Trophic niche overlap in stingless bees (Melipona Illiger, 1806) across season in Southern Brazil

Trophic niche overlap in stingless bees (Melipona Illiger, 1806) across season in Southern Brazil | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Trophic niche overlap in stingless bees (Melipona Illiger, 1806) across season in Southern Brazil Amanda Garçoa Raulik, João Marcelo Deliberador Miranda, Luiz Gabriel Ludwig, and 1 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8042458/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 5 You are reading this latest preprint version Abstract Understanding how stingless bee species share floral resources is essential for optimizing meliponiculture and supporting pollinator conservation. This study investigated the trophic niche overlap of four Melipona species ( M. quadrifasciata, M. bicolor, M. torrida , and M. mondury ) in Southern Brazil across warm and cold seasons. Pollen samples were collected monthly over ten months and analyzed to assess richness, trophic niche breadth, and overlap between species' trophic niches. Results showed seasonal variation in foraging patterns, with all species (except M. mondury ) reducing their niche breadth in colder months. Melipona mondury , a species native to warmer regions of the Atlantic Forest and lacking reproductive diapause, maintained a broad niche year-round, possibly increasing its vulnerability in colder climates. The highest overlap occurred between M. quadrifasciata and M. mondury during the cold season, raising concerns about managing these species in consorted meliponiculture systems. The most frequently visited botanical families included Myrtaceae, Arecaceae, Asteraceae, Moraceae, and Fabaceae highlighting their importance as key floral resources. These findings highlight the importance of considering seasonal dynamics and species-specific traits in meliponiculture, including the need for targeted supplemental feeding and resource planning. Promoting key botanical resources such as Myrtaceae and Arecaceae can support sustainable colony development in high-density environments. Meliponiculture Overlap niche Pollination Seasonality Trophic ecology Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction The Neotropical region hosts the richest bee fauna in the world, and bees play a vital role in maintaining plant biodiversity (Moure 2007). Among them, stingless bees of the tribe Meliponini stand out for their high diversity, abundance, and their crucial role in plant pollination in the Neotropics (Nates-Parra et al. 2008 ). Currently, there are 538 known species of stingless bees distributed across tropical and subtropical regions worldwide (Ascher & Pickering 2022). Within this group, bees of the genus Melipona are among the most commonly managed by beekeepers due to their larger body size and easy of handling (Jaffé et al. 2015 ). Several species also play an important role in the local economy, as farmers commercialize their hives and bee products such as honey, pollen, wax, and propolis (Reyes-González et al. 2016; Gemim & De Melo Silva 2017). The loss and degradation of Atlantic Forest habitats are major threats to stingless bees (Roberts et al. 2017 ). However, meliponiculture - the rational breeding of stingless bees - has gained popularity among small-scale producers and may represent a promising strategy for species conservation (Nogueira-Neto et al. 1986 ). The survival of a bee colony depends on the success of foragers in gathering food resources such as nectar and pollen. Pollen collected by stingless bees is particularly informative for studying their trophic ecology (Roubik & Patiño 2018). Although most stingless bees are generalist foragers, even species with more specialized habits trend to exploit a wide range of plants throughout the year (Bueno et al. 2023 ). Given the high diversity of stingless bee species coexisting in tropical ecosystems, trophic niche overlap becomes an important factor to consider. In meliponaries, the artificial increase in colony density and the number of individuals foraging in the same area intensifies this overlap, highlighting the need to investigate how these species share floral resources. This study aimed to evaluate seasonal trophic niche overlap among four stingless bee species of the genus Melipona ( Melipona quadrifasciata , M. bicolor , M. torrida , and M. mondury ), which are commonly kept together in meliponaries. These species differ in body size and coloration. Specifically, we sought to answer the following questions: (1) How do the floral resource choices of M. quadrifasciata , M. bicolor , M. torrida , and M. mondury vary between warm (spring and summer) and cold (autumn and winter) seasons? (2) Do body size, coloration, and phylogenetic distance influence trophic niche overlap among these species? Methods Study area The study was carried out at the "Meliponário Abelhas da Mata, located in the municipality of Guarapuava - Paraná - Brasil (25°22’23,69”S; 51°26’17,967”W, 1.119 m altitude) in a peri-urban region of the city. The region is located in a transitional area between Araucaria Moist Forest (Mixed Ombrophilous Forest) and high-altitude grasslands, originally characterized by natural fields interspersed with patches of Atlantic Forest. The climate is classified as humid subtropical and mesothermal, with an average annual temperature of 16.8°C (Costa & Andrade 2017 ), corresponding to the Cfb type in the Köppen–Geiger climate classification. The southern region of Brazil does not have a well-defined rainy season, having a uniform distribution of precipitation throughout the year. Stingless bee species selected The four species chosen for this study are endemic to the Atlantic Forest (Nieh 2003; Camargo et al. 2013 ; Lopes et al. 2022), commonly bred together in meliponaries (Nogueira-Neto 1997 ) and were housed in boxes of wood for over a year at the meliponary. These target species were: Melipona quadrifasciata quadrifasciata Lepeletier, 1836; Melipona bicolor schencki Gribodo, 1896; Melipona torrida Friese, 1916; and Melipona mondury Smith, 1893. The selection of these stingless bee species was based on two criteria: 1) their common use in consorted meliponiculture, and 2) the differences in foraging activities, reflected in their flight ranges and the resource selection. Although the four species are considered generalists, we evaluated three aspects such as size and color of the body and phylogenetic distance between them. Stingless bee monitoring and sample processing The study region does not have a well-defined dry season, with precipitation being evenly distributed throughout the year (Pereira et al. 2008 ). Therefore, the data collection was organized according to warm (spring–summer) and cold (autumn–winter) seasons. Field sampling was conducted monthly over a ten-month period (from November 2022 to August 2023), ensuring temporal coverage of both warm and cold seasons. Each sampling event consisted of standardized collections performed under suitable weather conditions (temperatures between 18–20 ºC and absence of rainfall), when stingless bees are known to be most active (Gianinni & Imperatriz-Fonseca 1986; Hilário et al. 2000 ). For each of the four focal species, pollen was collected from five foraging individuals originating from five distinct colonies, totaling 20 samples per month. Sampling effort was evenly distributed across the study period, resulting in 200 pollen samples in total. While most collections could be completed within a single field day, during colder months additional sampling time was occasionally required to ensure sufficient representation of all species. To obtain the pollen samples used by stingless bees, the entrances were blocked with small sticks, preventing forage bees from entering. As soon as the foraging bees approached and tried to enter the hive, only those that were carrying pollen on their corbiculae were selected and captured. This capture was done manually or with the help of a small plastic container with a lid. The collected pollen was stored in microtubes (1 ml) (Eppendorf type), containing AFA solution (5% acetic acid, 5% formaldehyde and 90% of 50% alcohol), and the bees were released to return to their activities. Palynological analysis The samples underwent the acetolysis process described by Erdtman ( 1952 ) with some modifications. Microscopes slides were prepared using glycerinated gelatin (Johansen 1940) and examined under a bright field microscopy to identify pollen types to the lowest possible taxonomic level. Taxons were identified to the lowest taxonomic level possible, in most cases to families and, occasionally, to different genera or morphotypes within the same family (Appendix). The pollen identification was conducted with the assistance of a botanic specialist and virtual platforms such as Online Pollen Catalog Network (RCPol) and Global Pollen Project (Martin & Harvey 2017 ). Diagnostic features considered included the size, aperture type, and surface characteristics of the pollen exine. Morphological characterization of stingless bees Ten individuals from each of the four stingless bee species were collected at the entrance of their nests. Each specimen was euthanized in a killing chamber using ethyl acetate. Each specimen was measured for its mass (mg) using a precision scale. The average weight of the 10 individuals from each species was used as a proxy for body size. Regarding coloration, one representative specimen per species was analyzed and classified based on the percentage of the body surface (dorsal view) displaying light and dark colors, using the R packages magick and imager. This analysis was intended to provide a representative overview of interspecific differences in coloration patterns. Table 1 – Morphological character matrix of the four analyzed stingless bee species ( Melipona quadrifasciata , M. bicolor , M. torrida , and M. mondury ). Bee species Average weight (mg) Dark coloration (%) Light coloration (%) M. quadrifasciata 62,1 ± 4,04 87,21 12,79 M. bicolor 51,1 ± 4,43 90,21 9,78 M. torrida 27,2 ± 2,44 86,51 13,49 M. mondury 60,6 ± 8,49 2,55 97,45 Phylogenetic distance To calculate the evolutionary or phylogenetic distance among the four stingless bee species, the molecular phylogeny of Melipona proposed by Ramírez et al. ( 2010 ) was used. To reconstruct the phylogeny and the distances between each taxon, the software FigTree® was used to generate an ultrametric phylogenetic tree. To update the nomenclature and evolutionary relationships, substitute taxa were used for the species in the present study: Melipona quadrifasciata anthidioides was used as a substitute for M. quadrifasciata quadrifasciata ; M. rufiventris rufiventris as a substitute for M. mondury ; and M. marginata as a substitute for M. torrida . The criteria for these substitutions were as follows: the possible evolutionary proximity between the subspecies of M. quadrifasciata (Batalha-Filho et al. 2010 ); the possible evolutionary proximity between M. marginata and M. torrida , since the latter replaced the name previously given to a former subspecies of M. marginata , M. marginata obscurior (Melo 2013 ); and the possible evolutionary proximity between M. rufiventris and M. mondury , since the latter was previously considered a subspecies of the former (Melo 2003). The phylogenetic distances between species pairs were calculated based on the number of evolutionary steps between each species pair. Data analysis For each stingless bee species, both pollen richness (i.e., the number of pollen morphotypes) and feeding niche breadth were calculated. Niche breadth was calculated using Levin’s index (B), with the standardized value indicated as BA. Based on the relative abundances of pollen morphotypes, trophic niche overlap among the four stingless bee species was analyzed. Niche overlap for Warm and Cold Season was assessed using the Pianka Index and a Non-Metric Multidimensional Scaling (NMDS) perceptual map. A permutation test with 1,000 randomizations was applied to evaluate whether the observed niche overlaps were significantly higher than expected by chance. All analyses were conducted in R using the vegan package (v. 2.5-5; Oksanen et al. 2019). Based on the size and color data of the four stingless bee species (Table 1 ), two distance matrices were constructed: one for size distance and one for color distance, both based on Euclidean Distance. Additionally, from the phylogeny of the genus Melipona (Ramírez et al. 2010 ), a phylogenetic distance matrix was constructed for the four stingless bee species. To test which variables best explain the feeding niche overlap among the four stingless bee species, a Generalized Linear Model (GLM) approach was used. The response variable was the trophic niche overlap values, and the predictor variables were Size Distance (DSize), Color Distance (DColor), and Phylogenetic Distance (DPhyl). To avoid the effect of collinearity among predictors in the GLM, a Variance Inflation Factor (VIF) analysis was conducted using the “fmsb” 0.7.3 package. In this analysis, predictor variables with VIF values greater than 3 are considered collinear and should be removed from subsequent analyses. Since all VIF values were below 3 (Table 2 ), all predictor variables were retained for model construction. Table 2 Variance inflation factors for the predictor variables of Trophic Niche Overlap among four stingless bee species from the Brazilian Atlantic Forest ( Melipona quadrifasciata , M. bicolor, M. torrida , and M. mondury ). GVIF = Generalized Variance Inflation Factor; DSize = Size distance among the four species (Euclidean Distance); DColor = Color distance among the four species (Euclidean Distance); and DPhyl = Phylogenetic distance among the four species. Variable GVIF DSize 1,1602 DColor 2,1148 DPhyl 2,0239 Eight models were constructed, including all predictor variables (DSize, DColor, and Phyl), all possible combinations among them, a global model (with all variables), and a null model (with none). The best model was selected based on the Akaike Information Criterion (AIC) and the AIC model weights. In this analysis, the lower the AIC value and the higher the AIC weight, the better the model fit for predicting the response variable (trophic niche overlap). When the difference between the best model and other models (ΔAIC) was less than 2, those models were considered equally predictive. In such cases, we performed model averaging to combine their estimates and account for model selection uncertainty. These analyses were performed using the “AICcmodavgpa” 2.3.2 package (Mazerolle 2023 ). The GLM analyses were carried out in the R environment (R Core Team, 2013). Results Melipona bicolor exhibited the highest richness of pollen morphotypes used (S = 18), followed closely by M. torrida (S = 17), while M. mondury showed the lowest richness (S = 13). In terms of trophic niche breadth, M. quadrifasciata presented the highest value (B = 10.1), whereas M. mondury had the lowest (B = 3.5) during the Warm Season (Table 3 ). The most represented plant family in the pollen samples was Myrtaceae, which dominated the diets of all species. Other frequent families included Arecaceae, Fabaceae, and Moraceae. Notably, Asteraceae appeared only during the Warm Season and was consumed exclusively by M. quadrifasciata and M. torrida (Fig. 2 ). Table 3 – Breadth of trophic niches (Levin’s index: B and standardized BA), richness of visited plant morphotypes (S), and pairwise niche overlap values (Pianka index) among stingless bee species raised in meliponaries in southern Brazil (Warm season). P-values are shown only for comparisons with statistically significant differences (based on permutation tests with 1,000 randomizations). B BA S Niche overlap (Warm season) M. quadrifasciata M. torrida M. bicolor M. mondury M. quadrifasciata 10.1 0.36 16 - 0.77 (p = 0,004) 0.90 (p = 0,001) 0.70 (p = 0,006) M. torrida 9.2 0.32 15 0.77 - 0.85 (p = 0,003) 0.86 (p = 0,001) M. bicolor 8.3 0.29 17 0.90 0.85 - 0.82 (p = 0,001) M. mondury 3.5 0.10 9 0.70 0.86 0.82 - Table 4 – Breadth of trophic niches (Levin’s index: B and standardized BA), richness of visited plant morphotypes (S), and pairwise niche overlap values (Pianka index) among stingless bee species raised in meliponaries in southern Brazil (Cold season). P-values are shown only for comparisons with statistically significant differences (based on permutation tests with 1,000 randomizations). B BA S Niche overlap (Cold season) M. quadrifasciata M. torrida M. bicolor M. mondury M. quadrifasciata 5.4 0.49 8 - 0.59 0.87 (p = 0,017) 0.86 (p = 0,026) M. torrida 4.4 0.37 8 0.59 - 0.61 0.81 M. bicolor 5 0.45 6 0.87 0.61 - 0.71 M. mondury 5 0.44 9 0.86 0.81 0.71 - Seasonal variation Throughout the year, M. mondury collected significantly fewer pollen morphotypes than M. quadrifasciata, M. bicolor , and M. torrida . Similarly, M. mondury also exhibited lower trophic niche breadth compared to the other species (Fig. 3 A). This same pattern was observed during the Warm Season, but during the Cold Season there were no differences in either the number of pollen morphotypes or the niche breadths among the species (Tables 3 and 4 ). In both seasons, the highest trophic niche overlap was observed between M. quadrifasciat a and M. bicolor (91% in the Warm Season and 87% in the Cold Season), indicating resource sharing between these species. During the Warm Season, all species pairs exhibited significantly higher niche overlap than expected by chance (p < 0.01 for all pairs), with particularly high values between M. torrida and M. mondury (86%, p = 0.001), and between M. bicolor and M. torrida (85%, p = 0.003). The lowest overlap occurred between M. quadrifasciata and M. mondury (69%, p = 0.006). In contrast, during the Cold Season, only two species pairs exhibited significant niche overlap: M. quadrifasciata with M. bicolor (87%, p = 0.017) and with M. mondury (87%, p = 0.026). Other pairs, such as M. quadrifasciata and M. torrida (60%, p = 0.27), and M. torrida and M. bicolor (61%, p = 0.22), did not show significant overlap, suggesting more distinct foraging patterns in those interactions during this period (Fig. 4 ). Overall, in the Cold Season, stingless bee species reduced both the number of pollen morphotypes collected and their trophic niche breadth compared to the Warm Season. However, M. mondury did not follow this same trend (Fig. 3 C). Models 4, 5, and 6 were equally predictive in explaining trophic niche overlap among the stingless bee species. Even after model averaging, the GLM analysis did not support any significant relationships between the predictive variables and trophic niche overlap. Color, body size, and phylogenetic distance were not significantly associated with variations in trophic niche overlap when considering all samples combined (Tables 5 and 6 ), as well as in the separate analyses for the warm and cold seasons. Table 5 – Generalized Linear Models (GLM) evaluated for their predictive power of Trophic Niche Overlap in four stingless bee species from the Brazilian Atlantic Forest ( Melipona quadrifasciata , M. bicolor , M. torrida , and M. mondury ). AICc = Akaike Information Criterion corrected for small sample sizes; ΔAICc = Difference in AICc relative to the best model; AICcWt = AICc model weight. Model AICc ΔAICc AICcWt 5 -2,17 0 0,41 4 -1,94 0,22 0,37 6 -0,92 1,25 0,22 Null 5,95 8,12 0,01 2 25,47 27,64 0 1 25,53 27,70 0 3 27,83 30 0 Global > 1.000 > 1.000 0 Table 6 – Coefficient of the predictor variable (DCor) for the best-fitted Generalized Linear Model (GLM) predicting Trophic Niche Overlap in four stingless bee species from the Brazilian Atlantic Forest ( Melipona quadrifasciata, M. bicolor, M. torrida , and M. mondury). DPhyl = Phylogenetic distance; DSize = Size distance; DColor = Color distance; GLM = Generalized Linear Models; TNO = Trophic Niche Overlap. Coefficient Standad error t-value p-value Intercept -0, 4822371 0,1209295 2,988 0,00281* DPhyl -0,0512033 0,0856634 0,489 0,62468 DSize -1,5423716 2,8637592 0,439 0,66067 DColor -0,0001646 0,0005282 0,242 0,80893 Discussion Climatic factors are intrinsically linked to the availability and density of resources for stingless bees (Keppner & Jarau 2016 ; Moura et al. 2022 ). Most studies evaluating the effects of seasonality on niche overlap in stingless bees analyze similarities and differences between the rainy and dry seasons (Brito et al. 2012 ; Santos et al. 2013 ; Escobedo-Kenefic et al. 2020 ; Ferreira et al. 2023 ). The results presented here demonstrate that stingless bees follow the pattern observed in previous studies, foraging more intensively for pollen resources during the warm season, due to increased floral abundance and higher temperatures (Souza et al. 2006 ; Ferreira-Junior et al. 2010 ). In contrast, during the cold season, a reduction in species richness and trophic niche breadth was observed in three of the studied species — Melipona quadrifasciata, M. bicolor , and M. torrida — all of which conformed to this pattern. However, M. mondury deviated from it, maintaining similar niche breadth values across both seasons, suggesting a distinct foraging strategy. This behavior may be related to its biogeographic origin: although native to the Atlantic Forest, M. mondury is not originally found in the study region, being more commonly distributed in lower-altitude areas of the Atlantic Forest along the Brazilian coast, where the climate is warmer and more humid (Mouga 2009 ; Mouga 2015 ; Mouga & Nogueira-Neto 2015). Furthermore, this species does not undergo reproductive diapause, which sustains its year-round demand for pollen and may increase its vulnerability in subtropical climates with pronounced winters. The increase in niche overlap between M. quadrifasciata and M. mondury during cold season may lead to challenges in colony development due to the reduced availability of floral resources during this period. These findings have important implications for meliponiculture, particularly regarding colony maintenance during colder periods. Supplemental feeding, including sugar syrup and pollen-based food, may be essential to support the survival and proper development of colonies maintained outside their natural range. Some botanical taxa form the basis of stingless bees’ diet (e.g., Myrtaceae, Arecaceae), while others are used more occasionally, which helps explain the trophic relationships observed in the networks of both warm and cold seasons. During the cold season, the availability of flowering plants decreases, which strengthens the bee–plant interactions (Ferreira et al. 2023 ). Additionally, several taxa recorded correspond to herbaceous species likely native to grasslands or forest edges (i.e. Asteraceae, Begoniaceae, Labiateae, Urticaceae, Verbenaceae) reflecting the transitional nature of the study area between Araucaria Moist Forest (Mixed Ombrophilous Forest) and high-altitude grasslands. The concept of a key resource can be understood as a food resource capable of determining the survival potential of organisms. Additionally, it can influence the composition of biological communities, as its absence or low availability may affect species’ population density, long-term viability, or even lead to local extinction (Primack 2014 ). Stingless bees exhibit a certain affinity for resources considered key to colony maintenance. The collection of morphotypes from botanical families that flower abundantly throughout the year, such as Myrtaceae, facilitates species coexistence in environments like meliponaries, where there is an artificially increased local density of bees. Identifying key resources may therefore serve as an important tool for effective long-term conservation and management strategies. Bees of the genus Melipona exhibit social behavior and live in populous, perennial colonies, requiring them to exploit a wide range of floral resources throughout the year. Most flowering plants are ephemeral sources, blooming for only a few days (Ramalho et al. 2007 ). Therefore, a generalist foraging strategy is considered the norm among eusocial bees of the family Apidae. Their foraging patterns vary seasonally in response to environmental factors such as flowering periods — with summer and spring being seasons of abundant floral availability (Winston 2003 ). Melipona bees have been recorded visiting a variety of botanical families, depending on the surrounding vegetation (Bueno et al. 2023 ). Although stingless bees forage on many species from different plant families, some families are more frequently utilized. Floristic surveys conducted in the Araucaria Moist Forest have identified the Myrtaceae family as one of the richest or most abundant (Cordeiro & Rodrigues 2007 ; Albuquerque et al. 2011 ; Vieira & May 2020 ) with about 246 species belonging to 20 genera recorded solely within the Atlantic Forest domain, and 220 species belonging to 16 genera recorded in Paraná state alone (Flora e Funga do Brasil 2025). In the Atlantic Forest, Meliponini bees frequently visit flowers of Myrcia tomentosa , particularly species of Melipona and Plebeia (Ramalho 2004 ). In São Paulo, Melipona mondury (previously cited as M. rufiventris ) has been identified as one of the most effective pollinators of Myrtaceae species (Fidalgo 2002 ). Other studies have consistently reported Myrtaceae as one of the most visited plant families by bees in Brazil (Carvalho et al. 2001 ; Antonini et al. 2006 ; Ramalho et al. 2007 ), with Melipona species being the most frequent pollinators of Myrtaceae in the country. This is largely attributed to the presence of nectaries in these plants, which offer nectar in addition to pollen (Antonini et al. 2006 ; Gressler et al. 2006 ). The Arecaceae family exhibits asynchronous flowering, providing floral resources during several months of the year. In Brazil, approximately 270 species and 38 native genera are recorded, standing out for their abundance and species richness in Neotropical forests (Lara et al. 2017 ). In the state of Paraná, 23 Arecaceae species have been reported (Flora do Brasil 2020 ). In Paraná state, 17 species from 11 genera of Arecaceae have been recorded. The importance of Arecaceae in the pollen diet of Melipona bees has already been highlighted by Oliveira et al. (2009). The Araucariaceae family, in turn, was recorded exclusively in the samples of M. mondury and is noteworthy since these plants do not produce flowers, yet their pollen is protein-rich and nutritionally valuable. Other studies have also reported the use of anemophilous plants (e.g., Cecropiaceae) by stingless bees in the Atlantic Forest (Morgado et al. 2011 ; Braga et al. 2014 ). The trophic niche breadth of the four stingless bee species endemic to the Atlantic Forest analyzed in this study reveals a generalist feeding behavior, which is common among eusocial bees and reflects their plasticity in the use of pollen resources. Body size and coloration are important traits in the trophic niche segregation of stingless bees (Hilário et al. 2001 ; Pereboom & Biesmeijer 2003; Teixeira & Campos 2005 ; Biesmeijer & Slaa 2006). However, these traits were not effective in predicting the niche overlap observed in this study. This may be due to the fact that the analysis included only four species from the same genus, Melipona . Although the Melipona genus is highly diverse, the selected species are among the largest stingless bees and show little variation in body size, as well as a close evolutionary relationship. Despite morphological differences, this variation could not be linked to the patterns of niche overlap among the analyzed species. However, increasing the sampling effort to include species from other genera, with greater variation in body size, coloration, and phylogenetic distance, may help to better understand the dietary differences among these bees. The results obtained in this study can directly support the management and strategic organization of meliponaries located in the Atlantic Forest ecosystem, particularly in colder regions such as the Araucaria Moist Forest. The identification of key resources, such as plant species from the Myrtaceae and Arecaceae families, which proved highly attractive to the studied bee species, suggests that cultivating these plants may serve as an important nutritional support strategy throughout the year. Moreover, data on niche overlap indicate that the coexistence of certain species may require special attention. For example, M. quadrifasciata and M. mondury showed a marked increase in niche overlap during the cold season, suggesting that keeping both species in the same meliponary should be approached with caution. If this combination is adopted, it is recommended to increase the availability of supplemental feeding for both species. It is also noteworthy that M. mondury exhibited a distinct foraging strategy, maintaining a wide niche breadth even in the cold season, likely due to the absence of reproductive diapause. This trait may render the species more vulnerable in colder climates, as it must maintain a high demand for pollen even when floral resources are scarce, which could compromise colony viability without adequate support. Declarations Ethical Approval Not applicable. Funding This research was supported by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES – Brazil) . Proccess number 88887.649588/2021-00. Authors' contributions Amanda Garçoa Raulik conducted the fieldwork, performed data analyses, wrote, and revised the manuscript. Luiz Gabriel Ludwig assisted with the statistical analyses. Adriano contributed to the palynological analyses. João critically revised the manuscript. Conflicts of interest/Competing interests The authors declare that they have no conflicts of interest. References Albuquerque JMD, Watzlawick LF, Mesquita NSD (2011) Efeitos do uso em sistema faxinal na florística e estrutura em duas áreas da Floresta Ombrófila Mista no munícipio de Rebouças. 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Revista de Biología Trop 56(3):1295–1308 Nieh JC et al (2003) Effect of food location and quality on recruitment sounds and success in two stingless bees, Melipona mandacaia and Melipona bicolor . Behav Ecol Sociobiol 55:87–94 Nogueira-Neto P (1997) Vida e criação de abelhas indígenas sem ferrão. Nogueirapis, São Paulo Nogueira-Neto P, Imperatriz-Fonseca VL, Kleinert ADMP, Viana BF, Castro MD (1986) Biologia e manejo de abelhas sem ferrão. Online Pollen Catalog Network – RCPol. http://chaves.rcpol.org.br/ Ooms J (2024) magick: Advanced Graphics and Image-Processing in R. R package version 2.8.5. https://CRAN.R-project.org/package=magick Pereira LMP et al (2008) Determinação do início e término da estação chuvosa no Estado do Paraná. Revista Geografar 3(2):1–12 Primack RB (2014) Essentials of conservation biology, 6th edn. Sinauer Associates Ramalho M (2004) Stingless bees and mass flowering trees in the canopy of Atlantic Forest: a tight relationship. Acta Bot Brasilica 18:37–47 Ramalho M, Silva MD, Carvalho CA (2007) Dinâmica de uso de fontes de pólen por Melipona scutellaris Latreille (Hymenoptera: Apidae): uma análise comparativa com Apis mellifera L. (Hymenoptera: Apidae), no Domínio Tropical Atlântico. Neotrop Entomol 36:38–45 Ramírez SR et al (2010) A molecular phylogeny of the stingless bee genus Melipona (Hymenoptera: Apidae). Mol Phylogenet Evol 56:519–525 Roberts HP, King DI, Milam J (2017) Factors affecting bee communities in forest openings and adjacent mature forest. For Ecol Manag 394:111–122 Roubik DW, Moreno Patiño JE (2018) The stingless honey bees (Apidae, Apinae: Meliponini) in Panama and pollination ecology from pollen analysis. Pot-Pollen in Stingless Bee Melittology. Springer Santos M et al (2013) Overlap in trophic and temporal niches in the flower-visiting bee guild (Hymenoptera, Apoidea) of a tropical dry forest. Apidologie 44:64–74 Silva JM, Oliveira RT (2009) A importância das abelhas na polinização de culturas agrícolas. Acta Biológica Catarinense 1(1):45–52 Souza BA, Carvalho CAL, Alves RMO (2006) Flight activity of Melipona asilvai moure (Hymenoptera: Apidae). Brazilian J Biology 66:731–737 Teixeira LV, Campos FNM (2005) Início da atividade de vôo em abelhas sem ferrão (Hymenoptera, Apidae): influência do tamanho da abelha e da temperatura ambiente. Revista Brasileira de Zoociências 7:195–202 Vieira NC, May D (2020) Regeneração natural de Myrtaceae em uma unidade de conservação urbana, Curitiba, PR. Brasil REVSBAU 15(1):39–49 Winston ML (2003) A Biologia da Abelha. Magister, Porto Alegre Supplementary Files Appendix.docx Cite Share Download PDF Status: Under Review Version 1 posted Reviewers agreed at journal 22 Dec, 2025 Reviewers invited by journal 24 Nov, 2025 Editor invited by journal 13 Nov, 2025 Editor assigned by journal 11 Nov, 2025 First submitted to journal 10 Nov, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. 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1","display":"","copyAsset":false,"role":"figure","size":166064,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eLocation of the study area in the municipality of Guarapuava, state of Paraná, southern Brazil, where the data were collected. The marked point represents the sample collection site. Base map: IBGE (2024). Reference system: SIRGAS 2000 / UTM Zone 22S (EPSG: 4974).\u003c/strong\u003e\u003c/p\u003e","description":"","filename":"floatimage1.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-8042458/v1/ec2b558d8f9e8b661ab65bd7.jpeg"},{"id":96976024,"identity":"d172886e-14ad-4d84-b282-2abf5d5ff4e8","added_by":"auto","created_at":"2025-11-28 08:34:42","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":62123,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eRelative abundance of pollen from the five most representative plant families found in the diet of four stingless bee species (\u003c/strong\u003e\u003cem\u003e\u003cstrong\u003eMelipona quadrifasciata, M. torrida, M. bicolor\u003c/strong\u003e\u003c/em\u003e\u003cstrong\u003e, and \u003c/strong\u003e\u003cem\u003e\u003cstrong\u003eM. mondury\u003c/strong\u003e\u003c/em\u003e\u003cstrong\u003e) during the Cold and Warm Seasons.\u003c/strong\u003e\u003c/p\u003e","description":"","filename":"floatimage2.png","url":"https://assets-eu.researchsquare.com/files/rs-8042458/v1/0c6ea48600cbd67482210282.png"},{"id":96976031,"identity":"4353db6f-aadf-42f0-a631-e580f216805f","added_by":"auto","created_at":"2025-11-28 08:34:42","extension":"jpeg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":612328,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eA. Trophic niche breadth (Levins index), B. Trophic niche overlap (Pianka index) between pairs of stingless bee species in Warm and Cold Season and C. Richness of pollen morphotypes per month throughout the study.\u003c/strong\u003e\u003c/p\u003e","description":"","filename":"floatimage3.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-8042458/v1/dc7c3ed06ab4d72a636a39cf.jpeg"},{"id":97138019,"identity":"e3fdd846-5611-45e9-a2c1-754c3d94d770","added_by":"auto","created_at":"2025-12-01 09:58:25","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":102821,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eTrophic niche overlap of the four stingless bee species in the Warm Season (red label) and the Cold Season (blue label), based on non-metric multidimensional scaling (NMDS) analysis using the Pianka similarity index.\u003c/strong\u003e\u003c/p\u003e","description":"","filename":"4.png","url":"https://assets-eu.researchsquare.com/files/rs-8042458/v1/32f80eb8188740af583d6c70.png"},{"id":97144773,"identity":"0d56ca26-ee5e-4b85-ab69-3e9cb008f24d","added_by":"auto","created_at":"2025-12-01 10:11:58","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2152031,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8042458/v1/9e88b4c0-b8eb-4796-ad0a-7080e7efc935.pdf"},{"id":96976021,"identity":"05f8d7f3-f1a1-4a08-9e2c-893a13956f8c","added_by":"auto","created_at":"2025-11-28 08:34:42","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":19865,"visible":true,"origin":"","legend":"","description":"","filename":"Appendix.docx","url":"https://assets-eu.researchsquare.com/files/rs-8042458/v1/2c019980f9c8665017e3ce51.docx"}],"financialInterests":"","formattedTitle":"Trophic niche overlap in stingless bees (Melipona Illiger, 1806) across season in Southern Brazil","fulltext":[{"header":"Introduction","content":"\u003cp\u003eThe Neotropical region hosts the richest bee fauna in the world, and bees play a vital role in maintaining plant biodiversity (Moure 2007). Among them, stingless bees of the tribe Meliponini stand out for their high diversity, abundance, and their crucial role in plant pollination in the Neotropics (Nates-Parra et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Currently, there are 538 known species of stingless bees distributed across tropical and subtropical regions worldwide (Ascher \u0026amp; Pickering 2022).\u003c/p\u003e\u003cp\u003eWithin this group, bees of the genus \u003cem\u003eMelipona\u003c/em\u003e are among the most commonly managed by beekeepers due to their larger body size and easy of handling (Jaff\u0026eacute; et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Several species also play an important role in the local economy, as farmers commercialize their hives and bee products such as honey, pollen, wax, and propolis (Reyes-Gonz\u0026aacute;lez et al. 2016; Gemim \u0026amp; De Melo Silva 2017). The loss and degradation of Atlantic Forest habitats are major threats to stingless bees (Roberts et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). However, meliponiculture - the rational breeding of stingless bees - has gained popularity among small-scale producers and may represent a promising strategy for species conservation (Nogueira-Neto et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e1986\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe survival of a bee colony depends on the success of foragers in gathering food resources such as nectar and pollen. Pollen collected by stingless bees is particularly informative for studying their trophic ecology (Roubik \u0026amp; Pati\u0026ntilde;o 2018). Although most stingless bees are generalist foragers, even species with more specialized habits trend to exploit a wide range of plants throughout the year (Bueno et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Given the high diversity of stingless bee species coexisting in tropical ecosystems, trophic niche overlap becomes an important factor to consider. In meliponaries, the artificial increase in colony density and the number of individuals foraging in the same area intensifies this overlap, highlighting the need to investigate how these species share floral resources.\u003c/p\u003e\u003cp\u003eThis study aimed to evaluate seasonal trophic niche overlap among four stingless bee species of the genus \u003cem\u003eMelipona\u003c/em\u003e (\u003cem\u003eMelipona quadrifasciata\u003c/em\u003e, \u003cem\u003eM. bicolor\u003c/em\u003e, \u003cem\u003eM. torrida\u003c/em\u003e, and \u003cem\u003eM. mondury\u003c/em\u003e), which are commonly kept together in meliponaries. These species differ in body size and coloration. Specifically, we sought to answer the following questions: (1) How do the floral resource choices of \u003cem\u003eM. quadrifasciata\u003c/em\u003e, \u003cem\u003eM. bicolor\u003c/em\u003e, \u003cem\u003eM. torrida\u003c/em\u003e, and \u003cem\u003eM. mondury\u003c/em\u003e vary between warm (spring and summer) and cold (autumn and winter) seasons? (2) Do body size, coloration, and phylogenetic distance influence trophic niche overlap among these species?\u003c/p\u003e"},{"header":"Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e\u003ch2\u003eStudy area\u003c/h2\u003e\u003cp\u003eThe study was carried out at the \"Melipon\u0026aacute;rio Abelhas da Mata, located in the municipality of Guarapuava - Paran\u0026aacute; - Brasil (25\u0026deg;22\u0026rsquo;23,69\u0026rdquo;S; 51\u0026deg;26\u0026rsquo;17,967\u0026rdquo;W, 1.119 m altitude) in a peri-urban region of the city. The region is located in a transitional area between Araucaria Moist Forest (Mixed Ombrophilous Forest) and high-altitude grasslands, originally characterized by natural fields interspersed with patches of Atlantic Forest. The climate is classified as humid subtropical and mesothermal, with an average annual temperature of 16.8\u0026deg;C (Costa \u0026amp; Andrade \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), corresponding to the Cfb type in the K\u0026ouml;ppen\u0026ndash;Geiger climate classification. The southern region of Brazil does not have a well-defined rainy season, having a uniform distribution of precipitation throughout the year.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003c/div\u003e\n\u003ch3\u003eStingless bee species selected\u003c/h3\u003e\n\u003cp\u003eThe four species chosen for this study are endemic to the Atlantic Forest (Nieh 2003; Camargo et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Lopes et al. 2022), commonly bred together in meliponaries (Nogueira-Neto \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e1997\u003c/span\u003e) and were housed in boxes of wood for over a year at the meliponary. These target species were: \u003cem\u003eMelipona quadrifasciata quadrifasciata\u003c/em\u003e Lepeletier, 1836; \u003cem\u003eMelipona bicolor schencki\u003c/em\u003e Gribodo, 1896; \u003cem\u003eMelipona torrida\u003c/em\u003e Friese, 1916; and \u003cem\u003eMelipona mondury\u003c/em\u003e Smith, 1893. The selection of these stingless bee species was based on two criteria: 1) their common use in consorted meliponiculture, and 2) the differences in foraging activities, reflected in their flight ranges and the resource selection. Although the four species are considered generalists, we evaluated three aspects such as size and color of the body and phylogenetic distance between them.\u003c/p\u003e\n\u003ch3\u003eStingless bee monitoring and sample processing\u003c/h3\u003e\n\u003cp\u003eThe study region does not have a well-defined dry season, with precipitation being evenly distributed throughout the year (Pereira et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Therefore, the data collection was organized according to warm (spring\u0026ndash;summer) and cold (autumn\u0026ndash;winter) seasons.\u003c/p\u003e\u003cp\u003eField sampling was conducted monthly over a ten-month period (from November 2022 to August 2023), ensuring temporal coverage of both warm and cold seasons. Each sampling event consisted of standardized collections performed under suitable weather conditions (temperatures between 18\u0026ndash;20 \u0026ordm;C and absence of rainfall), when stingless bees are known to be most active (Gianinni \u0026amp; Imperatriz-Fonseca 1986; Hil\u0026aacute;rio et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2000\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eFor each of the four focal species, pollen was collected from five foraging individuals originating from five distinct colonies, totaling 20 samples per month. Sampling effort was evenly distributed across the study period, resulting in 200 pollen samples in total. While most collections could be completed within a single field day, during colder months additional sampling time was occasionally required to ensure sufficient representation of all species. To obtain the pollen samples used by stingless bees, the entrances were blocked with small sticks, preventing forage bees from entering. As soon as the foraging bees approached and tried to enter the hive, only those that were carrying pollen on their corbiculae were selected and captured. This capture was done manually or with the help of a small plastic container with a lid. The collected pollen was stored in microtubes (1 ml) (Eppendorf type), containing AFA solution (5% acetic acid, 5% formaldehyde and 90% of 50% alcohol), and the bees were released to return to their activities.\u003c/p\u003e\n\u003ch3\u003ePalynological analysis\u003c/h3\u003e\n\u003cp\u003eThe samples underwent the acetolysis process described by Erdtman (\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e1952\u003c/span\u003e) with some modifications. Microscopes slides were prepared using glycerinated gelatin (Johansen 1940) and examined under a bright field microscopy to identify pollen types to the lowest possible taxonomic level. Taxons were identified to the lowest taxonomic level possible, in most cases to families and, occasionally, to different genera or morphotypes within the same family (Appendix).\u003c/p\u003e\u003cp\u003eThe pollen identification was conducted with the assistance of a botanic specialist and virtual platforms such as Online Pollen Catalog Network (RCPol) and Global Pollen Project (Martin \u0026amp; Harvey \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Diagnostic features considered included the size, aperture type, and surface characteristics of the pollen exine.\u003c/p\u003e\n\u003ch3\u003eMorphological characterization of stingless bees\u003c/h3\u003e\n\u003cp\u003eTen individuals from each of the four stingless bee species were collected at the entrance of their nests. Each specimen was euthanized in a killing chamber using ethyl acetate. Each specimen was measured for its mass (mg) using a precision scale. The average weight of the 10 individuals from each species was used as a proxy for body size. Regarding coloration, one representative specimen per species was analyzed and classified based on the percentage of the body surface (dorsal view) displaying light and dark colors, using the R packages \u003cem\u003emagick\u003c/em\u003e and \u003cem\u003eimager.\u003c/em\u003e This analysis was intended to provide a representative overview of interspecific differences in coloration patterns.\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\u003e\u0026ndash; Morphological character matrix of the four analyzed stingless bee species (\u003cem\u003eMelipona quadrifasciata\u003c/em\u003e, \u003cem\u003eM. bicolor\u003c/em\u003e, \u003cem\u003eM. torrida\u003c/em\u003e, and \u003cem\u003eM. mondury\u003c/em\u003e).\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"4\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eBee species\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eAverage weight (mg)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eDark coloration (%)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eLight coloration (%)\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eM. quadrifasciata\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c2\"\u003e\u003cp\u003e62,1\u0026thinsp;\u0026plusmn;\u0026thinsp;4,04\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e87,21\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e12,79\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eM. bicolor\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c2\"\u003e\u003cp\u003e51,1\u0026thinsp;\u0026plusmn;\u0026thinsp;4,43\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e90,21\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e9,78\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eM. torrida\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c2\"\u003e\u003cp\u003e27,2\u0026thinsp;\u0026plusmn;\u0026thinsp;2,44\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e86,51\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e13,49\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eM. mondury\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c2\"\u003e\u003cp\u003e60,6\u0026thinsp;\u0026plusmn;\u0026thinsp;8,49\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2,55\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e97,45\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cdiv id=\"Sec8\" class=\"Section2\"\u003e\u003ch2\u003ePhylogenetic distance\u003c/h2\u003e\u003cp\u003eTo calculate the evolutionary or phylogenetic distance among the four stingless bee species, the molecular phylogeny of \u003cem\u003eMelipona\u003c/em\u003e proposed by Ram\u0026iacute;rez et al. (\u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2010\u003c/span\u003e) was used. To reconstruct the phylogeny and the distances between each taxon, the software FigTree\u0026reg; was used to generate an ultrametric phylogenetic tree. To update the nomenclature and evolutionary relationships, substitute taxa were used for the species in the present study: \u003cem\u003eMelipona quadrifasciata anthidioides\u003c/em\u003e was used as a substitute for \u003cem\u003eM. quadrifasciata quadrifasciata\u003c/em\u003e; \u003cem\u003eM. rufiventris rufiventris\u003c/em\u003e as a substitute for \u003cem\u003eM. mondury\u003c/em\u003e; and \u003cem\u003eM. marginata\u003c/em\u003e as a substitute for \u003cem\u003eM. torrida\u003c/em\u003e. The criteria for these substitutions were as follows: the possible evolutionary proximity between the subspecies of \u003cem\u003eM. quadrifasciata\u003c/em\u003e (Batalha-Filho et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2010\u003c/span\u003e); the possible evolutionary proximity between \u003cem\u003eM. marginata\u003c/em\u003e and \u003cem\u003eM. torrida\u003c/em\u003e, since the latter replaced the name previously given to a former subspecies of \u003cem\u003eM. marginata\u003c/em\u003e, \u003cem\u003eM. marginata obscurior\u003c/em\u003e (Melo \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2013\u003c/span\u003e); and the possible evolutionary proximity between \u003cem\u003eM. rufiventris\u003c/em\u003e and \u003cem\u003eM. mondury\u003c/em\u003e, since the latter was previously considered a subspecies of the former (Melo 2003). The phylogenetic distances between species pairs were calculated based on the number of evolutionary steps between each species pair.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec9\" class=\"Section2\"\u003e\u003ch2\u003eData analysis\u003c/h2\u003e\u003cp\u003eFor each stingless bee species, both pollen richness (i.e., the number of pollen morphotypes) and feeding niche breadth were calculated. Niche breadth was calculated using Levin\u0026rsquo;s index (B), with the standardized value indicated as BA.\u003c/p\u003e\u003cp\u003eBased on the relative abundances of pollen morphotypes, trophic niche overlap among the four stingless bee species was analyzed. Niche overlap for Warm and Cold Season was assessed using the Pianka Index and a Non-Metric Multidimensional Scaling (NMDS) perceptual map. A permutation test with 1,000 randomizations was applied to evaluate whether the observed niche overlaps were significantly higher than expected by chance. All analyses were conducted in R using the vegan package (v. 2.5-5; Oksanen et al. 2019).\u003c/p\u003e\u003cp\u003eBased on the size and color data of the four stingless bee species (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e), two distance matrices were constructed: one for size distance and one for color distance, both based on Euclidean Distance. Additionally, from the phylogeny of the genus \u003cem\u003eMelipona\u003c/em\u003e (Ram\u0026iacute;rez et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), a phylogenetic distance matrix was constructed for the four stingless bee species.\u003c/p\u003e\u003cp\u003eTo test which variables best explain the feeding niche overlap among the four stingless bee species, a Generalized Linear Model (GLM) approach was used. The response variable was the trophic niche overlap values, and the predictor variables were Size Distance (DSize), Color Distance (DColor), and Phylogenetic Distance (DPhyl). To avoid the effect of collinearity among predictors in the GLM, a Variance Inflation Factor (VIF) analysis was conducted using the \u0026ldquo;fmsb\u0026rdquo; 0.7.3 package. In this analysis, predictor variables with VIF values greater than 3 are considered collinear and should be removed from subsequent analyses. Since all VIF values were below 3 (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e), all predictor variables were retained for model construction.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eVariance inflation factors for the predictor variables of Trophic Niche Overlap among four stingless bee species from the Brazilian Atlantic Forest (\u003cem\u003eMelipona quadrifasciata\u003c/em\u003e, \u003cem\u003eM. bicolor, M. torrida\u003c/em\u003e, and \u003cem\u003eM. mondury\u003c/em\u003e). GVIF\u0026thinsp;=\u0026thinsp;Generalized Variance Inflation Factor; DSize\u0026thinsp;=\u0026thinsp;Size distance among the four species (Euclidean Distance); DColor\u0026thinsp;=\u0026thinsp;Color distance among the four species (Euclidean Distance); and DPhyl\u0026thinsp;=\u0026thinsp;Phylogenetic distance among the four species.\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=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eVariable\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eGVIF\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDSize\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1,1602\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDColor\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e2,1148\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDPhyl\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e2,0239\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\u003eEight models were constructed, including all predictor variables (DSize, DColor, and Phyl), all possible combinations among them, a global model (with all variables), and a null model (with none). The best model was selected based on the Akaike Information Criterion (AIC) and the AIC model weights. In this analysis, the lower the AIC value and the higher the AIC weight, the better the model fit for predicting the response variable (trophic niche overlap). When the difference between the best model and other models (ΔAIC) was less than 2, those models were considered equally predictive. In such cases, we performed model averaging to combine their estimates and account for model selection uncertainty. These analyses were performed using the \u0026ldquo;AICcmodavgpa\u0026rdquo; 2.3.2 package (Mazerolle \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The GLM analyses were carried out in the R environment (R Core Team, 2013).\u003c/p\u003e\u003c/div\u003e"},{"header":"Results","content":"\u003cp\u003e\u003cem\u003eMelipona bicolor\u003c/em\u003e exhibited the highest richness of pollen morphotypes used (S\u0026thinsp;=\u0026thinsp;18), followed closely by \u003cem\u003eM. torrida\u003c/em\u003e (S\u0026thinsp;=\u0026thinsp;17), while \u003cem\u003eM. mondury\u003c/em\u003e showed the lowest richness (S\u0026thinsp;=\u0026thinsp;13). In terms of trophic niche breadth, \u003cem\u003eM. quadrifasciata\u003c/em\u003e presented the highest value (B\u0026thinsp;=\u0026thinsp;10.1), whereas \u003cem\u003eM. mondury\u003c/em\u003e had the lowest (B\u0026thinsp;=\u0026thinsp;3.5) during the Warm Season (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). The most represented plant family in the pollen samples was Myrtaceae, which dominated the diets of all species. Other frequent families included Arecaceae, Fabaceae, and Moraceae. Notably, Asteraceae appeared only during the Warm Season and was consumed exclusively by \u003cem\u003eM. quadrifasciata\u003c/em\u003e and \u003cem\u003eM. torrida\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003e\u0026ndash; Breadth of trophic niches (Levin\u0026rsquo;s index: B and standardized BA), richness of visited plant morphotypes (S), and pairwise niche overlap values (Pianka index) among stingless bee species raised in meliponaries in southern Brazil (Warm season). P-values are shown only for comparisons with statistically significant differences (based on permutation tests with 1,000 randomizations).\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"8\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eB\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eBA\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eS\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"4\" nameend=\"c8\" namest=\"c5\"\u003e\u003cp\u003eNiche overlap (Warm season)\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u003cem\u003eM. quadrifasciata\u003c/em\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u003cem\u003eM. torrida\u003c/em\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003e\u003cem\u003eM. bicolor\u003c/em\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c8\"\u003e\u003cp\u003e\u003cem\u003eM. mondury\u003c/em\u003e\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eM. quadrifasciata\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e10.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.36\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e16\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u003cb\u003e0.77 (p\u0026thinsp;=\u0026thinsp;0,004)\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e\u003cb\u003e0.90 (p\u0026thinsp;=\u0026thinsp;0,001)\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e\u003cb\u003e0.70 (p\u0026thinsp;=\u0026thinsp;0,006)\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eM. torrida\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e9.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.32\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e15\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.77\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e\u003cb\u003e0.85 (p\u0026thinsp;=\u0026thinsp;0,003)\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e\u003cb\u003e0.86 (p\u0026thinsp;=\u0026thinsp;0,001)\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eM. bicolor\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e8.3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.29\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e17\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.90\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.85\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e\u003cb\u003e0.82 (p\u0026thinsp;=\u0026thinsp;0,001)\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eM. mondury\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.70\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.86\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.82\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e-\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\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab4\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 4\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003e\u0026ndash; Breadth of trophic niches (Levin\u0026rsquo;s index: B and standardized BA), richness of visited plant morphotypes (S), and pairwise niche overlap values (Pianka index) among stingless bee species raised in meliponaries in southern Brazil (Cold season). P-values are shown only for comparisons with statistically significant differences (based on permutation tests with 1,000 randomizations).\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"8\"\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\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eB\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eBA\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eS\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"4\" nameend=\"c8\" namest=\"c5\"\u003e\u003cp\u003eNiche overlap (Cold season)\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u003cem\u003eM. quadrifasciata\u003c/em\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u003cem\u003eM. torrida\u003c/em\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003e\u003cem\u003eM. bicolor\u003c/em\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c8\"\u003e\u003cp\u003e\u003cem\u003eM. mondury\u003c/em\u003e\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eM. quadrifasciata\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5.4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.49\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.59\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e\u003cb\u003e0.87 (p\u0026thinsp;=\u0026thinsp;0,017)\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e\u003cb\u003e0.86 (p\u0026thinsp;=\u0026thinsp;0,026)\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eM. torrida\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e4.4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.37\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.59\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.61\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.81\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eM. bicolor\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.45\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.87\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.61\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.71\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eM. mondury\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.44\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.86\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.81\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.71\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cdiv id=\"Sec11\" class=\"Section2\"\u003e\u003ch2\u003eSeasonal variation\u003c/h2\u003e\u003cp\u003eThroughout the year, \u003cem\u003eM. mondury\u003c/em\u003e collected significantly fewer pollen morphotypes than \u003cem\u003eM. quadrifasciata, M. bicolor\u003c/em\u003e, and \u003cem\u003eM. torrida\u003c/em\u003e. Similarly, \u003cem\u003eM. mondury\u003c/em\u003e also exhibited lower trophic niche breadth compared to the other species (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eA). This same pattern was observed during the Warm Season, but during the Cold Season there were no differences in either the number of pollen morphotypes or the niche breadths among the species (Tables\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e and \u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eIn both seasons, the highest trophic niche overlap was observed between \u003cem\u003eM. quadrifasciat\u003c/em\u003ea and \u003cem\u003eM. bicolor\u003c/em\u003e (91% in the Warm Season and 87% in the Cold Season), indicating resource sharing between these species. During the Warm Season, all species pairs exhibited significantly higher niche overlap than expected by chance (p\u0026thinsp;\u0026lt;\u0026thinsp;0.01 for all pairs), with particularly high values between \u003cem\u003eM. torrida\u003c/em\u003e and \u003cem\u003eM. mondury\u003c/em\u003e (86%, p\u0026thinsp;=\u0026thinsp;0.001), and between \u003cem\u003eM. bicolor\u003c/em\u003e and \u003cem\u003eM. torrida\u003c/em\u003e (85%, p\u0026thinsp;=\u0026thinsp;0.003). The lowest overlap occurred between \u003cem\u003eM. quadrifasciata\u003c/em\u003e and \u003cem\u003eM. mondury\u003c/em\u003e (69%, p\u0026thinsp;=\u0026thinsp;0.006).\u003c/p\u003e\u003cp\u003eIn contrast, during the Cold Season, only two species pairs exhibited significant niche overlap: \u003cem\u003eM. quadrifasciata\u003c/em\u003e with \u003cem\u003eM. bicolor\u003c/em\u003e (87%, p\u0026thinsp;=\u0026thinsp;0.017) and with \u003cem\u003eM. mondury\u003c/em\u003e (87%, p\u0026thinsp;=\u0026thinsp;0.026). Other pairs, such as \u003cem\u003eM. quadrifasciata\u003c/em\u003e and \u003cem\u003eM. torrida\u003c/em\u003e (60%, p\u0026thinsp;=\u0026thinsp;0.27), and \u003cem\u003eM. torrida\u003c/em\u003e and \u003cem\u003eM. bicolor\u003c/em\u003e (61%, p\u0026thinsp;=\u0026thinsp;0.22), did not show significant overlap, suggesting more distinct foraging patterns in those interactions during this period (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eOverall, in the Cold Season, stingless bee species reduced both the number of pollen morphotypes collected and their trophic niche breadth compared to the Warm Season. However, \u003cem\u003eM. mondury\u003c/em\u003e did not follow this same trend (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eC).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eModels 4, 5, and 6 were equally predictive in explaining trophic niche overlap among the stingless bee species. Even after model averaging, the GLM analysis did not support any significant relationships between the predictive variables and trophic niche overlap. Color, body size, and phylogenetic distance were not significantly associated with variations in trophic niche overlap when considering all samples combined (Tables\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e and \u003cspan refid=\"Tab6\" class=\"InternalRef\"\u003e6\u003c/span\u003e), as well as in the separate analyses for the warm and cold seasons.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab5\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 5\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003e\u0026ndash; Generalized Linear Models (GLM) evaluated for their predictive power of Trophic Niche Overlap in four stingless bee species from the Brazilian Atlantic Forest (\u003cem\u003eMelipona quadrifasciata\u003c/em\u003e, \u003cem\u003eM. bicolor\u003c/em\u003e, \u003cem\u003eM. torrida\u003c/em\u003e, and \u003cem\u003eM. mondury\u003c/em\u003e). AICc\u0026thinsp;=\u0026thinsp;Akaike Information Criterion corrected for small sample sizes; ΔAICc\u0026thinsp;=\u0026thinsp;Difference in AICc relative to the best model; AICcWt\u0026thinsp;=\u0026thinsp;AICc model weight.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"4\"\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\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eModel\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eAICc\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eΔAICc\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAICcWt\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-2,17\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0,41\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-1,94\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0,22\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0,37\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-0,92\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1,25\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0,22\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eNull\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5,95\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e8,12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0,01\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e25,47\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e27,64\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e25,53\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e27,70\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e27,83\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eGlobal\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u0026gt;\u0026thinsp;1.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u0026gt;\u0026thinsp;1.000\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0\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\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab6\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 6\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003e\u0026ndash; Coefficient of the predictor variable (DCor) for the best-fitted Generalized Linear Model (GLM) predicting Trophic Niche Overlap in four stingless bee species from the Brazilian Atlantic Forest (\u003cem\u003eMelipona quadrifasciata, M. bicolor, M. torrida\u003c/em\u003e, and \u003cem\u003eM. mondury).\u003c/em\u003e DPhyl\u0026thinsp;=\u0026thinsp;Phylogenetic distance; DSize\u0026thinsp;=\u0026thinsp;Size distance; DColor\u0026thinsp;=\u0026thinsp;Color distance; GLM\u0026thinsp;=\u0026thinsp;Generalized Linear Models; TNO\u0026thinsp;=\u0026thinsp;Trophic Niche Overlap.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"5\"\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\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eCoefficient\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eStandad error\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003et-value\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003ep-value\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eIntercept\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-0, 4822371\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0,1209295\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e2,988\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0,00281*\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eDPhyl\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-0,0512033\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0,0856634\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0,489\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0,62468\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eDSize\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-1,5423716\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2,8637592\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0,439\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0,66067\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eDColor\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-0,0001646\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0,0005282\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e\u003cp\u003e0,242\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0,80893\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003c/div\u003e"},{"header":"Discussion","content":"\u003cp\u003eClimatic factors are intrinsically linked to the availability and density of resources for stingless bees (Keppner \u0026amp; Jarau \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Moura et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Most studies evaluating the effects of seasonality on niche overlap in stingless bees analyze similarities and differences between the rainy and dry seasons (Brito et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Santos et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Escobedo-Kenefic et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Ferreira et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The results presented here demonstrate that stingless bees follow the pattern observed in previous studies, foraging more intensively for pollen resources during the warm season, due to increased floral abundance and higher temperatures (Souza et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Ferreira-Junior et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). In contrast, during the cold season, a reduction in species richness and trophic niche breadth was observed in three of the studied species \u0026mdash; \u003cem\u003eMelipona quadrifasciata, M. bicolor\u003c/em\u003e, and \u003cem\u003eM. torrida\u003c/em\u003e \u0026mdash; all of which conformed to this pattern. However, \u003cem\u003eM. mondury\u003c/em\u003e deviated from it, maintaining similar niche breadth values across both seasons, suggesting a distinct foraging strategy. This behavior may be related to its biogeographic origin: although native to the Atlantic Forest, \u003cem\u003eM. mondury\u003c/em\u003e is not originally found in the study region, being more commonly distributed in lower-altitude areas of the Atlantic Forest along the Brazilian coast, where the climate is warmer and more humid (Mouga \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Mouga \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Mouga \u0026amp; Nogueira-Neto 2015). Furthermore, this species does not undergo reproductive diapause, which sustains its year-round demand for pollen and may increase its vulnerability in subtropical climates with pronounced winters. The increase in niche overlap between \u003cem\u003eM. quadrifasciata\u003c/em\u003e and \u003cem\u003eM. mondury\u003c/em\u003e during cold season may lead to challenges in colony development due to the reduced availability of floral resources during this period. These findings have important implications for meliponiculture, particularly regarding colony maintenance during colder periods. Supplemental feeding, including sugar syrup and pollen-based food, may be essential to support the survival and proper development of colonies maintained outside their natural range.\u003c/p\u003e\u003cp\u003eSome botanical taxa form the basis of stingless bees\u0026rsquo; diet (e.g., Myrtaceae, Arecaceae), while others are used more occasionally, which helps explain the trophic relationships observed in the networks of both warm and cold seasons. During the cold season, the availability of flowering plants decreases, which strengthens the bee\u0026ndash;plant interactions (Ferreira et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Additionally, several taxa recorded correspond to herbaceous species likely native to grasslands or forest edges (i.e. Asteraceae, Begoniaceae, Labiateae, Urticaceae, Verbenaceae) reflecting the transitional nature of the study area between Araucaria Moist Forest (Mixed Ombrophilous Forest) and high-altitude grasslands.\u003c/p\u003e\u003cp\u003eThe concept of a key resource can be understood as a food resource capable of determining the survival potential of organisms. Additionally, it can influence the composition of biological communities, as its absence or low availability may affect species\u0026rsquo; population density, long-term viability, or even lead to local extinction (Primack \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Stingless bees exhibit a certain affinity for resources considered key to colony maintenance. The collection of morphotypes from botanical families that flower abundantly throughout the year, such as Myrtaceae, facilitates species coexistence in environments like meliponaries, where there is an artificially increased local density of bees. Identifying key resources may therefore serve as an important tool for effective long-term conservation and management strategies.\u003c/p\u003e\u003cp\u003eBees of the genus \u003cem\u003eMelipona\u003c/em\u003e exhibit social behavior and live in populous, perennial colonies, requiring them to exploit a wide range of floral resources throughout the year. Most flowering plants are ephemeral sources, blooming for only a few days (Ramalho et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). Therefore, a generalist foraging strategy is considered the norm among eusocial bees of the family Apidae. Their foraging patterns vary seasonally in response to environmental factors such as flowering periods \u0026mdash; with summer and spring being seasons of abundant floral availability (Winston \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2003\u003c/span\u003e). \u003cem\u003eMelipona\u003c/em\u003e bees have been recorded visiting a variety of botanical families, depending on the surrounding vegetation (Bueno et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Although stingless bees forage on many species from different plant families, some families are more frequently utilized. Floristic surveys conducted in the Araucaria Moist Forest have identified the Myrtaceae family as one of the richest or most abundant (Cordeiro \u0026amp; Rodrigues \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Albuquerque et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Vieira \u0026amp; May \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) with about 246 species belonging to 20 genera recorded solely within the Atlantic Forest domain, and 220 species belonging to 16 genera recorded in Paran\u0026aacute; state alone (Flora e Funga do Brasil 2025). In the Atlantic Forest, Meliponini bees frequently visit flowers of \u003cem\u003eMyrcia tomentosa\u003c/em\u003e, particularly species of \u003cem\u003eMelipona\u003c/em\u003e and \u003cem\u003ePlebeia\u003c/em\u003e (Ramalho \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). In S\u0026atilde;o Paulo, \u003cem\u003eMelipona mondury\u003c/em\u003e (previously cited as \u003cem\u003eM. rufiventris\u003c/em\u003e) has been identified as one of the most effective pollinators of Myrtaceae species (Fidalgo \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). Other studies have consistently reported Myrtaceae as one of the most visited plant families by bees in Brazil (Carvalho et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Antonini et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Ramalho et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2007\u003c/span\u003e), with \u003cem\u003eMelipona\u003c/em\u003e species being the most frequent pollinators of Myrtaceae in the country. This is largely attributed to the presence of nectaries in these plants, which offer nectar in addition to pollen (Antonini et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Gressler et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2006\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe Arecaceae family exhibits asynchronous flowering, providing floral resources during several months of the year. In Brazil, approximately 270 species and 38 native genera are recorded, standing out for their abundance and species richness in Neotropical forests (Lara et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). In the state of Paran\u0026aacute;, 23 Arecaceae species have been reported (Flora do Brasil \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). In Paran\u0026aacute; state, 17 species from 11 genera of Arecaceae have been recorded. The importance of Arecaceae in the pollen diet of \u003cem\u003eMelipona\u003c/em\u003e bees has already been highlighted by Oliveira et al. (2009). The Araucariaceae family, in turn, was recorded exclusively in the samples of \u003cem\u003eM. mondury\u003c/em\u003e and is noteworthy since these plants do not produce flowers, yet their pollen is protein-rich and nutritionally valuable. Other studies have also reported the use of anemophilous plants (e.g., Cecropiaceae) by stingless bees in the Atlantic Forest (Morgado et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Braga et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2014\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe trophic niche breadth of the four stingless bee species endemic to the Atlantic Forest analyzed in this study reveals a generalist feeding behavior, which is common among eusocial bees and reflects their plasticity in the use of pollen resources. Body size and coloration are important traits in the trophic niche segregation of stingless bees (Hil\u0026aacute;rio et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Pereboom \u0026amp; Biesmeijer 2003; Teixeira \u0026amp; Campos \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Biesmeijer \u0026amp; Slaa 2006). However, these traits were not effective in predicting the niche overlap observed in this study. This may be due to the fact that the analysis included only four species from the same genus, \u003cem\u003eMelipona\u003c/em\u003e. Although the \u003cem\u003eMelipona\u003c/em\u003e genus is highly diverse, the selected species are among the largest stingless bees and show little variation in body size, as well as a close evolutionary relationship. Despite morphological differences, this variation could not be linked to the patterns of niche overlap among the analyzed species. However, increasing the sampling effort to include species from other genera, with greater variation in body size, coloration, and phylogenetic distance, may help to better understand the dietary differences among these bees.\u003c/p\u003e\u003cp\u003eThe results obtained in this study can directly support the management and strategic organization of meliponaries located in the Atlantic Forest ecosystem, particularly in colder regions such as the Araucaria Moist Forest. The identification of key resources, such as plant species from the Myrtaceae and Arecaceae families, which proved highly attractive to the studied bee species, suggests that cultivating these plants may serve as an important nutritional support strategy throughout the year. Moreover, data on niche overlap indicate that the coexistence of certain species may require special attention. For example, \u003cem\u003eM. quadrifasciata\u003c/em\u003e and \u003cem\u003eM. mondury\u003c/em\u003e showed a marked increase in niche overlap during the cold season, suggesting that keeping both species in the same meliponary should be approached with caution. If this combination is adopted, it is recommended to increase the availability of supplemental feeding for both species. It is also noteworthy that \u003cem\u003eM. mondury\u003c/em\u003e exhibited a distinct foraging strategy, maintaining a wide niche breadth even in the cold season, likely due to the absence of reproductive diapause. This trait may render the species more vulnerable in colder climates, as it must maintain a high demand for pollen even when floral resources are scarce, which could compromise colony viability without adequate support.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eEthical Approval\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis research was supported by the \u003cstrong\u003eCoordena\u0026ccedil;\u0026atilde;o de Aperfei\u0026ccedil;oamento de Pessoal de N\u0026iacute;vel Superior (CAPES \u0026ndash; Brazil)\u003c/strong\u003e. Proccess number 88887.649588/2021-00.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthors\u0026apos; contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAmanda Gar\u0026ccedil;oa Raulik conducted the fieldwork, performed data analyses, wrote, and revised the manuscript. Luiz Gabriel Ludwig assisted with the statistical analyses. Adriano contributed to the palynological analyses. Jo\u0026atilde;o critically revised the manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConflicts of interest/Competing interests\u003c/strong\u003e\u003c/p\u003e\u003cp\u003eThe authors declare that they have no conflicts of interest.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAlbuquerque JMD, Watzlawick LF, Mesquita NSD (2011) Efeitos do uso em sistema faxinal na flor\u0026iacute;stica e estrutura em duas \u0026aacute;reas da Floresta Ombr\u0026oacute;fila Mista no mun\u0026iacute;cipio de Rebou\u0026ccedil;as. 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Brazilian J Biology 66:731\u0026ndash;737\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eTeixeira LV, Campos FNM (2005) In\u0026iacute;cio da atividade de v\u0026ocirc;o em abelhas sem ferr\u0026atilde;o (Hymenoptera, Apidae): influ\u0026ecirc;ncia do tamanho da abelha e da temperatura ambiente. Revista Brasileira de Zooci\u0026ecirc;ncias 7:195\u0026ndash;202\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eVieira NC, May D (2020) Regenera\u0026ccedil;\u0026atilde;o natural de Myrtaceae em uma unidade de conserva\u0026ccedil;\u0026atilde;o urbana, Curitiba, PR. Brasil REVSBAU 15(1):39\u0026ndash;49\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eWinston ML (2003) A Biologia da Abelha. Magister, Porto Alegre\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":true,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"neotropical-entomology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"nent","sideBox":"Learn more about [Neotropical Entomology](https://www.springer.com/journal/13744)","snPcode":"13744","submissionUrl":"https://www.editorialmanager.com/nent/default2.aspx","title":"Neotropical Entomology","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Meliponiculture, Overlap niche, Pollination, Seasonality, Trophic ecology","lastPublishedDoi":"10.21203/rs.3.rs-8042458/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-8042458/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eUnderstanding how stingless bee species share floral resources is essential for optimizing meliponiculture and supporting pollinator conservation. This study investigated the trophic niche overlap of four \u003cem\u003eMelipona\u003c/em\u003e species (\u003cem\u003eM. quadrifasciata, M. bicolor, M. torrida\u003c/em\u003e, and \u003cem\u003eM. mondury\u003c/em\u003e) in Southern Brazil across warm and cold seasons. Pollen samples were collected monthly over ten months and analyzed to assess richness, trophic niche breadth, and overlap between species' trophic niches. Results showed seasonal variation in foraging patterns, with all species (except \u003cem\u003eM. mondury\u003c/em\u003e) reducing their niche breadth in colder months. \u003cem\u003eMelipona mondury\u003c/em\u003e, a species native to warmer regions of the Atlantic Forest and lacking reproductive diapause, maintained a broad niche year-round, possibly increasing its vulnerability in colder climates. The highest overlap occurred between \u003cem\u003eM. quadrifasciata\u003c/em\u003e and \u003cem\u003eM. mondury\u003c/em\u003e during the cold season, raising concerns about managing these species in consorted meliponiculture systems. The most frequently visited botanical families included Myrtaceae, Arecaceae, Asteraceae, Moraceae, and Fabaceae highlighting their importance as key floral resources. These findings highlight the importance of considering seasonal dynamics and species-specific traits in meliponiculture, including the need for targeted supplemental feeding and resource planning. Promoting key botanical resources such as Myrtaceae and Arecaceae can support sustainable colony development in high-density environments.\u003c/p\u003e","manuscriptTitle":"Trophic niche overlap in stingless bees (Melipona Illiger, 1806) across season in Southern Brazil","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-11-28 08:34:37","doi":"10.21203/rs.3.rs-8042458/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewerAgreed","content":"","date":"2025-12-22T13:49:09+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-11-24T12:16:27+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"Neotropical Entomology","date":"2025-11-13T21:27:22+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-11-11T18:53:04+00:00","index":"","fulltext":""},{"type":"submitted","content":"Neotropical Entomology","date":"2025-11-10T11:27:57+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"neotropical-entomology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"nent","sideBox":"Learn more about [Neotropical Entomology](https://www.springer.com/journal/13744)","snPcode":"13744","submissionUrl":"https://www.editorialmanager.com/nent/default2.aspx","title":"Neotropical Entomology","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"20389926-7ecd-41fd-8e7a-90e5d4521d77","owner":[],"postedDate":"November 28th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-05-22T11:10:40+00:00","versionOfRecord":[],"versionCreatedAt":"2025-11-28 08:34:37","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-8042458","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-8042458","identity":"rs-8042458","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}