Conservation Challenges of the Hyacinth Macaw (Anodorhynchus hyacinthinus (Latham, 1790)) in Brazil: An Integrative Review

Conservation Challenges of the Hyacinth Macaw (Anodorhynchus hyacinthinus (Latham, 1790)) in Brazil: An Integrative Review | 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 Systematic Review Conservation Challenges of the Hyacinth Macaw (Anodorhynchus hyacinthinus (Latham, 1790)) in Brazil: An Integrative Review Vitor Matheus Morais de Oliveira, Vinicíus de Avelar São Pedro This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8695459/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 4 You are reading this latest preprint version Abstract The Hyacinth Macaw ( Anodorhynchus hyacinthinus ), the world's largest psittacid is threatened by illegal trade and habitat loss. Conservation efforts in Brazil, particularly the Arara Azul Project, have contributed to the recovery of the Hyacinth Macaw population in the Pantanal. However, scientific knowledge on the species’ conservation remains scattered across the literature, which hampers the development of effective long-term strategies. This study aimed to systematize and critically analyze scientific research on the Hyacinth Macaw in Brazil through an integrative review. Seventy one papers were retrieved from six academic databases. The results revealed a significant increase in research in the past three decades, with a predominance of empirical studies, particularly in ecology, most of them conducted in the southern Pantanal. The reviewed literature indicates that Hyacinth Macaw populations exhibit recent gene flow and relatively high genetic diversity compared with other threatened psittacid species. Additionally, interpopulation variations were identified in diet and reproductive behavior, depending on the region. Major knowledge gaps include the lack of studies on the species’ ecology and behavior in the Central and Northern regions of the country, as well as the limited number of studies involving individuals in captivity. These gaps limit accurate demographic, genetic, and ecological assessments, as well as conservation planning. The findings highlight the need for region-specific approaches to conservation and emphasize the central role of the Arara Azul Institute in scientific research and population management. Despite significant progress, important gaps remain in understudied populations, requiring targeted conservation strategies and actions. Psittacidae animal conservation bird ecology extinction threats Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 1. INTRODUCTION The family Psittacidae is among the most threatened bird families worldwide, with 29% of its species classified as globally threatened and 58% showing declining population trends (Chan et al. 2021 ). Due to their vibrant plumage, high intelligence, longevity, and the ability to mimic the human voice, psittacids have become frequent targets of illegal wildlife trade. These attributes make them highly desirable as pets, generating thousands of dollars annually (Olah et al. 2016 ; Chan et al. 2021 ). The Hyacinth Macaw ( Anodorhynchus hyacinthinus (Latham, 1790)) is the largest species in the family Psittacidae, reaching approximately 1 m in length (Guedes et al. 2022 ). The species has a wide distribution in Brazil, occurring in three main biomes: the Pantanal, Amazon and Cerrado, and is also found in parts of Bolivia and Paraguay (Vilaça et al. 2024 ). In Brazil, four genetically distinct populations are recognised: the Southern Pantanal (Mato Grosso do Sul), the Northern Pantanal (Mato Grosso), the Northern region (including Pará, Amazonas, Piauí and northern Tocantins), and the Central region (southern Tocantins, Goiás, Bahia and Maranhão) (Almeida et al. 2019 ; Vilaça et al. 2024 ). The Hyacinth Macaw is specialised in feeding on the seeds of palm trees of the genera Attalea and Acrocomia , as well as on the fruits of Mauritia flexuosa Mart., depending on the biome in which it occurs (Guedes 1993 ; Tella et al. 2021). The species also shows specialised reproductive behaviour, nesting primarily in cavities of specific tree species, mainly species of the genus Sterculia (Guedes 1993 ; Silva et al. 2019 ). However, in Central Brazil, it is also capable of nesting in sandstone cliffs (Pacheco and Olmos 2010 ; Dornas et al. 2013 ). Due to its large body size, high intelligence, longevity and vibrant cobalt-blue plumage, the Hyacinth Macaw has become a frequent target of illegal wildlife trade (Guedes 2009 ). These factors, combined with the species’ low reproductive rate and high degree of specialisation in feeding and nesting sites, have led to a marked population decline (Guedes 2009 ). In the 1990s, the estimated wild population comprised only about 2,500 individuals (Guedes et al. 2022 ). Against this background, studies on the biology and reproductive success of the species in the Pantanal, which historically supported the largest population, were fundamental to its conservation. The establishment of the Arara Azul Project in 1990, informed by the knowledge generated by these studies, enabled the implementation of a range of conservation actions (Guedes 2004 ; Kuniy et al. 2006 ). Following these efforts, a substantial increase in the Hyacinth Macaw population was observed, with estimates of approximately 6,500 individuals in the wild (Guedes 2004 ; Guedes 2009 ). In 2014, the species was removed from Brazil’s official list of threatened species and reclassified as ‘Near Threatened’. However, at the global level, it remains categorised as ‘Vulnerable’ by the IUCN (IUCN 2025). Thus, the Arara Azul Project has become a leading reference for the conservation of the Hyacinth Macaw in Brazil, playing a central role in knowledge generation and fostering numerous studies over recent decades. In addition, the project has supported educational initiatives and established partnerships with ranchers, schools and local communities. While many of these studies involved direct or indirect participation by the Arara Azul Institute, others were conducted by independent research institutions. However, the resulting knowledge remains fragmented across the scientific literature, and to our knowledge no study has yet synthesised this information in a comprehensive way, despite its importance for advancing conservation strategies for the species in Brazil. This study aims to synthesise and critically assess the current state of knowledge on the Hyacinth Macaw in Brazil through an integrative review of the scientific literature published over recent decades, with a particular focus on the main challenges to the species’ conservation. Specifically, we aim to: (i) systematise available knowledge to support management actions, research and public policies for conservation; (ii) assess genetic aspects of Hyacinth Macaw populations; (iii) identify the main threats to the species and how they vary across its distribution; (iv) examine interpopulation variation in feeding and reproductive behaviour; and (v) highlight knowledge gaps and challenges for effective conservation across different biomes. 2. METHODS This study is an integrative review conducted following the methodological framework proposed by Whittemore and Knafl ( 2005 ). The review comprised five stages: (1) problem identification, as defined above; (2) literature search; (3) selection of scientific papers; (4) critical assessment and categorisation of the selected studies; and (5) presentation and synthesis of the results, with the final two stages addressed in the Results and Discussion sections of this paper. 2.1 Literature search This review focused exclusively on scientific articles published in indexed journals. This approach was adopted to ensure that the results analysed were derived from methodologically robust studies that are widely recognised by the scientific community. Although dissertations, theses, books, book chapters and conference abstracts may also contain relevant information, articles published in indexed journals were prioritised because they offer greater methodological standardisation, transparency and data traceability, as well as improved comparability across studies. These features are essential for integrative reviews that aim to produce a critical synthesis of knowledge. The literature search was conducted between October 2024 and February 2025 across six databases: ScienceDirect, Web of Science, SpringerLink, Scopus, SciELO and Google Scholar. The following keywords were used: ‘Anodorhynchus hyacinthinus’, ‘Hyacinth Macaw’, ‘Genetics’, ‘Threats’, ‘Ecology’, ‘Conservation’, ‘Conservation of Psittacidae in Brazil’ and ‘Psittacidae behaviour in Brazil’, combined using the Boolean operators ‘AND’ and ‘OR’. To ensure consistency and rigour in the selection process, specific inclusion and exclusion criteria were applied to each database. Only peer-reviewed articles published in indexed journals were considered, including empirical studies, reviews and scientific notes. Studies that focused directly on the Hyacinth Macaw were included, as well as studies addressing ecological components essential to the species, such as plant species used for feeding and nesting, which are fundamental for understanding its ecology and conservation. Duplicate records and publications in languages other than Portuguese or English were excluded. 2.2 Selection of scientific studies All records retrieved from the databases were exported and compiled into a single spreadsheet. Duplicate records across databases were then removed using the duplicate removal tool in Google Sheets, and any remaining duplicates were eliminated manually. Subsequently, the selection process was conducted in three screening phases: (1) title screening, to exclude papers clearly unrelated to the topic; (2) abstract screening, applying the inclusion and exclusion criteria to refine the selection; and (3) full-text screening of the remaining articles to confirm their relevance and eligibility. Only studies that met all criteria following this final assessment were included in the review. 2.3. Search outcomes In total, 5,648 publications were identified, of which 2,951 were duplicates, leaving 2,697 unique records (Fig. 1 ). Of these, 350 were retrieved from ScienceDirect, 489 from SpringerLink, 582 from Web of Science, 251 from Scopus, 25 from SciELO and 1,000 from Google Scholar. During the first screening stage (title screening), 2,522 publications were excluded because they did not focus on the Hyacinth Macaw, leaving 175 articles selected based on their titles. In the second stage (abstract screening), 84 articles were excluded, resulting in 91 studies. Finally, during full-text screening, 20 articles were excluded because they were unrelated to the species, unavailable or not peer-reviewed. Thus, 71 papers were selected for inclusion in this review (see Supporting Information). 3.3 Critical assessment and categorization of the selected studies For each of the 71 studies included in the review, information was compiled and categorised in a spreadsheet, including the title, authors, year of publication, from which the study was retrieved, population studied, biome, animal classification (wild or captive) and study type (review, empirical study or scientific note). The species’ populations were classified following Vilaça et al. ( 2024 ) and divided into Northern and Southern Pantanal, the Central region, and the Northern region of Brazil. In addition, the articles were grouped into five main categories, as shown in Table 1 . Table 1 Themes of the studies selected for the review and their respective aspects. 2 Themes Aspects Ecology Studies on species’ spatial distribution, feeding habits, reproduction, ecological interactions (intra- and interspecific), and population density estimates. Genetics Articles addressing the genetic aspects of the species, including genetic variation within populations. Behaviour Articles published in journals specialized in animal behaviour and/or containing behaviour-related terms in the keywords, title, or abstract. Animal Health Studies addressing health aspects, including clinical, physiological, or sanitary conditions of the species. Conservation Articles focused on the identification of threats, implementation of conservation strategies, population management, and environmental education initiatives aimed at the conservation of the species. a Each study was assigned to a single thematic category according to its primary focus. Based on full-text assessment of the selected studies, the main threats to the species, genetic aspects of its populations, current conservation strategies and initiatives, interpopulation variation in feeding and reproductive behaviour, as well as key knowledge gaps and challenges for effective conservation of the Hyacinth Macaw across different biomes were systematically identified and synthesised. 2.4 Data analysis To analyze the relationship between the number of publications over time, a Poisson regression was applied, with results graphically represented using the ‘ggplot2’ package in R v. 4.2.2. A significance level of p < 0.05 was adopted. A map showing the spatial distribution of studies across Hyacinth Macaw populations in Brazil was produced using QGIS v. 3.28.12. The distribution ranges of each population were obtained from the IUCN Red List, where the species is classified as resident, and Pantanal populations were subdivided following the regional delimitation proposed by Silva and Abdon (1998). Additional graphs summarising the results were also generated using ‘ggplot2’ to support the quantitative analyses. 3. RESULTS AND DISCUSSION Among the 71 articles analysed, most were classified as empirical studies (74%, n = 53), followed by reviews (13%, n = 9) and scientific notes (13%, n = 9) (Fig. 2 ). Research conducted on wild individuals predominated, accounting for 60% (n = 42) of the publications, whereas studies involving captive individuals represented only 14% (n = 10) of the total (Fig. 2 ). In addition, 26% (n = 19) of the selected articles did not focus directly on the species itself, consisting mainly of theoretical reviews or studies on plant species ecologically associated with the Hyacinth Macaw, which provide essential resources for feeding and nesting. Studies based on captive individuals addressed topics such as pathogen occurrence (Barros et al. 2002 ), bacterial resistance to antibiotics (Ramos et al. 2024 ), longevity and sexual maturation (Young et al. 2011 ; Castaldo et al. 2019 ; Barros et al. 2023 ; Barros et al. 2024 ), tool use (Borsari and Ottoni 2005 ; Paim et al. 2008 ), karyotyping (Lunardi et al. 2003 ), bite force (Harrison et al. 2024) and factors influencing species selection for ex situ conservation programmes (Colléony et al. 2017). However, none of these studies directly addressed conservation actions targeting the species, such as reintroduction, restocking or integrated management of captive populations for conservation purposes. The limited number of studies involving captive individuals therefore represents an important limitation for ex situ conservation strategies and, indirectly, for in situ conservation of the Hyacinth Macaw in Brazil. This gap hampers, for example, the development of genetic conservation measures, such as planned reproductive management aimed at maintaining adequate genetic variability within captive populations. Such variability is essential both for the genetic reinforcement of wild populations and for reintroduction programmes in areas where the species has become locally extinct (Vilaça et al. 2024 ). These activities appear to remain a low priority within management strategies for Neotropical psittacids (Berkunsky et al. 2017 ). Based on the articles included in this review, 294 individual authors were identified. Neiva M. R. Guedes, president of the Arara Azul Institute, which coordinates the Arara Azul Project, was the most frequently cited author, appearing in 38% (n = 27) of the publications, followed by Flávia T. Presti (8%, n = 6) and Fernanda M. Fontoura (7%, n = 5). The prominent role of the Institute is also reflected in the temporal pattern of scientific publications on the species. A trend analysis using Poisson regression (Fig. 3 ) revealed a significant increase in the number of publications between 1997 and 2024 (p < 0.05). This sustained growth in scientific output appears to be associated with the long-term research and monitoring activities conducted by the Arara Azul Institute and the dissemination of results generated by its researchers. Overall, the findings of this review highlight the relevance of the Arara Azul Project in consolidating scientific knowledge on the species in Brazil and its importance as one of the main in situ conservation initiatives. The knowledge generated by the project has contributed substantially to understanding the ecological requirements of the Hyacinth Macaw, identifying key threats across its range and supporting evidence-based management strategies aimed at conserving the species in its natural environment. 3.1 Genetic aspects of Hyacinth Macaw populations in Brazil Studies focusing on genetics accounted for approximately 11% (n = 8) of the articles selected in this review (Fig. 3 ). The first genetic studies on Hyacinth Macaw populations in Brazil were conducted by Faria et al. ( 2008 ), who analysed the genetic structure of wild individuals from two regions of the country: Piauí and Mato Grosso do Sul. This pioneering study revealed genetic differences between geographically distant populations and demonstrated the value of genetic approaches for identifying individuals seized from the illegal wildlife trade. Subsequently, Presti et al. ( 2015 ) and Almeida et al. ( 2019 ) expanded these analyses by incorporating samples from previously unstudied regions. Using nuclear and mitochondrial markers, these studies identified genetic differentiation among the currently recognised populations. Both studies also reported relatively low genetic diversity compared to other threatened psittacine species. However, they present methodological limitations that may have influenced these results, such as the use of heterologous markers (Presti et al. 2015 ) and the exclusive use of mitochondrial markers, which do not capture the full genetic variability of the species (Almeida et al. 2019 ). Although informative, these findings represent only a fraction of the species’ total genetic variability, highlighting the need to complement existing data with analyses based on species-specific nuclear markers, including those developed by Silva et al. ( 2015 ), or with genomic approaches. A recent study by Vilaça et al. ( 2024 ), using low-coverage genomic data, produced results broadly consistent with previous studies. However, the authors identified relatively high genetic diversity, particularly in the Southern Pantanal population, as well as evidence of recent gene flow among populations, mainly involving the Northern Pantanal. The high genetic diversity observed in the Southern Pantanal may be associated, at least in part, with gene flow between different regions. The ecological dynamics of the Northern Pantanal provide important insights into the patterns of gene flow reported by Vilaça et al. ( 2024 ). One of the main factors that may explain this pattern is the high emigration rate of individuals, as suggested by Neto, Guedes and Toledo ( 2019 ), who identified the largest known roost of the species in Brazil in this region, comprising approximately one thousand individuals. Such high concentrations may promote dispersal to other areas, particularly in response to the scarcity of nesting sites and food resources, conditions that tend to intensify movement. These movements may influence gene flow and, consequently, genetic variability by enhancing connectivity among populations and partially reducing genetic isolation. Despite these potential movements, the study by Vilaça et al. ( 2024 ) reported high levels of inbreeding, suggesting that the observed gene flow may be recent, limited or insufficient to mitigate the established genetic structure. This highlights the need for continued genetic studies involving representative samples from different parts of the species’ distribution. Such analyses are important for determining whether high levels of inbreeding pose a risk to population viability and whether strategies such as genetic reinforcement would be justified as part of species conservation efforts. 3.2 Ecological aspects of Hyacinth Macaw populations in Brazil Studies related to ecology accounted for 44% (n = 31) of the analysed publications, whereas those focusing on behaviour represented 6% (n = 4), making ecology the most frequently addressed research topic on the species (Fig. 2 ). Most ecological studies (61%, n = 19) were conducted on Pantanal populations. In addition, 13% (n = 4) of these studies did not focus directly on the species itself, consisting mainly of theoretical reviews or studies on plant species ecologically associated with the Hyacinth Macaw, which provide essential resources for feeding and nesting. Analysis of the geographical distribution of studies revealed that most research focused on the Southern Pantanal population, which accounted for 44% (n = 31) of the records, followed by the Northern Pantanal population with 18% (n = 13) of the studies (Fig. 4 ). In addition, 38% (n = 27) of the studies did not address specific populations or focused on plant species ecologically associated with the Hyacinth Macaw. Overall, Pantanal populations emerge as the most extensively studied in Brazil, a pattern that appears to be associated with the presence of the Arara Azul Project in the region, which has enabled continuous, long-term monitoring. Consequently, a substantial portion of the available literature on the ecology and behaviour of the Hyacinth Macaw reflects the sustained research efforts undertaken within the scope of this project. By contrast, this review highlights a marked shortage of information on the ecology and behaviour of the species in the Central and Northern populations of Brazil. This gap limits comprehensive assessments and the formulation of effective, population-specific conservation strategies. The few studies conducted in these regions (n = 4 in the Central population and n = 1 in the Northern population) consisted primarily of faunal inventories which, although valuable, do not provide detailed or systematic information on the species’ ecology. This highlights the need to expand research efforts focused on the Central and Northern populations in order to better understand their specific ecological requirements and local threats. Such efforts would contribute to the development of more effective and targeted conservation strategies for each population. The results of this review indicate that the Pantanal was the most frequently studied biome, representing 48% (n = 34) of the publications, followed by the Cerrado (20%, n = 14) and the Amazon (11%, n = 11) (Fig. 5 ). Notably, 38% of the analysed papers addressed the Neotropical region more broadly. When considering only studies focused on Brazilian populations, the Pantanal holds the largest volume of available information, making it central to current understanding of the species’ ecology and conservation in Brazil. In contrast, populations in the Central region, located in the Cerrado, and in the Northern region of the country, located in the Amazon, remain poorly studied. These knowledge gaps could be reduced through increased sampling effort and targeted research in these regions, aimed at better understanding the species’ ecological requirements and ensuring population viability across its range. Citizen science initiatives represent a promising approach to expand knowledge of the Hyacinth Macaw by involving local communities, guides, birdwatchers and partner institutions in the collection of ecological and occurrence data. Such collaborative efforts can help address monitoring gaps and strengthen participatory conservation strategies. 3.2.1 Diet and Feeding Ecology The studies analysed in this review indicate that the diet of Hyacinth Macaws in the Southern and Northern Pantanal populations consists largely of the seeds of two palm species that are widely distributed in the region: Attalea phalerata Mart. ex Spreng and Acrocomia aculeata (Jacq.) Lodd. ex Mart. (Tella et al. 2020 ; Guedes et al. 2022 ). A. phalerata is the most frequently consumed food item in the Pantanal, a pattern that can be explained by a combination of traits of this palm species, including its high abundance, continuous fruiting throughout the year, the high lipid content of its seeds, which is among the highest within the Arecaceae, and low competition pressure, as relatively few species feed on its seeds (Negrelle 2015 ). The Hyacinth Macaw has morphological specializations that allow it to handle the A. phalerata fruits, including adaptations of the skull, beak, and tongue, which result in one of the strongest bite forces among Psittacidae (Blanco et al. 2017 ; Harrison et al. 2024). Although A. phalerata fruits have a very hard endocarp, individuals from these populations tend to select fruits with thinner endocarps (Barros and Pires 2021 ). This preference may be interpreted as an evolutionary response to the structural defences of the plant, likely arising from a coevolutionary relationship between the two species. In addition to morphological and evolutionary adaptations, some behavioral strategies related to A. phalerata consumption have also been documented. Schneider et al. ( 2006 ) observed that some macaws remove pieces of A. phalerata leaves to facilitate fruit handling and make it easier to crack the endocarp. The authors also reported that macaws frequently drop fruits while manipulating the bunches and later return to the site, approximately one month later, to feed on the seeds of the fallen fruits, which contained insect larvae that were likely consumed. A similar behaviour was reported by Paula et al. (2015), who observed a pair of Hyacinth Macaws consuming termites from pieces of wood from a nest infested by these insects in a transition area between the Southern Pantanal and the Cerrado. This behaviour may be interpreted as a form of protein supplementation, particularly during the breeding season, when protein demands are higher. Based on the literature analysed, some studies report that in the Pantanal, particularly in cattle ranching areas, groups of Hyacinth Macaws are commonly observed foraging on the ground in search of A. phalerata fruits (Schneider et al. 2006 ; Tella et al. 2020 ). Both cattle and Tapirus terrestris Linnaeus, 1758 consume the mesocarp of A. phalerata fruits and subsequently regurgitate or excrete the endocarps, thereby facilitating access to the seeds by macaws (Schneider et al. 2006 ; Barros and Pires 2021 ). Regarding the diet of the Northern population, no studies were found that directly investigated the diet of the Hyacinth Macaw in this region or the foraging behaviours associated with it. Available information is limited to indirect evidence reported in isolated studies, in which groups of individuals were observed in Mauritia palm swamps and in ecosystems with a high abundance of palms of the genus Attalea , particularly Attalea maripa (Aubl.) Mart. (Pacheco and Olmos 2005 ; Dornas et al. 2013 ). These observations suggest that these palm species may represent the main food resources for the population in this region. For the Central population, the limited available data indicate that Mauritia flexuosa is likely to represent a primary food resource for the Hyacinth Macaw in this region. This palm can occur dominantly in certain Cerrado areas, substantially contributing to local food availability (Dornas et al. 2013 ; Tella et al. 2020 ). In addition to M. flexuosa , other palms used as food resources in this region include Attalea eichleri (Drude) A. J. Hend. and Attalea barreirensis Glassman (Pacheco and Olmos 2010 ; Tella et al. 2020 ). Unlike what is observed in other regions, such as the Pantanal, where the species predominantly consumes palm seeds, Tella et al. ( 2020 ) reported that individuals from this population feed mainly on the pericarp of M. flexuosa fruits, thereby playing a significant role in the dispersal of this species’ seeds. Although initially considered a purely antagonistic relationship, the interaction between the Hyacinth Macaw and the palm species it feeds on in different regions represents a continuum between antagonism and mutualism (Blanco et al. 2017 ; Dracxler and Kissling 2022 ). Consequently, the species can act as an important seed disperser, as fruits are often removed directly from palms and transported to safer locations for consumption, where fruits or seeds may be dropped. This process can significantly contribute to the colonisation and persistence of these palm species across the landscape (Blanco et al. 2017 ; Tella et al. 2020 ). Thus, the Hyacinth Macaw also performs an ecological role analogous to that once played by members of the extinct megafauna, since few current species are able to carry out this interaction. Therefore, future studies should investigate these ecological relationships in greater depth, assessing the impact of the absence of the Hyacinth Macaw on the demography and population dynamics of Attalea species and other palms consumed by the macaw. 3.2.2 Reproductive Biology In addition to its highly specialised diet, the Hyacinth Macaw also exhibits strong specialisation in the selection of nesting sites. In the Pantanal, the species primarily nests in natural cavities of Sterculia apetala (Jacq.) H. Karst., a tree species typical of this biome. According to Guedes et al. ( 2022 ), approximately 95% of natural nests in the Southern Pantanal are found exclusively in S. apetala trees, whereas in the Northern Pantanal this proportion is slightly lower, at around 85% (Pinho and Nogueira 2003 ). S. apetala exhibits a set of physical and chemical traits that favour its selection as a nesting site, including soft wood, rapid growth, and the frequent presence of natural cavities suitable for the reproduction of several Pantanal species. Furthermore, this species contains chemical compounds with fungicidal properties that inhibit pathogen proliferation within nests, thereby contributing to increased nestling survival (Fontoura et al. 2015 ). Natural cavities formed in S. apetala trees function as important nesting sites, being used not only by the Hyacinth Macaw but also by several other wildlife species in the Pantanal (Carrara et al. 2007 ). However, for these cavities to be suitable for Hyacinth Macaws, trees must be at least 60 years old (Júnior et al. 2006 ). Despite this, it is estimated that only about 5% of adult S. apetala trees in the Southern Pantanal contain cavities with characteristics suitable for the species, making this a scarce and highly contested resource during the breeding season (Johnson et al. 1997 ; Júnior et al. 2007 ). Future studies could therefore focus on constructing interaction networks centred on cavity availability in S. apetala , aiming to better understand the relationships between the Hyacinth Macaw and other cavity-dependent species, as well as to characterise patterns of agonistic interactions and degrees of dependence among Pantanal fauna. Such an approach would allow a more precise assessment of the relative dependence of different species on S. apetala , which is recognised as a keystone species in Pantanal ecosystems. In addition, the application of population dynamics models for S. apetala could help project the future availability of nesting cavities in key areas of the species’ range, thereby providing valuable support for conservation planning. Regarding the Northern population, among the few studies addressing reproduction, the work of Silva et al. ( 2019 ) stands out. While investigating the species’ ecology within a mosaic of conservation units in Carajás, southeastern Pará, the authors observed that more than 80% of the nests used by this population were located in natural cavities of Sterculia pruriens (Aubl.) K. Schum. In addition, most nests were found in open environments (87.5%), followed by forest interiors (8.3%) and forest edges (4.2%). With respect to reproduction in the Central population, the available information is largely based on field observations and is therefore inconclusive (Pacheco and Olmos 2010 ; Dornas et al. 2013 ). Reports indicate nesting on sandstone cliffs in eastern Tocantins, particularly in the Jalapão region, as well as possible nesting in cavities of Mauritia flexuosa palms (Pacheco and Olmos 2010 ; Rego et al. 2011 ). However, no study specifically addressing the reproductive biology of the species in this region was included in this review, representing a significant gap in understanding the factors influencing local reproductive success. The potential use of sandstone cliffs as nesting sites suggests that the Hyacinth Macaw population in this region may exhibit ecological adaptations distinct from those of the Pantanal and Amazon populations. Such differentiation may be associated with local factors, including a low availability of natural tree cavities, reduced predation pressure leading to higher nestling survival, or lower interspecific competition for nesting sites. Nonetheless, these hypotheses remain untested, highlighting the need for targeted studies on the species’ ecology in the Central region to generate systematic data and enable comparisons with other populations. According to available data from studies conducted in the Pantanal, the Hyacinth Macaw exhibits relatively low reproductive success. Females may lay between one and three eggs, with an average of two eggs per pair (Ramalho et al. 2024 ). However, of every 100 eggs laid, only approximately 25% hatch and survive to the age at which nestlings typically leave the nest (Guedes et al. 2022 ). Egg laying in this species is asynchronous, meaning that after the first egg is laid, there may be an interval of 1 to 16 days before the second egg is laid. This strategy is thought to reduce the effects of predation and increase the likelihood that at least one nestling survives (Kuniy et al. 2006 ; Ramalho et al. 2024 ). Nevertheless, when both eggs hatch, the second nestling is often rejected by the parents. There are, however, records of pairs successfully raising two nestlings, depending on factors such as parental experience, food availability, and the age difference between chicks (Guedes et al. 2022 ). During incubation, the female remains inside the nest for most of the time, where she protects and incubates the eggs. The male, in turn, acts as a sentinel, defending the nest against predators and competitors, and is responsible for foraging and feeding the female (Pinho and Nogueira 2003 ; Schneider et al. 2006 ). An additional noteworthy aspect is that slight sexual dimorphism is observable only at the beginning of the breeding season, when females may exhibit a slightly curved tail as a result of prolonged periods spent incubating the eggs inside the nest (Schneider et al. 2006 ). Egg predation represents a significant threat to reproductive success, with Ramphastos toco Statius Muller, 1776 accounting for approximately 53% of predation events recorded in the Southern Pantanal (Pizo et al. 2008 ). Paradoxically, R. toco is also the most important seed disperser of Sterculia apetala , being responsible for about 86% of the species’ seed dispersal in the Pantanal (Pizo et al. 2008 ). This dispersal plays a crucial role in the regeneration and maintenance of trees that develop natural cavities suitable for nesting. The incubation period of Hyacinth Macaw eggs lasts on average 28 to 30 days. During the final stages of incubation, the embryo begins to vocalise inside the egg, which likely serves as a stimulus for parental assistance with thermoregulation and the hatching process (Ramalho et al. 2024 ). After hatching, nestling development occurs in three stages, with fledging taking place at an average age of 107 days. However, juveniles remain under parental care for up to 12 months, a period that is essential for survival in the wild (Guedes et al. 2022 ). In the Northern population, nests containing nestlings at different developmental stages have been reported, indicating asynchrony in the reproductive cycle among breeding pairs. This pattern contrasts with that observed in Pantanal populations, where greater synchrony among pairs has been documented (Silva et al. 2019 ). Moreover, some individuals exhibit developmental abnormalities, characterised by body mass and overall weight gain that are approximately 54% lower than those of typically sized individuals, and are therefore referred to as “dwarfs” (Guedes et al. 2022 ). Given that these individuals represent a small fraction of the population and that the ecological and reproductive implications of this condition remain poorly understood, targeted studies are needed to assess whether it affects individual fitness. 3.3 Threats and Challenges to the Conservation of the Hyacinth Macaw in Brazil Among studies addressing threats to the Hyacinth Macaw in Brazil, Devenish et al. ( 2021 ) reported an expansion of the species’ range between 1990 and 2019. However, approximately 30% of this area was converted into pastures and agricultural land, and only 8% is currently located within Protected Areas, raising concerns about the effectiveness of existing conservation measures. In addition, previous population estimates indicating around 6,500 individuals in the wild (Guedes 2004 ) reflect a past demographic scenario. Therefore, updated population surveys are required to provide more accurate estimates of the species’ current status. Such surveys could be conducted through collaborations between South American non-governmental organizations and Citizen Science initiatives, promoting the involvement of local communities in data collection, expanding monitoring efforts, and strengthening participatory conservation. Despite significant advances in knowledge about the species, the available data still reveal important knowledge gaps, reflecting broader spatial and thematic biases reported for biodiversity research in the Pantanal (Frota et al. 2020 ). The Central region of the Pantanal, for instance, remains poorly sampled and studied. In this review, only one article related to this area was identified, reporting a case of cavity competition between a pair of Hyacinth Macaws and Tyto furcata (Temminck, 1827), which suggests a possible scarcity of suitable natural cavities (Tortato and Bonanomi 2012 ). Given this scenario, it is essential that future studies conduct surveys not only in areas already known to host the Hyacinth Macaw but also in poorly sampled or still unexplored regions, such as the municipality of Parintins in the state of Amazonas (Barreiros and Gomes 2010 ), as well as in the Central population, which currently lacks studies focused on the species’ biology and ecology. Such surveys are crucial to obtain more accurate estimates of wild population size, identify priority areas for conservation, recognize region-specific threats, and better understand the ecological requirements of the species across different population contexts. This information is also fundamental to support the creation of new Protected Areas and the management of potentially isolated populations within the landscape. In this sense, the Hyacinth Macaw represents a symbol of Brazilian biodiversity, and its conservation is essential not only for future generations to understand the importance of the species but also for the preservation of entire ecosystems on which many other species depend (Douglas and Veríssimo 2013 ). As proposed by Berkunsky et al. ( 2017 ) and Vilaça et al. ( 2024 ), a conservation approach based on population management units is recommended for the Hyacinth Macaw. This strategy allows the identification and mitigation of local threats, the containment of potential population declines, and the preservation of the species’ genetic variability. Each biome in which the Hyacinth Macaw occurs is subject to distinct anthropogenic pressures and specific socio-environmental contexts, which require the development of differentiated management strategies and the definition of priority areas for monitoring. In the following subsections, the main threats and challenges faced by the species’ populations in the Pantanal, Amazon and Cerrado biomes will be addressed, highlighting the importance of conservation actions adapted to regional specificities. 3.3.1 Pantanal As previously noted in this review, the Pantanal stands out as the most important biome for the conservation of the Hyacinth Macaw in Brazil. Complementary data from the literature, such as those presented by Oliveira et al. ( 2021 a), indicate that only 8.8% of the occurrence areas of the three key species, the Hyacinth Macaw, S. apetala , and A. phalerata , are located within conservation units in the biome. Moreover, these authors reported a 13% loss of these occurrence areas between 2002 and 2017, mainly attributed to deforestation and the expansion of agricultural activities. Due to the characteristic flood regime of the Pantanal, agricultural activities are concentrated in ecosystems not subject to flooding, such as capões and cordilheiras , which largely overlap with the occurrence areas of plant species essential to the Hyacinth Macaw’s life cycle (Oliveira et al. 2021 b). Cattle pressure in these areas directly affects the density of S. apetala seedlings and compromises the regeneration of A. phalerata , mainly through soil compaction that inhibits seed emergence (Johnson et al. 1997 ; Tella et al. 2020 ). In addition, these regions have been deforested and converted into pastures or monoculture crops, reducing the availability of breeding and feeding sites for the species. The data analysed in this review and in the literature indicate the need to establish new conservation units in non-flooded areas, particularly in strategic regions that promote connectivity between the Northern and Southern Pantanal populations, such as the Central Pantanal. Another essential measure is the development of public policies and specific legislation that restrict, discourage, and penalise deforestation in capões and cordilheiras , especially in the southern portion of the Pantanal. Regulations aimed at controlling logging and vegetation suppression in these areas can strengthen oversight and environmental licensing, thereby preventing irregular and illegal activities. In addition to deforestation, fire represents one of the greatest threats to Hyacinth Macaw populations in the Pantanal, as it directly affects key resources used for nesting and feeding. According to Ferreira et al. (2021), most of the distribution of the Hyacinth Macaw, Sterculia apetala and Attalea phalerata historically occurs in areas that have been little affected by fire. However, due to climate change and the conversion of native vegetation into pastures, fires have become more intense and widespread, driven by increasingly frequent and severe droughts in recent decades. In 2020, approximately 28% of the areas considered suitable for the Hyacinth Macaw were affected by wildfires, as well as 25% of the areas suitable for S. apetala and A. phalerata . These values were about five times higher than the annual average recorded between 2003 and 2019 (Ferreira et al. 2021). Fires directly compromise food availability, as the fruits of A. phalerata and A. aculeata become unsuitable for consumption after burning. In addition, fire negatively affects reproductive success, since the peak of fire occurrence in the Pantanal, in September, coincides with the peak of the species’ breeding season. Natural cavities used as nests may be destroyed or structurally compromised during fires, rendering them unusable in subsequent breeding seasons and further reducing the availability of suitable nesting sites. Among the effective mitigation strategies, integrated fire management stands out, including preventive actions such as environmental education campaigns, coordinated and structured firefighting efforts, and the controlled use of fire in areas with high loads of dry combustible material, particularly around capões and cordilheiras . Complementary to these measures, geotechnologies such as Remotely Piloted Aircraft Systems (RPAS) and Geographic Information Systems (GIS) have proven to be essential tools. These technologies enable spatial monitoring of risk areas, integration of environmental variables, and the production of easily interpretable information to support decision-making. Their effectiveness in conservation actions in the Pantanal has already been demonstrated (Guedes et al. 2006 ). The use of aerial imagery obtained from RPAS or satellite platforms, combined with data on relative humidity and temperature, allows the identification of areas with high fire risk and the continuous monitoring of environmental conditions, which are fundamental measures for the protection of terrestrial ecosystems and the conservation of the Hyacinth Macaw. Another emerging threat to the Hyacinth Macaw is environmental contamination associated with agricultural activities. Studies conducted by Marchesi et al. ( 2015 ) identified high cadmium concentrations in nestlings from the Southern Pantanal, indicating exposure to heavy metal contamination. In the same region, Vicente and Guedes ( 2021 ) also reported mortality events among individuals of the species resulting from the improper use of pesticides. Although these occurrences appear to be sporadic, they are alarming and highlight the need for systematic monitoring in areas under intensive agricultural use, in order to assess whether irregular applications of agrochemicals may compromise reproductive success. In addition, environmental changes, whether natural or anthropogenic, can promote the emergence of pathogens in nestlings, weakening their immune system and facilitating infections by opportunistic yeasts and bacteria (Loiko et al. 2007 ; Allgayer et al. 2009 b), as well as by free-living microorganisms such as Chlamydophila psittaci (Raso et al. 2006 ; Raso et al. 2013 ) and Salmonella spp . (Allgayer et al. 2009 a), which are capable of causing severe diseases in wild populations. Given the multiple threat factors identified, the results of this review highlight the need for integrated conservation actions for Hyacinth Macaw populations in the Pantanal. Key priorities include strengthening public policies for the protection of capões and cordilheiras , expanding protected areas in strategic regions, implementing integrated fire management supported by geotechnologies, monitoring environmental contamination, and assessing population health, all of which are essential to ensure the long-term viability of the species. 3.3.2 Amazon According to the results presented above, the Amazon biome and the Hyacinth Macaw population occurring in this region remain the least studied compared to the other biomes. This scarcity of scientific information limits the understanding of local threats and constrains the development of targeted conservation strategies. Among the few studies addressing the Amazon biome, Devenish et al. ( 2021 ) reported an apparent geographic expansion of the Hyacinth Macaw in the region. This expansion has been directly associated with landscape changes driven by deforestation, particularly the conversion of forested areas into savanna-like environments. Although such changes may initially favour the species, which in this region shows a preference for nesting in more open habitats, continued conversion of native vegetation into pastureland is likely to reverse this trend, leading to habitat loss and a reduction in the species’ area of occupancy. Therefore, the apparent expansion of the Hyacinth Macaw’s range in the Amazon should not be interpreted as a positive conservation signal, but rather as a consequence of ongoing land-use change and deforestation in the biome. Thus, further studies focusing on the demography of the Hyacinth Macaw in the Amazon are needed to estimate current population size and to support the development of effective strategies aimed at ensuring the species’ long-term viability in the region. In addition, awareness-raising initiatives involving local communities are essential, particularly those that highlight the species’ vulnerability and its ecological and economic relevance. A successful example is provided by Presti et al. ( 2017 ), who implemented environmental education activities in schools in the Carajás region, Pará, using educational games to promote student engagement in Hyacinth Macaw conservation. The scarcity of studies on the Amazonian population may be partly explained by the difficulty of access to these areas, as well as by the high logistical and financial costs involved, which pose major challenges for research. Nevertheless, the development of new studies in the region is essential to improve understanding of the species’ ecological requirements and the local threats faced by this population. It is noteworthy that the Arara Azul Institute is currently conducting continuous monitoring in the region, which is expected to substantially advance knowledge of the Amazonian population. Even so, the results of this review indicate that the Amazon biome still lacks fundamental information, underscoring the urgent need to expand research and conservation efforts to ensure the long-term persistence of the Hyacinth Macaw and the conservation of this biome in Brazil. 3.3.3 Cerrado According to the results of this review, the population in the Central region, within the Cerrado biome, has been the subject of relatively few studies, which limits the availability of information on the threats faced by this population. Among the studies addressing the Cerrado, Borges et al. ( 2019 ) stand out for highlighting the high vulnerability of the Hyacinth Macaw to environmental changes in this biome. These authors reported that only 14% of the species’ range in the Cerrado is located within Protected Areas, exposing the population to the expansion of the agricultural frontier and increasing habitat fragmentation. Under these conditions, actions such as systematic monitoring of the Central population, the establishment of new Protected Areas, and the implementation of ecological corridors are essential to facilitate individual movement and to mitigate the effects of land-use change and projected climate shifts during this century. Although the expansion of the agricultural frontier represents one of the main threats to the Central population through the loss of feeding and nesting areas, illegal wildlife trade also constitutes a major driver of population decline. According to Faria et al. ( 2008 ) and Presti et al. ( 2015 ), most Hyacinth Macaws seized during anti-trafficking operations originated from this population. This vulnerability is closely associated with the socioeconomic context of northeastern Brazil, which presents some of the lowest human development indices in the country. In areas characterized by low income and limited access to information, wildlife trade is often perceived as an economic alternative rather than as an illegal activity, being viewed as a means of exploiting what is considered an abundant natural resource (Herrera and Hennessey 2007 ; Pires 2012 ; Clarke and By 2013 ). In addition, the predatory methods used to capture eggs and nestlings further exacerbate conservation problems, as hunters frequently cut down nesting trees, thereby reducing the availability of natural cavities (Berkunsky et al. 2017 ; Bonaparte et al. 2024). Thus, the conservation measures adopted by the Arara Azul Project in the Pantanal and the Amazon, such as raising awareness among local communities about the species’ importance and vulnerability, establishing partnerships with landowners to protect nesting sites, and promoting tourism as a source of income for rural families, could also be implemented for this population. These actions have the potential to reduce illegal trade and to enhance local engagement in the conservation of the Hyacinth Macaw (Corrêa and Guedes 2006 ; Mercado et al. 2021 ). Another emerging factor that deserves attention is the use of social media platforms for illegal wildlife trade, which has expanded across several countries in the Americas. These platforms facilitate illegal transactions and may partially replace physical markets following their closure, allowing wildlife trafficking to persist within national borders (Pires et al. 2015 ; Picazo et al. 2023 ). However, studies in Brazil examining the role of social media in Hyacinth Macaw trafficking remain scarce, representing an important gap for future research. Lastly, it is important to note that although the Hyacinth Macaw has been removed from Brazil’s official list of threatened species, this reclassification may result in less stringent enforcement and weaker penalties for illegal trade. Such a scenario could increase harvesting pressure, particularly in regions with limited monitoring and high socioeconomic vulnerability. Accordingly, the results of this review highlight the need for further studies focused on the Central population, with particular emphasis on socio-environmental threats, feeding and reproductive ecology, and local community perceptions of the species. Integrated approaches that consider the socio-ecological systems of the region are essential to develop effective conservation strategies, reduce illegal trade, and ensure the long-term viability of the Hyacinth Macaw in the Cerrado. 3.4 Limitations of the review Although this review was carefully developed, some limitations should be acknowledged. First, the distribution of available information on Hyacinth Macaw populations in Brazil remains uneven, with a particularly low number of studies focusing on populations in the Central and Northern regions of the country. This scarcity of information limits the understanding of ecological requirements and regional threats, thereby constraining the development of conservation strategies aimed at ensuring the species’ long-term viability. In addition, this review focused exclusively on peer-reviewed articles published in indexed journals, a criterion adopted to ensure methodological rigour and comparability among the studies analysed. Consequently, relevant information contained in theses, dissertations, book chapters, technical reports, conference abstracts, and citizen-science databases was not included, although such sources may provide valuable insights into the species. Acknowledging these limitations is essential to guide future research and to support more comprehensive and effective conservation strategies. 4. CONCLUSIONS This study represents one of the first reviews dedicated exclusively to the Hyacinth Macaw in Brazil, integrating information on its biology, distribution, and conservation, and providing a valuable foundation to guide future conservation strategies for the species. The results indicate a marked increase in the number of studies on the Hyacinth Macaw in Brazil between 1997 and 2024, particularly in the fields of ecology, conservation, and animal health. However, this scientific production has been largely concentrated in the Pantanal biome, especially in the Southern Pantanal, where long-term monitoring and management actions have been conducted by the Arara Azul Project. These efforts have substantially advanced knowledge of the species’ threats, reproductive biology, genetics, and ecology, and have contributed to the observed recovery of Hyacinth Macaw populations in the country. Furthermore, this review identified regional variation in threats, feeding and reproductive behaviour, and ecological interactions, indicating that conservation strategies should be tailored to the specific conditions of each biome. Despite the overall increase in scientific output, substantial knowledge gaps remain, particularly for populations in the Northern and Central regions of Brazil, where information on reproductive biology, local ecology, and population-specific threats is still limited. In addition, the small number of studies involving captive individuals represents a major constraint for the development of future reintroduction or restocking programmes in areas strongly affected by anthropogenic pressures. Consequently, it is essential to direct future research efforts toward these regions and to promote studies focused on the ex situ conservation of the species. Overall, this review provides a consolidated framework to support future research, management actions, and the formulation of public policies aimed at conserving the Hyacinth Macaw in Brazil. By fostering effective and regionally adapted strategies that actively engage local communities through citizen science initiatives and environmental education, it is possible to mitigate the environmental threats that continue to compromise the integrity of the species’ populations. Declarations CONFLICT OF INTEREST STATEMENT The authors have no competing interests. ACKNOWLEDGEMENTS We are grateful to the members of the Hyacinth Macaw Institute for their generous assistance throughout this study, particularly Fernanda M. Fontoura, who provided invaluable insights. We also thank Alexandra Sanches and Eduardo Roberto Alexandrino for their careful evaluation of the manuscript and for their valuable comments and suggestions, which were essential to the development of this article. FUNDING STATEMENT This research did not receive any specific grant from funding agencies in the public, commercial, or not-for-profit sectors. AUTHOR CONTRIBUTIONS Both authors contributed to the study conception and design. Material preparation, data collection and analysis were performed by Vitor Matheus Morais de Oliveira. The first draft of the manuscript was written by Vitor Matheus Morais de Oliveira, and Vinicíus de Avelar São Pedro reviewed the manuscript and contributed to the development of the study structure. 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Biodivers Conserv 30:1971–1990. https://doi.org/10.1007/s10531-021-02176-w Pacheco JF, Olmos F (2005) Birds Of A Latitudinal Transect In The Tapajós-Xingu Interfluvium, Eastern Brazilian Amazonia. Ararajuba 13:29–46 Pacheco JF, Olmos F (2006) Birds Of Tocantins 1. Southeast Region. Revista Brasileira De Ornitologia 14:85–100 Pacheco JF, Olmos F (2010) Birds Of Tocantins, Brazil – 2: Jalapão Region. Revista Brasileira De Ornitologia 18:1–18 Paim CS, Borsari A, Ottoni EB (2008) Means To An End: Neotropical Parrots Manage To Pull Strings To Meet Their Goals. Anim Cogn 13:287–301. https://doi.org/10.1007/s10071-008-0190-z Paula GA, Laps R, Fischer E (2017) Hyacinth Macaws ( Anodorhynchus Hyacinthinus , Psittacidae) Feeding On Termites. Ornitologia Neotropical 28:187–190. https://doi.org/10.58843/ornneo.v28i0.247 Picazo RIS, Bravo OER, Padilha IM, Rivera EEC (2023) The Role Of Social Media Groups On Illegal Wildlife Trade In Four Mexican States: A Year-Long Assessment. Global Ecol Conserv 45. https://doi.org/10.1016/j.gecco.2023.e02539 Pinho JB, Nogueira FMB (2003) Hyacinth Macaw ( Anodorhynchus Hyacinthinus ) Reproduction In The Northern Pantanal, Mato Grosso. Brazil Ornitologia Neotropical 14:29–38 Pires SF (2012) The Illegal Parrot Trade: A Literature Review. Global Crime 13:1–15. https://doi.org/10.1080/17440572.2012.700180 Pires SF, Schneider JL, Herrera M (2015) Organized Crime Or Crime That Is Organized? The Parrot Trade In The Neotropics. Trends Organized Crime 19:4–20. https://doi.org/10.1007/s12117-015-9259-7 Pizo MA, Donatti CI, Guedes NMR, Galetti M (2008) Conservation Puzzle: Endangered Hyacinth Macaw Depends On Its Nest Predator For Reproduction. Biol Conserv 141:792–796. https://doi.org/10.1016/j.biocon.2007.12.023 Presti FT, Almeida TRA, Silva GF, Silva HE, Conrado LP, Cespede L, Rdrigues TM, Barbirato M, Wasko AP (2017) Conhecendo A Arara-Azul-Grande: Confecção E Aplicação De Um Jogo Didático Como Parte Das Ações De Educação Ambiental Visando A Conservação Da Espécie. Revista Brasileira De Educação Ambiental 12:259–273. https://doi.org/10.34024/revbea.2017.v12.1982 Presti FT, Guedes NMR, Antas PTZ, Miyaki CY (2015) Population Genetic Structure In Hyacinth Macaws ( Anodorhynchus Hyacinthinus ) And Identification Of The Probable Origin Of Confiscated Individuals. J Hered 106:491–502. https://doi.org/10.1093/jhered/esv038 Ramalho KRA, Fontoura FM, Guedes NMR (2024) First Record Of Free-Living Hyacinth Macaw ( Anodorhynchus Hyacinthinus ) Eggs Hatching Using Camera Traps In Southern Pantanal, Brazil. Ornithol Res 33. https://doi.org/10.1007/s43388-024-00207-y Ramos CA, Ferreira CJ, Ballaben AS, Filho RACP, Darini ALC (2024) Analysis Of Antibiotic Resistance In Gram-Negative Bacilli In Wild And Exotic Healthy Birds In Brazil: A Warning Sign. Veterinary Microbiology 296. https://doi.org/10.1016/j.vetmic.2024.110196 Raso TF, Seixas GHF, Guedes NMR, Pinto AA (2006) Chlamydophila Psittaci In Free-Living Blue-Fronted Amazon Parrots ( Amazona Aestiva ) And Hyacinth Macaws ( Anodorhynchus Hyacinthinus ). Brazil Veterinary Microbiol 117:235–241. https://doi.org/10.1016/j.vetmic.2006.06.025 . The Pantanal Of Mato Grosso Do Sul Raso TF, Teixeira RHF, Carrasco AOT, Júnior JPA, Pinto AA (2013) Chlamydophila Psittaci Infections In Hyacinth Macaws ( Anodorhynchus Hyacinthinus ) Confiscated In Brazil. J Zoo Wildl Med 44:169–172. https://doi.org/10.1638/1042-7260-44.1.169 Rego MA, Silveira LF, Piacentini VQ, Schunck F, Machado E, Pinheiro RT, Reis E (2011) As Aves Da Estação Ecológica Serra Geral Do Tocantins, Centro Do Brasil. https://doi.org/10.1590/S1676-06032011000100027 . Biota Neotropical 11 Santos MPD, Santana A, Soares LMS, Sousa SA (2012) Avifauna of Serra Vermelha, Southern Piaui, Brazil. Revista Brasileira De Ornitologia 20:199–214 Schneider L, Serbena AL, Guedes NMR (2006) Behavioral Categories Of Hyacinth Macaws ( Anodorhynchus Hyacinthinus ) During The Reproductive Period, At South Pantanal, Brazil. Revista De Etologia 8:71–80 Silva GF, Presti FT, Rechetelo J, Guedes NMR, Wasko AP, Donatelli RJ (2019) Hyacinth Macaw ( Anodorhynchus Hyacinthinus ) Nests In A Mosaic Of Protected Areas In Carajás And Surrounding Areas, State Of Pará, Brazil. Revista Brasileira de Ornitologia 27:187–194. https://doi.org/10.1007/BF03544469 Silva HE, Presti FT, Wasko AP, Pinhal D (2015) Development Of Microsatellite Markers For Hyacinth Macaw ( Anodorhynchus Hyacinthinus ) And Their Cross-Amplification In Other Parrot Species Genetics. Bmc Research Notes 8. https://doi.org/10.1186/s13104-015-1749-9 Tella JL, Hiraldo F, Pacifico E, Luque JAD, Dénes FV, Fontoura FM, Guedes NMR, Blanco G (2020) Conserving The Diversity Of Ecological Interactions: The Role Of Two Threatened Macaw Species As Legitimate Dispersers Of Megafaunal Fruits. Diversity 12. https://doi.org/10.3390/d12020045 Tortato FR, Bonanomi J (2012) Competition For Cavity Between Anodorhynchus Hyacinthinus (Latham, 1790) (Psittacidae) And Tyto Alba (Scopoli, 1769) (Tytonidae) In The Pantanal Of Paiaguas Region, Corumba, Mato Grosso Do Sul, Brazil. Revista Brasileira de Ornitologia 20:22–25 Vicente EC, Guedes NMR (2021) Organophosphate Poisoning of Hyacinth Macaws In The Southern Pantanal, Brazil. Scientific Reports 11. https://doi.org/10.1038/s41598-021-84228-3 Vilaça ST, Dalapicolla J, Soares R, Guedes NMR, Miyaki CY, Aleixo A (2024) Prioritizing Conservation Areas For The Hyacinth Macaw ( Anodorhynchus Hyacinthinus ) In Brazil From Low-Coverage Genomic Data. Evolutionary Applications 17. https://doi.org/10.1111/eva.70039 Whittemore R, Knafl K (2005) The Integrative Review: Updated Methodology. J Adv Nurs 52:546–553. https://doi.org/10.1111/j.1365-2648.2005.03621.x Young AM, Hobson A, Lackey LB, Wright TF (2011) Survival On The Ark: Life-History Trends. Captive Parrots Anim Conserv 15:28–43. https://doi.org/10.1111/j.1469-1795.2011.00477.x Additional Declarations No competing interests reported. Supplementary Files DatabaseSpreadsheetMainDocument.xlsx SUPPLEMENTARY MATERIAL Supplementary material supporting this article is available online at the end of the article. Cite Share Download PDF Status: Under Review Version 1 posted Reviewers invited by journal 18 Mar, 2026 Editor assigned by journal 06 Feb, 2026 Submission checks completed at journal 06 Feb, 2026 First submitted to journal 25 Jan, 2026 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-8695459","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Systematic Review","associatedPublications":[],"authors":[{"id":608144580,"identity":"7e38f261-e175-4b75-b87f-58e074163ae9","order_by":0,"name":"Vitor Matheus Morais de Oliveira","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA+ElEQVRIie3PMWrDMBSAYRmDsii7IL1CISYgOhj7IF0kDMrUvUMGZ5Gn7LlE184Ggb2IenXJItMLuFsGU/psugXbyVaI/kUI3seTEHK5/mscIQKHZ9vXsD/3+VUEjB8cjexJOk/6gODVUunhMkkes0PT2k4/xIvDV3DEVfSWadiyC5/HCDPlhgqlCSFlYFtySt6NAFLIl3SM1BJRkUpCqIS/0FPCciBeqqeIf+bdQPCKrD8SVjWzBFOOwz/C84jVc1tMgZ+EAmIKeFiecFbDFj71l1L5n+eOxotMefb7J4pZtW1suwtHyUVimOTXjvfFtwy7XC7XffQL6bdhUlh/1EsAAAAASUVORK5CYII=","orcid":"","institution":"Federal University of São Carlos","correspondingAuthor":true,"prefix":"","firstName":"Vitor","middleName":"Matheus Morais","lastName":"de Oliveira","suffix":""},{"id":608144581,"identity":"9d38e473-794a-4899-a84e-5558f1f752ac","order_by":1,"name":"Vinicíus de Avelar São Pedro","email":"","orcid":"","institution":"Federal University of São Carlos","correspondingAuthor":false,"prefix":"","firstName":"Vinicíus","middleName":"de Avelar São","lastName":"Pedro","suffix":""}],"badges":[],"createdAt":"2026-01-26 01:24:01","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-8695459/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-8695459/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":105564286,"identity":"3fac7855-285c-4200-bad8-c1113d2c2a6c","added_by":"auto","created_at":"2026-03-27 12:49:10","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":442785,"visible":true,"origin":"","legend":"\u003cp\u003ePRISMA flowchart illustrating the steps carried out for the integrative review\u003c/p\u003e","description":"","filename":"floatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-8695459/v1/f60118721d45967b67a6e717.png"},{"id":105201197,"identity":"bfc7cc97-e97a-40ea-b1d9-7858d388f376","added_by":"auto","created_at":"2026-03-23 11:28:40","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":259885,"visible":true,"origin":"","legend":"\u003cp\u003eFigures showing: (a) Percentage distribution of empirical studies, notes, and reviews among the analyzed publications; (b) Main research themes addressed in studies on the Hyacinth Macaw, classified according to sampling type (in situ, ex situ, or not applicable)\u003c/p\u003e","description":"","filename":"floatimage2.png","url":"https://assets-eu.researchsquare.com/files/rs-8695459/v1/aaab6e6cbd0cb5496999cb86.png"},{"id":105201191,"identity":"d6bbcf49-245c-4068-9cc9-196ce3c9fb05","added_by":"auto","created_at":"2026-03-23 11:28:39","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":383309,"visible":true,"origin":"","legend":"\u003cp\u003eVariation in the number of publications about the Hyacinth Macaw over the years. The model indicated a statistically significant increasing trend in publications over time (p \u0026lt; 0.05)\u003c/p\u003e","description":"","filename":"floatimage3.png","url":"https://assets-eu.researchsquare.com/files/rs-8695459/v1/24965613a3fce9a933750176.png"},{"id":105201194,"identity":"fa7f6ffc-2c9a-4acf-afa8-8227b24853b7","added_by":"auto","created_at":"2026-03-23 11:28:39","extension":"jpeg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":221845,"visible":true,"origin":"","legend":"\u003cp\u003eSpatial distribution of scientific studies on the Hyacinth Macaw across populations in Brazil, shown as absolute numbers per population on the map and as relative percentages of publications in the pie chart. The species’ range was obtained from IUCN data, where it is classified as “resident.” The populations in the Pantanal were delimited according to the proposal by Silva and Abdon (1998)\u003c/p\u003e","description":"","filename":"floatimage4.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-8695459/v1/e1ca507c3ad53b0ea3e97cf2.jpeg"},{"id":105201193,"identity":"605cecae-ee8d-4863-90cc-b94fe9060841","added_by":"auto","created_at":"2026-03-23 11:28:39","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":283698,"visible":true,"origin":"","legend":"\u003cp\u003ePercentage representation of scientific studies on the Hyacinth Macaw conducted across different biomes of Brazil\u003c/p\u003e","description":"","filename":"floatimage5.png","url":"https://assets-eu.researchsquare.com/files/rs-8695459/v1/99ef21cc519e817d68009c77.png"},{"id":105569574,"identity":"3d57067d-bdb1-43de-890d-6499773f0623","added_by":"auto","created_at":"2026-03-27 13:12:47","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2395062,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8695459/v1/7e6996af-0b7b-4e9a-9eab-b05d65b41911.pdf"},{"id":105201196,"identity":"1c59a4e8-5474-463e-8c8b-ba335882d109","added_by":"auto","created_at":"2026-03-23 11:28:40","extension":"xlsx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":37291,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eSUPPLEMENTARY MATERIAL\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eSupplementary material supporting this article is available online at the end of the article.\u003c/p\u003e","description":"","filename":"DatabaseSpreadsheetMainDocument.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-8695459/v1/089835b1028c99e0e83459f4.xlsx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Conservation Challenges of the Hyacinth Macaw (Anodorhynchus hyacinthinus (Latham, 1790)) in Brazil: An Integrative Review","fulltext":[{"header":"1. INTRODUCTION","content":"\u003cp\u003eThe family Psittacidae is among the most threatened bird families worldwide, with 29% of its species classified as globally threatened and 58% showing declining population trends (Chan et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Due to their vibrant plumage, high intelligence, longevity, and the ability to mimic the human voice, psittacids have become frequent targets of illegal wildlife trade. These attributes make them highly desirable as pets, generating thousands of dollars annually (Olah et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Chan et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe Hyacinth Macaw (\u003cem\u003eAnodorhynchus hyacinthinus\u003c/em\u003e (Latham, 1790)) is the largest species in the family Psittacidae, reaching approximately 1 m in length (Guedes et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). The species has a wide distribution in Brazil, occurring in three main biomes: the Pantanal, Amazon and Cerrado, and is also found in parts of Bolivia and Paraguay (Vila\u0026ccedil;a et al. \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). In Brazil, four genetically distinct populations are recognised: the Southern Pantanal (Mato Grosso do Sul), the Northern Pantanal (Mato Grosso), the Northern region (including Par\u0026aacute;, Amazonas, Piau\u0026iacute; and northern Tocantins), and the Central region (southern Tocantins, Goi\u0026aacute;s, Bahia and Maranh\u0026atilde;o) (Almeida et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Vila\u0026ccedil;a et al. \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). The Hyacinth Macaw is specialised in feeding on the seeds of palm trees of the genera \u003cem\u003eAttalea\u003c/em\u003e and \u003cem\u003eAcrocomia\u003c/em\u003e, as well as on the fruits of \u003cem\u003eMauritia flexuosa\u003c/em\u003e Mart., depending on the biome in which it occurs (Guedes \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e1993\u003c/span\u003e; Tella et al. 2021). The species also shows specialised reproductive behaviour, nesting primarily in cavities of specific tree species, mainly species of the genus \u003cem\u003eSterculia\u003c/em\u003e (Guedes \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e1993\u003c/span\u003e; Silva et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). However, in Central Brazil, it is also capable of nesting in sandstone cliffs (Pacheco and Olmos \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Dornas et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2013\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eDue to its large body size, high intelligence, longevity and vibrant cobalt-blue plumage, the Hyacinth Macaw has become a frequent target of illegal wildlife trade (Guedes \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). These factors, combined with the species\u0026rsquo; low reproductive rate and high degree of specialisation in feeding and nesting sites, have led to a marked population decline (Guedes \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). In the 1990s, the estimated wild population comprised only about 2,500 individuals (Guedes et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Against this background, studies on the biology and reproductive success of the species in the Pantanal, which historically supported the largest population, were fundamental to its conservation. The establishment of the \u003cem\u003eArara Azul\u003c/em\u003e Project in 1990, informed by the knowledge generated by these studies, enabled the implementation of a range of conservation actions (Guedes \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Kuniy et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). Following these efforts, a substantial increase in the Hyacinth Macaw population was observed, with estimates of approximately 6,500 individuals in the wild (Guedes \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Guedes \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). In 2014, the species was removed from Brazil\u0026rsquo;s official list of threatened species and reclassified as \u0026lsquo;Near Threatened\u0026rsquo;. However, at the global level, it remains categorised as \u0026lsquo;Vulnerable\u0026rsquo; by the IUCN (IUCN 2025).\u003c/p\u003e \u003cp\u003eThus, the \u003cem\u003eArara Azul\u003c/em\u003e Project has become a leading reference for the conservation of the Hyacinth Macaw in Brazil, playing a central role in knowledge generation and fostering numerous studies over recent decades. In addition, the project has supported educational initiatives and established partnerships with ranchers, schools and local communities. While many of these studies involved direct or indirect participation by the \u003cem\u003eArara Azul\u003c/em\u003e Institute, others were conducted by independent research institutions. However, the resulting knowledge remains fragmented across the scientific literature, and to our knowledge no study has yet synthesised this information in a comprehensive way, despite its importance for advancing conservation strategies for the species in Brazil.\u003c/p\u003e \u003cp\u003eThis study aims to synthesise and critically assess the current state of knowledge on the Hyacinth Macaw in Brazil through an integrative review of the scientific literature published over recent decades, with a particular focus on the main challenges to the species\u0026rsquo; conservation. Specifically, we aim to: (i) systematise available knowledge to support management actions, research and public policies for conservation; (ii) assess genetic aspects of Hyacinth Macaw populations; (iii) identify the main threats to the species and how they vary across its distribution; (iv) examine interpopulation variation in feeding and reproductive behaviour; and (v) highlight knowledge gaps and challenges for effective conservation across different biomes.\u003c/p\u003e"},{"header":"2. METHODS","content":"\u003cp\u003eThis study is an integrative review conducted following the methodological framework proposed by Whittemore and Knafl (\u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). The review comprised five stages: (1) problem identification, as defined above; (2) literature search; (3) selection of scientific papers; (4) critical assessment and categorisation of the selected studies; and (5) presentation and synthesis of the results, with the final two stages addressed in the Results and Discussion sections of this paper.\u003c/p\u003e \u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003e2.1 Literature search\u003c/h2\u003e \u003cp\u003eThis review focused exclusively on scientific articles published in indexed journals. This approach was adopted to ensure that the results analysed were derived from methodologically robust studies that are widely recognised by the scientific community. Although dissertations, theses, books, book chapters and conference abstracts may also contain relevant information, articles published in indexed journals were prioritised because they offer greater methodological standardisation, transparency and data traceability, as well as improved comparability across studies. These features are essential for integrative reviews that aim to produce a critical synthesis of knowledge.\u003c/p\u003e \u003cp\u003eThe literature search was conducted between October 2024 and February 2025 across six databases: ScienceDirect, Web of Science, SpringerLink, Scopus, SciELO and Google Scholar. The following keywords were used: \u0026lsquo;Anodorhynchus hyacinthinus\u0026rsquo;, \u0026lsquo;Hyacinth Macaw\u0026rsquo;, \u0026lsquo;Genetics\u0026rsquo;, \u0026lsquo;Threats\u0026rsquo;, \u0026lsquo;Ecology\u0026rsquo;, \u0026lsquo;Conservation\u0026rsquo;, \u0026lsquo;Conservation of Psittacidae in Brazil\u0026rsquo; and \u0026lsquo;Psittacidae behaviour in Brazil\u0026rsquo;, combined using the Boolean operators \u0026lsquo;AND\u0026rsquo; and \u0026lsquo;OR\u0026rsquo;. To ensure consistency and rigour in the selection process, specific inclusion and exclusion criteria were applied to each database. Only peer-reviewed articles published in indexed journals were considered, including empirical studies, reviews and scientific notes. Studies that focused directly on the Hyacinth Macaw were included, as well as studies addressing ecological components essential to the species, such as plant species used for feeding and nesting, which are fundamental for understanding its ecology and conservation. Duplicate records and publications in languages other than Portuguese or English were excluded.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003e2.2 Selection of scientific studies\u003c/h2\u003e \u003cp\u003eAll records retrieved from the databases were exported and compiled into a single spreadsheet. Duplicate records across databases were then removed using the duplicate removal tool in Google Sheets, and any remaining duplicates were eliminated manually. Subsequently, the selection process was conducted in three screening phases: (1) title screening, to exclude papers clearly unrelated to the topic; (2) abstract screening, applying the inclusion and exclusion criteria to refine the selection; and (3) full-text screening of the remaining articles to confirm their relevance and eligibility. Only studies that met all criteria following this final assessment were included in the review.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003e2.3. Search outcomes\u003c/h2\u003e \u003cp\u003eIn total, 5,648 publications were identified, of which 2,951 were duplicates, leaving 2,697 unique records (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Of these, 350 were retrieved from ScienceDirect, 489 from SpringerLink, 582 from Web of Science, 251 from Scopus, 25 from SciELO and 1,000 from Google Scholar. During the first screening stage (title screening), 2,522 publications were excluded because they did not focus on the Hyacinth Macaw, leaving 175 articles selected based on their titles. In the second stage (abstract screening), 84 articles were excluded, resulting in 91 studies. Finally, during full-text screening, 20 articles were excluded because they were unrelated to the species, unavailable or not peer-reviewed. Thus, 71 papers were selected for inclusion in this review (see Supporting Information).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003e3.3 Critical assessment and categorization of the selected studies\u003c/h2\u003e \u003cp\u003eFor each of the 71 studies included in the review, information was compiled and categorised in a spreadsheet, including the title, authors, year of publication, from which the study was retrieved, population studied, biome, animal classification (wild or captive) and study type (review, empirical study or scientific note). The species\u0026rsquo; populations were classified following Vila\u0026ccedil;a et al. (\u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) and divided into Northern and Southern Pantanal, the Central region, and the Northern region of Brazil. In addition, the articles were grouped into five main categories, as shown in Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eThemes of the studies selected for the review and their respective aspects.\u003csup\u003e2\u003c/sup\u003e\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"2\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eThemes\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAspects\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEcology\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eStudies on species\u0026rsquo; spatial distribution, feeding habits, reproduction, ecological interactions (intra- and interspecific), and population density estimates.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eGenetics\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eArticles addressing the genetic aspects of the species, including genetic variation within populations.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBehaviour\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eArticles published in journals specialized in animal behaviour and/or containing behaviour-related terms in the keywords, title, or abstract.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAnimal Health\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eStudies addressing health aspects, including clinical, physiological, or sanitary conditions of the species.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eConservation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eArticles focused on the identification of threats, implementation of conservation strategies, population management, and environmental education initiatives aimed at the conservation of the species.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003csup\u003ea\u003c/sup\u003e Each study was assigned to a single thematic category according to its primary focus.\u003c/p\u003e \u003cp\u003eBased on full-text assessment of the selected studies, the main threats to the species, genetic aspects of its populations, current conservation strategies and initiatives, interpopulation variation in feeding and reproductive behaviour, as well as key knowledge gaps and challenges for effective conservation of the Hyacinth Macaw across different biomes were systematically identified and synthesised.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003e2.4 Data analysis\u003c/h2\u003e \u003cp\u003eTo analyze the relationship between the number of publications over time, a Poisson regression was applied, with results graphically represented using the \u0026lsquo;ggplot2\u0026rsquo; package in R v. 4.2.2. A significance level of p\u0026thinsp;\u0026lt;\u0026thinsp;0.05 was adopted. A map showing the spatial distribution of studies across Hyacinth Macaw populations in Brazil was produced using QGIS v. 3.28.12. The distribution ranges of each population were obtained from the IUCN Red List, where the species is classified as resident, and Pantanal populations were subdivided following the regional delimitation proposed by Silva and Abdon (1998). Additional graphs summarising the results were also generated using \u0026lsquo;ggplot2\u0026rsquo; to support the quantitative analyses.\u003c/p\u003e \u003c/div\u003e"},{"header":"3. RESULTS AND DISCUSSION","content":"\u003cp\u003eAmong the 71 articles analysed, most were classified as empirical studies (74%, n\u0026thinsp;=\u0026thinsp;53), followed by reviews (13%, n\u0026thinsp;=\u0026thinsp;9) and scientific notes (13%, n\u0026thinsp;=\u0026thinsp;9) (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Research conducted on wild individuals predominated, accounting for 60% (n\u0026thinsp;=\u0026thinsp;42) of the publications, whereas studies involving captive individuals represented only 14% (n\u0026thinsp;=\u0026thinsp;10) of the total (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). In addition, 26% (n\u0026thinsp;=\u0026thinsp;19) of the selected articles did not focus directly on the species itself, consisting mainly of theoretical reviews or studies on plant species ecologically associated with the Hyacinth Macaw, which provide essential resources for feeding and nesting.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eStudies based on captive individuals addressed topics such as pathogen occurrence (Barros et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2002\u003c/span\u003e), bacterial resistance to antibiotics (Ramos et al. \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), longevity and sexual maturation (Young et al. \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Castaldo et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Barros et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Barros et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), tool use (Borsari and Ottoni \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Paim et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2008\u003c/span\u003e), karyotyping (Lunardi et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2003\u003c/span\u003e), bite force (Harrison et al. 2024) and factors influencing species selection for ex situ conservation programmes (Coll\u0026eacute;ony et al. 2017). However, none of these studies directly addressed conservation actions targeting the species, such as reintroduction, restocking or integrated management of captive populations for conservation purposes. The limited number of studies involving captive individuals therefore represents an important limitation for ex situ conservation strategies and, indirectly, for in situ conservation of the Hyacinth Macaw in Brazil. This gap hampers, for example, the development of genetic conservation measures, such as planned reproductive management aimed at maintaining adequate genetic variability within captive populations. Such variability is essential both for the genetic reinforcement of wild populations and for reintroduction programmes in areas where the species has become locally extinct (Vila\u0026ccedil;a et al. \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). These activities appear to remain a low priority within management strategies for Neotropical psittacids (Berkunsky et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eBased on the articles included in this review, 294 individual authors were identified. Neiva M. R. Guedes, president of the \u003cem\u003eArara Azul\u003c/em\u003e Institute, which coordinates the \u003cem\u003eArara Azul\u003c/em\u003e Project, was the most frequently cited author, appearing in 38% (n\u0026thinsp;=\u0026thinsp;27) of the publications, followed by Fl\u0026aacute;via T. Presti (8%, n\u0026thinsp;=\u0026thinsp;6) and Fernanda M. Fontoura (7%, n\u0026thinsp;=\u0026thinsp;5). The prominent role of the Institute is also reflected in the temporal pattern of scientific publications on the species. A trend analysis using Poisson regression (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e) revealed a significant increase in the number of publications between 1997 and 2024 (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05). This sustained growth in scientific output appears to be associated with the long-term research and monitoring activities conducted by the \u003cem\u003eArara Azul\u003c/em\u003e Institute and the dissemination of results generated by its researchers. Overall, the findings of this review highlight the relevance of the \u003cem\u003eArara Azul\u003c/em\u003e Project in consolidating scientific knowledge on the species in Brazil and its importance as one of the main in situ conservation initiatives. The knowledge generated by the project has contributed substantially to understanding the ecological requirements of the Hyacinth Macaw, identifying key threats across its range and supporting evidence-based management strategies aimed at conserving the species in its natural environment.\u003c/p\u003e \u003cdiv id=\"Sec9\" class=\"Section2\"\u003e \u003ch2\u003e3.1 Genetic aspects of Hyacinth Macaw populations in Brazil\u003c/h2\u003e \u003cp\u003eStudies focusing on genetics accounted for approximately 11% (n\u0026thinsp;=\u0026thinsp;8) of the articles selected in this review (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). The first genetic studies on Hyacinth Macaw populations in Brazil were conducted by Faria et al. (\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2008\u003c/span\u003e), who analysed the genetic structure of wild individuals from two regions of the country: Piau\u0026iacute; and Mato Grosso do Sul. This pioneering study revealed genetic differences between geographically distant populations and demonstrated the value of genetic approaches for identifying individuals seized from the illegal wildlife trade. Subsequently, Presti et al. (\u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) and Almeida et al. (\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) expanded these analyses by incorporating samples from previously unstudied regions. Using nuclear and mitochondrial markers, these studies identified genetic differentiation among the currently recognised populations. Both studies also reported relatively low genetic diversity compared to other threatened psittacine species. However, they present methodological limitations that may have influenced these results, such as the use of heterologous markers (Presti et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) and the exclusive use of mitochondrial markers, which do not capture the full genetic variability of the species (Almeida et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Although informative, these findings represent only a fraction of the species\u0026rsquo; total genetic variability, highlighting the need to complement existing data with analyses based on species-specific nuclear markers, including those developed by Silva et al. (\u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2015\u003c/span\u003e), or with genomic approaches.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eA recent study by Vila\u0026ccedil;a et al. (\u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), using low-coverage genomic data, produced results broadly consistent with previous studies. However, the authors identified relatively high genetic diversity, particularly in the Southern Pantanal population, as well as evidence of recent gene flow among populations, mainly involving the Northern Pantanal. The high genetic diversity observed in the Southern Pantanal may be associated, at least in part, with gene flow between different regions. The ecological dynamics of the Northern Pantanal provide important insights into the patterns of gene flow reported by Vila\u0026ccedil;a et al. (\u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). One of the main factors that may explain this pattern is the high emigration rate of individuals, as suggested by Neto, Guedes and Toledo (\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), who identified the largest known roost of the species in Brazil in this region, comprising approximately one thousand individuals. Such high concentrations may promote dispersal to other areas, particularly in response to the scarcity of nesting sites and food resources, conditions that tend to intensify movement. These movements may influence gene flow and, consequently, genetic variability by enhancing connectivity among populations and partially reducing genetic isolation.\u003c/p\u003e \u003cp\u003eDespite these potential movements, the study by Vila\u0026ccedil;a et al. (\u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) reported high levels of inbreeding, suggesting that the observed gene flow may be recent, limited or insufficient to mitigate the established genetic structure. This highlights the need for continued genetic studies involving representative samples from different parts of the species\u0026rsquo; distribution. Such analyses are important for determining whether high levels of inbreeding pose a risk to population viability and whether strategies such as genetic reinforcement would be justified as part of species conservation efforts.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec10\" class=\"Section2\"\u003e \u003ch2\u003e3.2 Ecological aspects of Hyacinth Macaw populations in Brazil\u003c/h2\u003e \u003cp\u003eStudies related to ecology accounted for 44% (n\u0026thinsp;=\u0026thinsp;31) of the analysed publications, whereas those focusing on behaviour represented 6% (n\u0026thinsp;=\u0026thinsp;4), making ecology the most frequently addressed research topic on the species (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Most ecological studies (61%, n\u0026thinsp;=\u0026thinsp;19) were conducted on Pantanal populations. In addition, 13% (n\u0026thinsp;=\u0026thinsp;4) of these studies did not focus directly on the species itself, consisting mainly of theoretical reviews or studies on plant species ecologically associated with the Hyacinth Macaw, which provide essential resources for feeding and nesting.\u003c/p\u003e \u003cp\u003eAnalysis of the geographical distribution of studies revealed that most research focused on the Southern Pantanal population, which accounted for 44% (n\u0026thinsp;=\u0026thinsp;31) of the records, followed by the Northern Pantanal population with 18% (n\u0026thinsp;=\u0026thinsp;13) of the studies (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). In addition, 38% (n\u0026thinsp;=\u0026thinsp;27) of the studies did not address specific populations or focused on plant species ecologically associated with the Hyacinth Macaw. Overall, Pantanal populations emerge as the most extensively studied in Brazil, a pattern that appears to be associated with the presence of the \u003cem\u003eArara Azul\u003c/em\u003e Project in the region, which has enabled continuous, long-term monitoring. Consequently, a substantial portion of the available literature on the ecology and behaviour of the Hyacinth Macaw reflects the sustained research efforts undertaken within the scope of this project.\u003c/p\u003e \u003cp\u003eBy contrast, this review highlights a marked shortage of information on the ecology and behaviour of the species in the Central and Northern populations of Brazil. This gap limits comprehensive assessments and the formulation of effective, population-specific conservation strategies. The few studies conducted in these regions (n\u0026thinsp;=\u0026thinsp;4 in the Central population and n\u0026thinsp;=\u0026thinsp;1 in the Northern population) consisted primarily of faunal inventories which, although valuable, do not provide detailed or systematic information on the species\u0026rsquo; ecology. This highlights the need to expand research efforts focused on the Central and Northern populations in order to better understand their specific ecological requirements and local threats. Such efforts would contribute to the development of more effective and targeted conservation strategies for each population.\u003c/p\u003e \u003cp\u003eThe results of this review indicate that the Pantanal was the most frequently studied biome, representing 48% (n\u0026thinsp;=\u0026thinsp;34) of the publications, followed by the Cerrado (20%, n\u0026thinsp;=\u0026thinsp;14) and the Amazon (11%, n\u0026thinsp;=\u0026thinsp;11) (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). Notably, 38% of the analysed papers addressed the Neotropical region more broadly. When considering only studies focused on Brazilian populations, the Pantanal holds the largest volume of available information, making it central to current understanding of the species\u0026rsquo; ecology and conservation in Brazil. In contrast, populations in the Central region, located in the Cerrado, and in the Northern region of the country, located in the Amazon, remain poorly studied. These knowledge gaps could be reduced through increased sampling effort and targeted research in these regions, aimed at better understanding the species\u0026rsquo; ecological requirements and ensuring population viability across its range. Citizen science initiatives represent a promising approach to expand knowledge of the Hyacinth Macaw by involving local communities, guides, birdwatchers and partner institutions in the collection of ecological and occurrence data. Such collaborative efforts can help address monitoring gaps and strengthen participatory conservation strategies.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cdiv id=\"Sec11\" class=\"Section3\"\u003e \u003ch2\u003e3.2.1 Diet and Feeding Ecology\u003c/h2\u003e \u003cp\u003eThe studies analysed in this review indicate that the diet of Hyacinth Macaws in the Southern and Northern Pantanal populations consists largely of the seeds of two palm species that are widely distributed in the region: \u003cem\u003eAttalea phalerata\u003c/em\u003e Mart. ex Spreng and \u003cem\u003eAcrocomia aculeata\u003c/em\u003e (Jacq.) Lodd. ex Mart. (Tella et al. \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Guedes et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). \u003cem\u003eA. phalerata\u003c/em\u003e is the most frequently consumed food item in the Pantanal, a pattern that can be explained by a combination of traits of this palm species, including its high abundance, continuous fruiting throughout the year, the high lipid content of its seeds, which is among the highest within the Arecaceae, and low competition pressure, as relatively few species feed on its seeds (Negrelle \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2015\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe Hyacinth Macaw has morphological specializations that allow it to handle the \u003cem\u003eA. phalerata\u003c/em\u003e fruits, including adaptations of the skull, beak, and tongue, which result in one of the strongest bite forces among Psittacidae (Blanco et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Harrison et al. 2024). Although \u003cem\u003eA. phalerata\u003c/em\u003e fruits have a very hard endocarp, individuals from these populations tend to select fruits with thinner endocarps (Barros and Pires \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). This preference may be interpreted as an evolutionary response to the structural defences of the plant, likely arising from a coevolutionary relationship between the two species.\u003c/p\u003e \u003cp\u003eIn addition to morphological and evolutionary adaptations, some behavioral strategies related to \u003cem\u003eA. phalerata\u003c/em\u003e consumption have also been documented. Schneider et al. (\u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2006\u003c/span\u003e) observed that some macaws remove pieces of \u003cem\u003eA. phalerata\u003c/em\u003e leaves to facilitate fruit handling and make it easier to crack the endocarp. The authors also reported that macaws frequently drop fruits while manipulating the bunches and later return to the site, approximately one month later, to feed on the seeds of the fallen fruits, which contained insect larvae that were likely consumed. A similar behaviour was reported by Paula et al. (2015), who observed a pair of Hyacinth Macaws consuming termites from pieces of wood from a nest infested by these insects in a transition area between the Southern Pantanal and the Cerrado. This behaviour may be interpreted as a form of protein supplementation, particularly during the breeding season, when protein demands are higher.\u003c/p\u003e \u003cp\u003eBased on the literature analysed, some studies report that in the Pantanal, particularly in cattle ranching areas, groups of Hyacinth Macaws are commonly observed foraging on the ground in search of \u003cem\u003eA. phalerata\u003c/em\u003e fruits (Schneider et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Tella et al. \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Both cattle and \u003cem\u003eTapirus terrestris\u003c/em\u003e Linnaeus, 1758 consume the mesocarp of \u003cem\u003eA. phalerata\u003c/em\u003e fruits and subsequently regurgitate or excrete the endocarps, thereby facilitating access to the seeds by macaws (Schneider et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Barros and Pires \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eRegarding the diet of the Northern population, no studies were found that directly investigated the diet of the Hyacinth Macaw in this region or the foraging behaviours associated with it. Available information is limited to indirect evidence reported in isolated studies, in which groups of individuals were observed in \u003cem\u003eMauritia\u003c/em\u003e palm swamps and in ecosystems with a high abundance of palms of the genus \u003cem\u003eAttalea\u003c/em\u003e, particularly \u003cem\u003eAttalea maripa\u003c/em\u003e (Aubl.) Mart. (Pacheco and Olmos \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Dornas et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). These observations suggest that these palm species may represent the main food resources for the population in this region.\u003c/p\u003e \u003cp\u003eFor the Central population, the limited available data indicate that \u003cem\u003eMauritia flexuosa\u003c/em\u003e is likely to represent a primary food resource for the Hyacinth Macaw in this region. This palm can occur dominantly in certain Cerrado areas, substantially contributing to local food availability (Dornas et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Tella et al. \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). In addition to \u003cem\u003eM. flexuosa\u003c/em\u003e, other palms used as food resources in this region include \u003cem\u003eAttalea eichleri\u003c/em\u003e (Drude) A. J. Hend. and \u003cem\u003eAttalea barreirensis\u003c/em\u003e Glassman (Pacheco and Olmos \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Tella et al. \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Unlike what is observed in other regions, such as the Pantanal, where the species predominantly consumes palm seeds, Tella et al. (\u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) reported that individuals from this population feed mainly on the pericarp of \u003cem\u003eM. flexuosa\u003c/em\u003e fruits, thereby playing a significant role in the dispersal of this species\u0026rsquo; seeds.\u003c/p\u003e \u003cp\u003eAlthough initially considered a purely antagonistic relationship, the interaction between the Hyacinth Macaw and the palm species it feeds on in different regions represents a continuum between antagonism and mutualism (Blanco et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Dracxler and Kissling \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Consequently, the species can act as an important seed disperser, as fruits are often removed directly from palms and transported to safer locations for consumption, where fruits or seeds may be dropped. This process can significantly contribute to the colonisation and persistence of these palm species across the landscape (Blanco et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Tella et al. \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Thus, the Hyacinth Macaw also performs an ecological role analogous to that once played by members of the extinct megafauna, since few current species are able to carry out this interaction. Therefore, future studies should investigate these ecological relationships in greater depth, assessing the impact of the absence of the Hyacinth Macaw on the demography and population dynamics of \u003cem\u003eAttalea\u003c/em\u003e species and other palms consumed by the macaw.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec12\" class=\"Section3\"\u003e \u003ch2\u003e3.2.2 Reproductive Biology\u003c/h2\u003e \u003cp\u003eIn addition to its highly specialised diet, the Hyacinth Macaw also exhibits strong specialisation in the selection of nesting sites. In the Pantanal, the species primarily nests in natural cavities of \u003cem\u003eSterculia apetala\u003c/em\u003e (Jacq.) H. Karst., a tree species typical of this biome. According to Guedes et al. (\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), approximately 95% of natural nests in the Southern Pantanal are found exclusively in \u003cem\u003eS. apetala\u003c/em\u003e trees, whereas in the Northern Pantanal this proportion is slightly lower, at around 85% (Pinho and Nogueira \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2003\u003c/span\u003e). \u003cem\u003eS. apetala\u003c/em\u003e exhibits a set of physical and chemical traits that favour its selection as a nesting site, including soft wood, rapid growth, and the frequent presence of natural cavities suitable for the reproduction of several Pantanal species. Furthermore, this species contains chemical compounds with fungicidal properties that inhibit pathogen proliferation within nests, thereby contributing to increased nestling survival (Fontoura et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2015\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eNatural cavities formed in \u003cem\u003eS. apetala\u003c/em\u003e trees function as important nesting sites, being used not only by the Hyacinth Macaw but also by several other wildlife species in the Pantanal (Carrara et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). However, for these cavities to be suitable for Hyacinth Macaws, trees must be at least 60 years old (J\u0026uacute;nior et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). Despite this, it is estimated that only about 5% of adult \u003cem\u003eS. apetala\u003c/em\u003e trees in the Southern Pantanal contain cavities with characteristics suitable for the species, making this a scarce and highly contested resource during the breeding season (Johnson et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; J\u0026uacute;nior et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). Future studies could therefore focus on constructing interaction networks centred on cavity availability in \u003cem\u003eS. apetala\u003c/em\u003e, aiming to better understand the relationships between the Hyacinth Macaw and other cavity-dependent species, as well as to characterise patterns of agonistic interactions and degrees of dependence among Pantanal fauna. Such an approach would allow a more precise assessment of the relative dependence of different species on \u003cem\u003eS. apetala\u003c/em\u003e, which is recognised as a keystone species in Pantanal ecosystems. In addition, the application of population dynamics models for \u003cem\u003eS. apetala\u003c/em\u003e could help project the future availability of nesting cavities in key areas of the species\u0026rsquo; range, thereby providing valuable support for conservation planning.\u003c/p\u003e \u003cp\u003eRegarding the Northern population, among the few studies addressing reproduction, the work of Silva et al. (\u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) stands out. While investigating the species\u0026rsquo; ecology within a mosaic of conservation units in Caraj\u0026aacute;s, southeastern Par\u0026aacute;, the authors observed that more than 80% of the nests used by this population were located in natural cavities of \u003cem\u003eSterculia pruriens\u003c/em\u003e (Aubl.) K. Schum. In addition, most nests were found in open environments (87.5%), followed by forest interiors (8.3%) and forest edges (4.2%).\u003c/p\u003e \u003cp\u003eWith respect to reproduction in the Central population, the available information is largely based on field observations and is therefore inconclusive (Pacheco and Olmos \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Dornas et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Reports indicate nesting on sandstone cliffs in eastern Tocantins, particularly in the Jalap\u0026atilde;o region, as well as possible nesting in cavities of \u003cem\u003eMauritia flexuosa\u003c/em\u003e palms (Pacheco and Olmos \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Rego et al. \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). However, no study specifically addressing the reproductive biology of the species in this region was included in this review, representing a significant gap in understanding the factors influencing local reproductive success. The potential use of sandstone cliffs as nesting sites suggests that the Hyacinth Macaw population in this region may exhibit ecological adaptations distinct from those of the Pantanal and Amazon populations. Such differentiation may be associated with local factors, including a low availability of natural tree cavities, reduced predation pressure leading to higher nestling survival, or lower interspecific competition for nesting sites. Nonetheless, these hypotheses remain untested, highlighting the need for targeted studies on the species\u0026rsquo; ecology in the Central region to generate systematic data and enable comparisons with other populations.\u003c/p\u003e \u003cp\u003eAccording to available data from studies conducted in the Pantanal, the Hyacinth Macaw exhibits relatively low reproductive success. Females may lay between one and three eggs, with an average of two eggs per pair (Ramalho et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). However, of every 100 eggs laid, only approximately 25% hatch and survive to the age at which nestlings typically leave the nest (Guedes et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Egg laying in this species is asynchronous, meaning that after the first egg is laid, there may be an interval of 1 to 16 days before the second egg is laid. This strategy is thought to reduce the effects of predation and increase the likelihood that at least one nestling survives (Kuniy et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Ramalho et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Nevertheless, when both eggs hatch, the second nestling is often rejected by the parents. There are, however, records of pairs successfully raising two nestlings, depending on factors such as parental experience, food availability, and the age difference between chicks (Guedes et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eDuring incubation, the female remains inside the nest for most of the time, where she protects and incubates the eggs. The male, in turn, acts as a sentinel, defending the nest against predators and competitors, and is responsible for foraging and feeding the female (Pinho and Nogueira \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Schneider et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). An additional noteworthy aspect is that slight sexual dimorphism is observable only at the beginning of the breeding season, when females may exhibit a slightly curved tail as a result of prolonged periods spent incubating the eggs inside the nest (Schneider et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2006\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eEgg predation represents a significant threat to reproductive success, with \u003cem\u003eRamphastos toco\u003c/em\u003e Statius Muller, 1776 accounting for approximately 53% of predation events recorded in the Southern Pantanal (Pizo et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Paradoxically, \u003cem\u003eR. toco\u003c/em\u003e is also the most important seed disperser of \u003cem\u003eSterculia apetala\u003c/em\u003e, being responsible for about 86% of the species\u0026rsquo; seed dispersal in the Pantanal (Pizo et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). This dispersal plays a crucial role in the regeneration and maintenance of trees that develop natural cavities suitable for nesting.\u003c/p\u003e \u003cp\u003eThe incubation period of Hyacinth Macaw eggs lasts on average 28 to 30 days. During the final stages of incubation, the embryo begins to vocalise inside the egg, which likely serves as a stimulus for parental assistance with thermoregulation and the hatching process (Ramalho et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). After hatching, nestling development occurs in three stages, with fledging taking place at an average age of 107 days. However, juveniles remain under parental care for up to 12 months, a period that is essential for survival in the wild (Guedes et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). In the Northern population, nests containing nestlings at different developmental stages have been reported, indicating asynchrony in the reproductive cycle among breeding pairs. This pattern contrasts with that observed in Pantanal populations, where greater synchrony among pairs has been documented (Silva et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eMoreover, some individuals exhibit developmental abnormalities, characterised by body mass and overall weight gain that are approximately 54% lower than those of typically sized individuals, and are therefore referred to as \u0026ldquo;dwarfs\u0026rdquo; (Guedes et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Given that these individuals represent a small fraction of the population and that the ecological and reproductive implications of this condition remain poorly understood, targeted studies are needed to assess whether it affects individual fitness.\u003c/p\u003e \u003c/div\u003e \u003c/div\u003e \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003e3.3 Threats and Challenges to the Conservation of the Hyacinth Macaw in Brazil\u003c/h2\u003e \u003cp\u003eAmong studies addressing threats to the Hyacinth Macaw in Brazil, Devenish et al. (\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) reported an expansion of the species\u0026rsquo; range between 1990 and 2019. However, approximately 30% of this area was converted into pastures and agricultural land, and only 8% is currently located within Protected Areas, raising concerns about the effectiveness of existing conservation measures. In addition, previous population estimates indicating around 6,500 individuals in the wild (Guedes \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2004\u003c/span\u003e) reflect a past demographic scenario. Therefore, updated population surveys are required to provide more accurate estimates of the species\u0026rsquo; current status. Such surveys could be conducted through collaborations between South American non-governmental organizations and Citizen Science initiatives, promoting the involvement of local communities in data collection, expanding monitoring efforts, and strengthening participatory conservation.\u003c/p\u003e \u003cp\u003eDespite significant advances in knowledge about the species, the available data still reveal important knowledge gaps, reflecting broader spatial and thematic biases reported for biodiversity research in the Pantanal (Frota et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The Central region of the Pantanal, for instance, remains poorly sampled and studied. In this review, only one article related to this area was identified, reporting a case of cavity competition between a pair of Hyacinth Macaws and \u003cem\u003eTyto furcata\u003c/em\u003e (Temminck, 1827), which suggests a possible scarcity of suitable natural cavities (Tortato and Bonanomi \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Given this scenario, it is essential that future studies conduct surveys not only in areas already known to host the Hyacinth Macaw but also in poorly sampled or still unexplored regions, such as the municipality of Parintins in the state of Amazonas (Barreiros and Gomes \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), as well as in the Central population, which currently lacks studies focused on the species\u0026rsquo; biology and ecology.\u003c/p\u003e \u003cp\u003eSuch surveys are crucial to obtain more accurate estimates of wild population size, identify priority areas for conservation, recognize region-specific threats, and better understand the ecological requirements of the species across different population contexts. This information is also fundamental to support the creation of new Protected Areas and the management of potentially isolated populations within the landscape. In this sense, the Hyacinth Macaw represents a symbol of Brazilian biodiversity, and its conservation is essential not only for future generations to understand the importance of the species but also for the preservation of entire ecosystems on which many other species depend (Douglas and Ver\u0026iacute;ssimo \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2013\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eAs proposed by Berkunsky et al. (\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) and Vila\u0026ccedil;a et al. (\u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), a conservation approach based on population management units is recommended for the Hyacinth Macaw. This strategy allows the identification and mitigation of local threats, the containment of potential population declines, and the preservation of the species\u0026rsquo; genetic variability. Each biome in which the Hyacinth Macaw occurs is subject to distinct anthropogenic pressures and specific socio-environmental contexts, which require the development of differentiated management strategies and the definition of priority areas for monitoring. In the following subsections, the main threats and challenges faced by the species\u0026rsquo; populations in the Pantanal, Amazon and Cerrado biomes will be addressed, highlighting the importance of conservation actions adapted to regional specificities.\u003c/p\u003e \u003cdiv id=\"Sec14\" class=\"Section3\"\u003e \u003ch2\u003e3.3.1 Pantanal\u003c/h2\u003e \u003cp\u003eAs previously noted in this review, the Pantanal stands out as the most important biome for the conservation of the Hyacinth Macaw in Brazil. Complementary data from the literature, such as those presented by Oliveira et al. (\u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2021\u003c/span\u003ea), indicate that only 8.8% of the occurrence areas of the three key species, the Hyacinth Macaw, \u003cem\u003eS. apetala\u003c/em\u003e, and \u003cem\u003eA. phalerata\u003c/em\u003e, are located within conservation units in the biome. Moreover, these authors reported a 13% loss of these occurrence areas between 2002 and 2017, mainly attributed to deforestation and the expansion of agricultural activities.\u003c/p\u003e \u003cp\u003eDue to the characteristic flood regime of the Pantanal, agricultural activities are concentrated in ecosystems not subject to flooding, such as \u003cem\u003ecap\u0026otilde;es\u003c/em\u003e and \u003cem\u003ecordilheiras\u003c/em\u003e, which largely overlap with the occurrence areas of plant species essential to the Hyacinth Macaw\u0026rsquo;s life cycle (Oliveira et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2021\u003c/span\u003eb). Cattle pressure in these areas directly affects the density of \u003cem\u003eS. apetala\u003c/em\u003e seedlings and compromises the regeneration of \u003cem\u003eA. phalerata\u003c/em\u003e, mainly through soil compaction that inhibits seed emergence (Johnson et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Tella et al. \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). In addition, these regions have been deforested and converted into pastures or monoculture crops, reducing the availability of breeding and feeding sites for the species. The data analysed in this review and in the literature indicate the need to establish new conservation units in non-flooded areas, particularly in strategic regions that promote connectivity between the Northern and Southern Pantanal populations, such as the Central Pantanal. Another essential measure is the development of public policies and specific legislation that restrict, discourage, and penalise deforestation in \u003cem\u003ecap\u0026otilde;es\u003c/em\u003e and \u003cem\u003ecordilheiras\u003c/em\u003e, especially in the southern portion of the Pantanal. Regulations aimed at controlling logging and vegetation suppression in these areas can strengthen oversight and environmental licensing, thereby preventing irregular and illegal activities.\u003c/p\u003e \u003cp\u003eIn addition to deforestation, fire represents one of the greatest threats to Hyacinth Macaw populations in the Pantanal, as it directly affects key resources used for nesting and feeding. According to Ferreira et al. (2021), most of the distribution of the Hyacinth Macaw, \u003cem\u003eSterculia apetala\u003c/em\u003e and \u003cem\u003eAttalea phalerata\u003c/em\u003e historically occurs in areas that have been little affected by fire. However, due to climate change and the conversion of native vegetation into pastures, fires have become more intense and widespread, driven by increasingly frequent and severe droughts in recent decades. In 2020, approximately 28% of the areas considered suitable for the Hyacinth Macaw were affected by wildfires, as well as 25% of the areas suitable for \u003cem\u003eS. apetala\u003c/em\u003e and \u003cem\u003eA. phalerata\u003c/em\u003e. These values were about five times higher than the annual average recorded between 2003 and 2019 (Ferreira et al. 2021). Fires directly compromise food availability, as the fruits of \u003cem\u003eA. phalerata\u003c/em\u003e and \u003cem\u003eA. aculeata\u003c/em\u003e become unsuitable for consumption after burning. In addition, fire negatively affects reproductive success, since the peak of fire occurrence in the Pantanal, in September, coincides with the peak of the species\u0026rsquo; breeding season. Natural cavities used as nests may be destroyed or structurally compromised during fires, rendering them unusable in subsequent breeding seasons and further reducing the availability of suitable nesting sites.\u003c/p\u003e \u003cp\u003eAmong the effective mitigation strategies, integrated fire management stands out, including preventive actions such as environmental education campaigns, coordinated and structured firefighting efforts, and the controlled use of fire in areas with high loads of dry combustible material, particularly around \u003cem\u003ecap\u0026otilde;es\u003c/em\u003e and \u003cem\u003ecordilheiras\u003c/em\u003e. Complementary to these measures, geotechnologies such as Remotely Piloted Aircraft Systems (RPAS) and Geographic Information Systems (GIS) have proven to be essential tools. These technologies enable spatial monitoring of risk areas, integration of environmental variables, and the production of easily interpretable information to support decision-making. Their effectiveness in conservation actions in the Pantanal has already been demonstrated (Guedes et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). The use of aerial imagery obtained from RPAS or satellite platforms, combined with data on relative humidity and temperature, allows the identification of areas with high fire risk and the continuous monitoring of environmental conditions, which are fundamental measures for the protection of terrestrial ecosystems and the conservation of the Hyacinth Macaw.\u003c/p\u003e \u003cp\u003eAnother emerging threat to the Hyacinth Macaw is environmental contamination associated with agricultural activities. Studies conducted by Marchesi et al. (\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) identified high cadmium concentrations in nestlings from the Southern Pantanal, indicating exposure to heavy metal contamination. In the same region, Vicente and Guedes (\u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) also reported mortality events among individuals of the species resulting from the improper use of pesticides. Although these occurrences appear to be sporadic, they are alarming and highlight the need for systematic monitoring in areas under intensive agricultural use, in order to assess whether irregular applications of agrochemicals may compromise reproductive success. In addition, environmental changes, whether natural or anthropogenic, can promote the emergence of pathogens in nestlings, weakening their immune system and facilitating infections by opportunistic yeasts and bacteria (Loiko et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Allgayer et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2009\u003c/span\u003eb), as well as by free-living microorganisms such as \u003cem\u003eChlamydophila psittaci\u003c/em\u003e (Raso et al. \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Raso et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) and \u003cem\u003eSalmonella spp\u003c/em\u003e. (Allgayer et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2009\u003c/span\u003ea), which are capable of causing severe diseases in wild populations.\u003c/p\u003e \u003cp\u003eGiven the multiple threat factors identified, the results of this review highlight the need for integrated conservation actions for Hyacinth Macaw populations in the Pantanal. Key priorities include strengthening public policies for the protection of \u003cem\u003ecap\u0026otilde;es\u003c/em\u003e and \u003cem\u003ecordilheiras\u003c/em\u003e, expanding protected areas in strategic regions, implementing integrated fire management supported by geotechnologies, monitoring environmental contamination, and assessing population health, all of which are essential to ensure the long-term viability of the species.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec15\" class=\"Section3\"\u003e \u003ch2\u003e3.3.2 Amazon\u003c/h2\u003e \u003cp\u003eAccording to the results presented above, the Amazon biome and the Hyacinth Macaw population occurring in this region remain the least studied compared to the other biomes. This scarcity of scientific information limits the understanding of local threats and constrains the development of targeted conservation strategies. Among the few studies addressing the Amazon biome, Devenish et al. (\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) reported an apparent geographic expansion of the Hyacinth Macaw in the region. This expansion has been directly associated with landscape changes driven by deforestation, particularly the conversion of forested areas into savanna-like environments. Although such changes may initially favour the species, which in this region shows a preference for nesting in more open habitats, continued conversion of native vegetation into pastureland is likely to reverse this trend, leading to habitat loss and a reduction in the species\u0026rsquo; area of occupancy. Therefore, the apparent expansion of the Hyacinth Macaw\u0026rsquo;s range in the Amazon should not be interpreted as a positive conservation signal, but rather as a consequence of ongoing land-use change and deforestation in the biome.\u003c/p\u003e \u003cp\u003eThus, further studies focusing on the demography of the Hyacinth Macaw in the Amazon are needed to estimate current population size and to support the development of effective strategies aimed at ensuring the species\u0026rsquo; long-term viability in the region. In addition, awareness-raising initiatives involving local communities are essential, particularly those that highlight the species\u0026rsquo; vulnerability and its ecological and economic relevance. A successful example is provided by Presti et al. (\u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), who implemented environmental education activities in schools in the Caraj\u0026aacute;s region, Par\u0026aacute;, using educational games to promote student engagement in Hyacinth Macaw conservation.\u003c/p\u003e \u003cp\u003eThe scarcity of studies on the Amazonian population may be partly explained by the difficulty of access to these areas, as well as by the high logistical and financial costs involved, which pose major challenges for research. Nevertheless, the development of new studies in the region is essential to improve understanding of the species\u0026rsquo; ecological requirements and the local threats faced by this population. It is noteworthy that the \u003cem\u003eArara Azul\u003c/em\u003e Institute is currently conducting continuous monitoring in the region, which is expected to substantially advance knowledge of the Amazonian population. Even so, the results of this review indicate that the Amazon biome still lacks fundamental information, underscoring the urgent need to expand research and conservation efforts to ensure the long-term persistence of the Hyacinth Macaw and the conservation of this biome in Brazil.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec16\" class=\"Section3\"\u003e \u003ch2\u003e3.3.3 Cerrado\u003c/h2\u003e \u003cp\u003eAccording to the results of this review, the population in the Central region, within the Cerrado biome, has been the subject of relatively few studies, which limits the availability of information on the threats faced by this population. Among the studies addressing the Cerrado, Borges et al. (\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) stand out for highlighting the high vulnerability of the Hyacinth Macaw to environmental changes in this biome. These authors reported that only 14% of the species\u0026rsquo; range in the Cerrado is located within Protected Areas, exposing the population to the expansion of the agricultural frontier and increasing habitat fragmentation. Under these conditions, actions such as systematic monitoring of the Central population, the establishment of new Protected Areas, and the implementation of ecological corridors are essential to facilitate individual movement and to mitigate the effects of land-use change and projected climate shifts during this century.\u003c/p\u003e \u003cp\u003eAlthough the expansion of the agricultural frontier represents one of the main threats to the Central population through the loss of feeding and nesting areas, illegal wildlife trade also constitutes a major driver of population decline. According to Faria et al. (\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2008\u003c/span\u003e) and Presti et al. (\u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2015\u003c/span\u003e), most Hyacinth Macaws seized during anti-trafficking operations originated from this population. This vulnerability is closely associated with the socioeconomic context of northeastern Brazil, which presents some of the lowest human development indices in the country. In areas characterized by low income and limited access to information, wildlife trade is often perceived as an economic alternative rather than as an illegal activity, being viewed as a means of exploiting what is considered an abundant natural resource (Herrera and Hennessey \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Pires \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Clarke and By \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). In addition, the predatory methods used to capture eggs and nestlings further exacerbate conservation problems, as hunters frequently cut down nesting trees, thereby reducing the availability of natural cavities (Berkunsky et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Bonaparte et al. 2024).\u003c/p\u003e \u003cp\u003eThus, the conservation measures adopted by the \u003cem\u003eArara Azul\u003c/em\u003e Project in the Pantanal and the Amazon, such as raising awareness among local communities about the species\u0026rsquo; importance and vulnerability, establishing partnerships with landowners to protect nesting sites, and promoting tourism as a source of income for rural families, could also be implemented for this population. These actions have the potential to reduce illegal trade and to enhance local engagement in the conservation of the Hyacinth Macaw (Corr\u0026ecirc;a and Guedes \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Mercado et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eAnother emerging factor that deserves attention is the use of social media platforms for illegal wildlife trade, which has expanded across several countries in the Americas. These platforms facilitate illegal transactions and may partially replace physical markets following their closure, allowing wildlife trafficking to persist within national borders (Pires et al. \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Picazo et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). However, studies in Brazil examining the role of social media in Hyacinth Macaw trafficking remain scarce, representing an important gap for future research.\u003c/p\u003e \u003cp\u003eLastly, it is important to note that although the Hyacinth Macaw has been removed from Brazil\u0026rsquo;s official list of threatened species, this reclassification may result in less stringent enforcement and weaker penalties for illegal trade. Such a scenario could increase harvesting pressure, particularly in regions with limited monitoring and high socioeconomic vulnerability. Accordingly, the results of this review highlight the need for further studies focused on the Central population, with particular emphasis on socio-environmental threats, feeding and reproductive ecology, and local community perceptions of the species. Integrated approaches that consider the socio-ecological systems of the region are essential to develop effective conservation strategies, reduce illegal trade, and ensure the long-term viability of the Hyacinth Macaw in the Cerrado.\u003c/p\u003e \u003c/div\u003e \u003c/div\u003e \u003cdiv id=\"Sec17\" class=\"Section2\"\u003e \u003ch2\u003e3.4 Limitations of the review\u003c/h2\u003e \u003cp\u003eAlthough this review was carefully developed, some limitations should be acknowledged. First, the distribution of available information on Hyacinth Macaw populations in Brazil remains uneven, with a particularly low number of studies focusing on populations in the Central and Northern regions of the country. This scarcity of information limits the understanding of ecological requirements and regional threats, thereby constraining the development of conservation strategies aimed at ensuring the species\u0026rsquo; long-term viability. In addition, this review focused exclusively on peer-reviewed articles published in indexed journals, a criterion adopted to ensure methodological rigour and comparability among the studies analysed. Consequently, relevant information contained in theses, dissertations, book chapters, technical reports, conference abstracts, and citizen-science databases was not included, although such sources may provide valuable insights into the species. Acknowledging these limitations is essential to guide future research and to support more comprehensive and effective conservation strategies.\u003c/p\u003e \u003c/div\u003e"},{"header":"4. CONCLUSIONS","content":"\u003cp\u003eThis study represents one of the first reviews dedicated exclusively to the Hyacinth Macaw in Brazil, integrating information on its biology, distribution, and conservation, and providing a valuable foundation to guide future conservation strategies for the species. The results indicate a marked increase in the number of studies on the Hyacinth Macaw in Brazil between 1997 and 2024, particularly in the fields of ecology, conservation, and animal health. However, this scientific production has been largely concentrated in the Pantanal biome, especially in the Southern Pantanal, where long-term monitoring and management actions have been conducted by the \u003cem\u003eArara Azul\u003c/em\u003e Project. These efforts have substantially advanced knowledge of the species\u0026rsquo; threats, reproductive biology, genetics, and ecology, and have contributed to the observed recovery of Hyacinth Macaw populations in the country.\u003c/p\u003e \u003cp\u003eFurthermore, this review identified regional variation in threats, feeding and reproductive behaviour, and ecological interactions, indicating that conservation strategies should be tailored to the specific conditions of each biome. Despite the overall increase in scientific output, substantial knowledge gaps remain, particularly for populations in the Northern and Central regions of Brazil, where information on reproductive biology, local ecology, and population-specific threats is still limited. In addition, the small number of studies involving captive individuals represents a major constraint for the development of future reintroduction or restocking programmes in areas strongly affected by anthropogenic pressures. Consequently, it is essential to direct future research efforts toward these regions and to promote studies focused on the ex situ conservation of the species.\u003c/p\u003e \u003cp\u003eOverall, this review provides a consolidated framework to support future research, management actions, and the formulation of public policies aimed at conserving the Hyacinth Macaw in Brazil. By fostering effective and regionally adapted strategies that actively engage local communities through citizen science initiatives and environmental education, it is possible to mitigate the environmental threats that continue to compromise the integrity of the species\u0026rsquo; populations.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eCONFLICT OF INTEREST STATEMENT\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors have no competing interests.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eACKNOWLEDGEMENTS\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe are grateful to the members of the Hyacinth Macaw Institute for their generous assistance throughout this study, particularly Fernanda M. Fontoura, who provided invaluable insights. We also thank Alexandra Sanches and Eduardo Roberto Alexandrino for their careful evaluation of the manuscript and for their valuable comments and suggestions, which were essential to the development of this article.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFUNDING STATEMENT\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis research did not receive any specific grant from funding agencies in the public, commercial, or not-for-profit sectors.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAUTHOR CONTRIBUTIONS\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eBoth authors contributed to the study conception and design. Material preparation, data collection and analysis were performed by Vitor Matheus Morais de Oliveira. The first draft of the manuscript was written by Vitor Matheus Morais de Oliveira, and Vinicíus de Avelar São Pedro reviewed the manuscript and contributed to the development of the study structure. Both authors commented on previous versions of the manuscript. Both authors read and approved the final manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eDATA AVAILABILITY STATEMENT\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAll data supporting the findings of this study are available within the article and its supplementary material.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAllgayer MGA, Guedes NMR, Chiminazzo C, Cziulik M, Weimer TA (2009) Clinical Pathology And Parasitologic Evaluation Of Free-Living Nestlings Of The Hyacinth Macaw (\u003cem\u003eAnodorhynchus Hyacinthinus\u003c/em\u003e). 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Captive Parrots Anim Conserv 15:28\u0026ndash;43. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/j.1469-1795.2011.00477.x\u003c/span\u003e\u003cspan address=\"10.1111/j.1469-1795.2011.00477.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"ornithology-research","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"orni","sideBox":"Learn more about [Ornithology Research](https://link.springer.com/journal/43388)","snPcode":"43388","submissionUrl":"https://submission.nature.com/new-submission/43388/3","title":"Ornithology Research","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Psittacidae, animal conservation, bird ecology, extinction threats","lastPublishedDoi":"10.21203/rs.3.rs-8695459/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-8695459/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThe Hyacinth Macaw (\u003cem\u003eAnodorhynchus hyacinthinus\u003c/em\u003e), the world's largest psittacid is threatened by illegal trade and habitat loss. Conservation efforts in Brazil, particularly the \u003cem\u003eArara Azul\u003c/em\u003e Project, have contributed to the recovery of the Hyacinth Macaw population in the Pantanal. However, scientific knowledge on the species\u0026rsquo; conservation remains scattered across the literature, which hampers the development of effective long-term strategies. This study aimed to systematize and critically analyze scientific research on the Hyacinth Macaw in Brazil through an integrative review. Seventy one papers were retrieved from six academic databases. The results revealed a significant increase in research in the past three decades, with a predominance of empirical studies, particularly in ecology, most of them conducted in the southern Pantanal. The reviewed literature indicates that Hyacinth Macaw populations exhibit recent gene flow and relatively high genetic diversity compared with other threatened psittacid species. Additionally, interpopulation variations were identified in diet and reproductive behavior, depending on the region. Major knowledge gaps include the lack of studies on the species\u0026rsquo; ecology and behavior in the Central and Northern regions of the country, as well as the limited number of studies involving individuals in captivity. These gaps limit accurate demographic, genetic, and ecological assessments, as well as conservation planning. The findings highlight the need for region-specific approaches to conservation and emphasize the central role of the \u003cem\u003eArara Azul\u003c/em\u003e Institute in scientific research and population management. Despite significant progress, important gaps remain in understudied populations, requiring targeted conservation strategies and actions.\u003c/p\u003e","manuscriptTitle":"Conservation Challenges of the Hyacinth Macaw (Anodorhynchus hyacinthinus (Latham, 1790)) in Brazil: An Integrative Review","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-03-23 11:28:34","doi":"10.21203/rs.3.rs-8695459/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewersInvited","content":"","date":"2026-03-18T09:23:01+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2026-02-06T09:28:17+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2026-02-06T09:22:28+00:00","index":"","fulltext":""},{"type":"submitted","content":"Ornithology Research","date":"2026-01-26T01:08:22+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":"78ccca7c-9ee8-4b60-96ec-4439c59b0d34","owner":[],"postedDate":"March 23rd, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-03-23T11:28:35+00:00","versionOfRecord":[],"versionCreatedAt":"2026-03-23 11:28:34","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-8695459","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-8695459","identity":"rs-8695459","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}