em\u003e306:295\u0026ndash;302. https://doi.org/10.3354/meps306295\u003c/li\u003e\n\u003cli\u003eTydecks L, Jeschke JM, Wolf M, Singer G, Tockner K (2018) Spatial and topical imbalances in biodiversity research. PLOS One 13:e0199327. https://doi.org/10.1371/journal.pone.0199327\u003c/li\u003e\n\u003cli\u003eViana PL, Filgueiras TS (2008) Invent\u0026aacute;rio e distribui\u0026ccedil;\u0026atilde;o geogr\u0026aacute;fica das gram\u0026iacute;neas (Poaceae) na Cadeia do Espinha\u0026ccedil;o, Brasil. Megadiversidade 4:71\u0026ndash;88.\u003c/li\u003e\n\u003cli\u003eViciedo DO, Prado RM, Mart\u0026iacute;nez CA, Habermann E, Piccolo MC (2019) Short-term warming and water stress affect \u003cem\u003ePanicum maximum\u003c/em\u003e Jacq. stoichiometric homeostasis and biomass production. Sci Total Environ\u003cem\u003e \u003c/em\u003e681:267\u0026ndash;274. https://doi.org/10.1016/j.scitotenv.2019.05.108\u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Table 1","content":"\u003cp\u003e\u003cstrong\u003eTable I.\u003c/strong\u003e Food items consumed by lined seedeaters (\u003cem\u003eSporophila lineola\u003c/em\u003e) based on literature data and field observations. The food items were marked as having been consumed at the breeding or wintering grounds, which also included individuals observed foraging along migratory pathways (i.e., transient birds).\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eFOOD ITEM\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eBREEDI\u003c/strong\u003e\u003cstrong\u003eNG\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eWINTE\u003c/strong\u003e\u003cstrong\u003eRING\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eSOURCES\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eVEGETABLE MATTER\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cstrong\u003ePoaceae Seeds\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cem\u003eAndropogon bicornis\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eIlha and Ragusa-Neto (2023)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cem\u003eAxonopus argentinus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eIlha and Ragusa-Neto (2023)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cem\u003eDigitaria horizontalis\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eThis study\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cem\u003eDigitaria insularis\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eIlha and Ragusa-Neto (2023)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cem\u003eEchinochloa colona\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eThis study; Sales Jr. and Major (2001)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cem\u003eEchinochloa crus-pavonis\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eBencke (2010)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cem\u003eEchinochloa polystachya\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eThis study; Olifiers et al. (2001); Areta and Almir\u0026oacute;n (2009); Bencke (2010)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cem\u003eEleusine indica\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eThis study\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cem\u003eHymenachne amplexicaulis\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eThis study\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cem\u003eMegathyrsus maximus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eThis study\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cem\u003ePanicum\u0026nbsp;\u003c/em\u003ecf.\u003cem\u003e\u0026nbsp;aquaticum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eThis study\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cem\u003ePanicum fasciculatum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eSales Jr. and Major (2001)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cem\u003ePaspalum hydrophilum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eIlha and Ragusa-Neto (2023)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cem\u003ePaspalum intermedium\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eDi Giacomo (2005)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cem\u003ePaspalum notatum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eThis study; Areta and Almir\u0026oacute;n (2009)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cem\u003ePaspalum plicatulum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eThis study\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cem\u003ePaspalum repens\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eThis study; Olifiers et al. (2001)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cem\u003ePaspalum urvillei\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eDi Giacomo (2005); Areta and Almir\u0026oacute;n (2009)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cem\u003eSetaria parviflora\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eIlha and Ragusa-Neto (2023)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cem\u003eSetaria verticillata\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eSales Jr. and Major (2001)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cem\u003eSteinchisma laxum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eIlha and Ragusa-Neto (2023)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cem\u003eUrochloa arrecta\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eThis study\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cem\u003eUrochloa brizantha\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eThis study\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cem\u003eUrochloa plantaginea\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eSales Jr. and Major (2001)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eAmaranthaceae Seeds\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cem\u003eAmaranthus viridis\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eThis study\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eAstereacae Seeds\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003eUnidentified species\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eThis study\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eCyperaceae Seeds\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cem\u003eCyperus\u0026nbsp;\u003c/em\u003esp.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eSales Jr. and Major (2001)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eANIMAL MATTER\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003eUnidentified spiders\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eThis study\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003eUnidentified arthropods\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 80px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 88px;\"\u003e\n \u003cp\u003eX\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 290px;\"\u003e\n \u003cp\u003eIlha and Ragusa-Neto (2023)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\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":"disturbed habitats, granivorous birds, invasive grasses, migration, Neotropics","lastPublishedDoi":"10.21203/rs.3.rs-8225100/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-8225100/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eMigratory animals face seasonal variations in energetic and nutritional demands, especially between breeding and wintering grounds. In this study, we investigated the foraging behavior and diet of lined seedeaters (\u003cem\u003eSporophila lineola\u003c/em\u003e) between their breeding grounds in southeastern Brazil and wintering grounds in the Amazonia. The habitat used is a disturbed dry grassland in the breeding grounds and a natural wetland in the wintering grounds. Thus, we hypothesized that attack maneuvers would vary between habitats and that food items taken would also shift seasonally, with higher arthropod intake when breeding and higher seed intake when wintering. For that we recorded 135 and 147 foraging events in the breeding and wintering grounds, respectively, also analyzing the contents of 11 stomachs of museum specimens. Lined seedeaters were more likely to use Glean and Hang-up attack maneuvers in both grounds studied, but the proportion between them differed, likely reflecting differences in grass morphology between areas. Food category composition did not differ between grounds, with seeds being the dominant item in both areas. The diet included mainly seeds of \u003cem\u003eUrochloa brizantha\u003c/em\u003e (an introduced African grass) in the breeding grounds, and seeds of \u003cem\u003eEchinochloa polystachya\u003c/em\u003e and \u003cem\u003ePaspalum repens\u003c/em\u003e (two native grasses from the Amazonia wetlands) in the wintering grounds. The stomach content analysis also confirmed the predominance of grass seeds in the diet of the species. We concluded that the lined seedeater is a generalist granivore capable of exploiting both native and invasive grasses. This flexibility may offer advantages over more specialized species in a changing world.\u003c/p\u003e","manuscriptTitle":"Foraging behavior and diet of lined seedeaters (Sporophila lineola) in the breeding and wintering grounds","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-03-11 06:49:34","doi":"10.21203/rs.3.rs-8225100/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewerAgreed","content":"151215537900853276061420485985576896574","date":"2026-03-25T14:21:03+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-03-23T20:41:51+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"143339283665089625513223396882232542446","date":"2026-03-05T17:34:06+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2026-03-05T14:24:41+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-12-02T06:28:25+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-12-02T06:26:51+00:00","index":"","fulltext":""},{"type":"submitted","content":"Ornithology Research","date":"2025-11-27T21:34:14+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":"bb17cd9d-4e67-43a3-9ccd-c8790ac64d5b","owner":[],"postedDate":"March 11th, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-03-11T06:49:34+00:00","versionOfRecord":[],"versionCreatedAt":"2026-03-11 06:49:34","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-8225100","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-8225100","identity":"rs-8225100","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}