Abstract
Honey bees (Apis mellifera), depend on the availability and diversity of floral resources within the landscape for their survival and productivity. This study characterized the floristic composition, richness and diversity of vegetation in a beekeeping area of the Peruvian Amazon in order to evaluate its potential to provide forage resources for honey bees. Vegetation was surveyed within a circular area of 28.3 km2 centered on an apiary, where 24 quadrats were distributed along four transects. Species composition analyses were conducted for the general floristic composition and for species with bee foraging potential, followed by PERMANOVA and non-metric multidimensional scaling (NMDS) analyses. A total of 124 plant species belonging to 103 genera were recorded. Among them, 90 species belonging to 75 genera were identified as bee forage species, of which 23% were classified as pollen sources, whereas 77% provided both nectar and pollen. Both the general floristic composition and the bee forage species assemblage were dominated by the low vegetation stratum (≥86%), indicating a high prevalence of herbaceous species. Shannon, Simpson, evenness, and Berger–Parker dominance indices for the general floristic composition were 2.05 (0.38), 0.81 (0.08), 0.79 (0.09), and 0.32 (0.11), respectively, with no significant differences among land-cover types, transect orientations, or distances from the apiary, except for the Berger–Parker dominance index, for which distance had a significant effect. PERMANOVA revealed significant differences in species composition among land-cover types and transects (p < 0.05) for both the general floristic composition and bee forage species, which was further supported by NMDS analysis (stress = 0.16). The high proportion of plant species with documented floral resources and the heterogeneous landscape indicate substantial potential floral resource availability. These findings provide a baseline for understanding the floral resources supporting beekeeping in the study area and highlight the need for future validation through honey bee visitation, palynological, phenological, and resin-resource studies.
Keywords:
bees; beekeeping; bee forage species; cropland; forest; floristic composition; diversity indices 1. Introduction
The honey bee (Apis mellifera) is one of the most important species due to its ecological and economic significance, as it contributes substantially to the pollination of both wild plants and agricultural crops [1,2]. The availability of floral resources, such as nectar, pollen and resins, is essential for the survival, development, and productivity of bee colonies. Nectar provides carbohydrates that serve as an energy source for adult bees, whereas pollen constitutes the primary source of proteins and other nutrients required for larval development and colony growth [3]. Therefore, the diversity and abundance of flowering plants within a landscape strongly influence honey bee foraging behavior and overall colony performance [4].
In recent years, studies have highlighted the importance of landscape composition and vegetation diversity in shaping the availability of floral resources for honey bees [5]. Agricultural landscapes, depending on their management practices and degree of heterogeneity, may either limit or enhance the availability plant resources. Landscapes composed of a mosaic of crops, forest, and semi-natural habitats can provide a continuous supply of floral resources throughout the year, thereby supporting pollinator communities [6]. In contrast, landscapes dominated by monocultures or characterized by low plant diversity may lead to temporary shortages of floral resources, negatively affecting pollinator populations [7]. Recent studies have further demonstrated that floral diversity and landscape composition play a key role in honey bee nutrition, foraging behavior, and colony health, reinforcing the importance of conserving diverse plant communities in agricultural landscapes [8].
The identification of plant species that may provide floral resources for honey bees is essential for understanding foraging patterns and the productive potential of beekeeping systems [9]. Floristic surveys conducted in tropical and subtropical regions have demonstrated that plant diversity strongly influences the quantity and quality of floral resources available to honey bees, ultimately affecting colony development and honey production [10]. However, previous studies have largely focused either on inventories of bee forage species or on the effects of landscape composition on honey bees [11], whereas fewer studies have integrated the general floristic composition of the landscape with the spatial distribution of potential bee forage species within the foraging range of A. mellifera. This gap is particularly relevant in the Andean–Amazonian landscapes of northern Peru, where information on vegetation resources that may support local beekeeping systems remains scarce [12]. Characterizing both the general floristic composition and the spatial availability of potential floral resources can therefore provide a baseline for understanding the ecological resources surrounding apiaries and for guiding future studies on actual honey bee resource use.
In this context, the present study aimed to characterize the general floristic composition, richness, and diversity of the vegetation surrounding an apiary in the Peruvian Amazon and to identify plant species with documented value as floral resources for Apis mellifera. By integrating floristic and spatial analyses, this study provides a landscape-scale assessment of the potential availability of nectar and pollen resources within the honey bee foraging range, establishing a baseline for future studies based on direct observations of honey bee visitation or palynological analyses to validate resource use under local conditions.
2. Materials and Methods
2.1. Study Area
The study was conducted in an apiary located in the district of Levanto (6°17′51.66″ S, 77°52′54.46″ W), Chachapoyas Province, Amazonas Region, Peru. The area is characterized by a humid tropical climate, with high rainfall (2.3 ± 4.3 mm day−1) and temperatures ranging from 11.5 ± 1.1 °C (minimum) to 21.4 ± 2.0 °C (maximum) during the sampling months (April and May 2025), according to records from the National Service of Meteorology and Hydrology of Peru [13]. The surrounding landscape consisted of a mosaic of agricultural fields and forest patches, forming a heterogeneous environment with potential for honey bee foraging. The apiary was selected because of its location within this heterogeneous Andean–Amazonian landscape and the suitable conditions it provided for systematic vegetation surveys within the potential foraging range of Apis mellifera. Permission to conduct the research was granted by the apiary owner.
2.2. Experimental Design and Vegetation Sampling
Vegetation sampling was conducted within a circular area with a radius of 3 km centered on the apiary, considering this distance to be representative of the foraging range of A. mellifera. A total of 24 sampling units (25 m2) were established using a Garmin GPS Montana 700 (Garmin Ltd., Olathe, KS, USA) along four transects oriented according to the cardinal directions (north, south, east, and west), using the apiary as the reference point. Along each transect, six quadrats were positioned at distances of 0.5, 1.0, 1.5, 2.0, 2.5, and 3.0 km from the apiary to assess potential spatial variations in floristic composition. The spatial arrangement of the sampling units is illustrated in Figure 1.
Figure 1.
Spatial arrangement of transects and sampling quadrats within the study area. Red lines represent concentric sampling distances established at 500 m intervals from the apiary.
Within each quadrat, all plant species present at the time of sampling were recorded, regardless of their pollination syndrome, floral reward, or documented visitation by A. mellifera. When necessary, botanical specimens were collected following the procedures described by [14] for subsequent identification using specialized literature and taxonomic databases. Specimens lacking flowers or fruits that could not be reliably identified to species level were assigned to the genus level and recorded as Genus sp. For each species, the number of individuals was recorded to estimate abundance and density. For each species, the number of individuals was recorded to estimate its abundance and density.
Plant species were classified according to vegetation stratum as low (<1.5 m), medium (1.5–5 m), or high (>5 m), following habitat structural stratification criteria that may influence resource accessibility and utilization by honey bees [15,16]. Based on published studies on A. mellifera, other bee species, and regional botanical databases, floral resources were classified as pollen (P), nectar (N), or both (N-P). Regional sources were considered when species-specific information for A. mellifera was unavailable [17,18]. Resin- and oil-producing plants were not included in this classification. No direct observations of A. mellifera visitation, corbicular pollen collection, or systematic phenological monitoring were conducted. Accordingly, the term “potential bee forage species” refers to plant species reported in the literature as providing floral resources and does not imply locally confirmed use or preference by honey bees.
2.3. Data Processing and Ecological Indices
A comprehensive species inventory was compiled and classified according to land-cover type (forest or cropland), transect orientation, and distance from the apiary. Additionally, species were grouped based on their potential value as foraging resources for honey bees.
Species richness was estimated as the total number of species recorded per quadrat. Diversity was assessed using the Shannon & Weaver [19] and Simpson [20] indices, whereas evenness and dominance were calculated using the Pielou [21] and Berger–Parker [22] indices, respectively. These metrics were used to characterize the structure of the general floristic composition and the availability of floral resources for honey bees. Species accumulation curves were generated using randomization and rarefaction methods to evaluate the adequacy and representativeness of the sampling effort.
2.4. Statistical Analysis
All statistical analyses were performed in RMarkdown (RStudio, version 2025.05.1 + 513, Boston, MA, USA) [23]. The vegan package [24] was used for the calculation of diversity indices and multivariate analyses. Prior to inferential analyses, the assumptions of normality and homogeneity of variances were evaluated using the Shapiro–Wilk and Levene tests, respectively. Because the data met these assumptions, analyses of variance (ANOVA) were performed for species richness, diversity, evenness, and dominance indices using a linear model that included land-cover type (forest vs. cropland), distance from the apiary (0.5, 1.0, 1.5, 2.0, 2.5, and 3.0 km), and transect orientation (north, south, east, and west) as factors. When significant differences were detected, means were compared using Tukey’s honestly significant difference (HSD) test. Statistical significance was assessed at α = 0.05.
Differences in species composition were evaluated using permutational multivariate analysis of variance (PERMANOVA), considering land-cover type, distance from the apiary, and transect orientation as factors. Analyses were based on Bray–Curtis dissimilarity matrices [25] with 10,000 permutations. When significant effects were detected, pairwise comparisons were conducted using the Benjamini–Hochberg adjustment method. Finally, non-metric multidimensional scaling (NMDS) was used to visualize patterns of similarity in species composition among land-cover types, transects, and distances from the apiary. The ordination was based on Bray–Curtis dissimilarities and 10,000 permutations, and model goodness-of-fit was assessed using stress values.
3. Results
3.1. General Floristic Composition
All plant species identified in the sampling area, irrespective of their vegetation stratum or their potential value as forage resources for A. mellifera, were included in the general floristic composition. A total of 124 species belonging to 103 genera were recorded. Of these, 50 species were shared between both land-cover types (forest and cropland), whereas 37 species were exclusively found in forest and another 37 were restricted to cropland (Table 1).
Table 1.
Plant species recorded in the study area, their taxonomic classification, vegetation stratum, and bee foraging potential.
A total of 86% of the recorded species (107 species) belonged to the low vegetation stratum, whereas 11% (13 species) and 3% (4 species) were classified within the medium and high strata, respectively, indicating a predominance of herbaceous and low-growing vegetation (Figure 2a). Regarding bee foraging potential, 73% of the recorded species (90 species) were identified as potential forage resources for honey bees, whereas 27% (34 species) did not provide significant floral resources (Figure 2b).
Figure 2.
Distribution of recorded plant species according to vegetation stratum (a), bee foraging potential (b), and generic composition (c).
The most representative genera were Lolium (11%), Dactylis (9%), Rumex (8%), Digitaria (6%), Galinsoga (5%), Pennisetum (5%), Baccharis (4%), Bidens (3%), and Plantago (3%), whereas Elaphoglossum (2%) and Trifolium (2%) were less represented. The remaining 42% comprised genera with relative frequencies below 2% (Figure 2c).
3.2. Bee Forage Species
A total of 90 species with bee foraging potential, belonging to 75 genera, were recorded. Of these, 74 species (82%) were present in four or fewer quadrats, whereas 16 species (18%) occurred in five or more quadrats. In terms of abundance, the most numerous bee forage species were Lolium multiflorum (603 individuals), Dactylis glomerata (513 individuals), Rumex obtusifolius (433 individuals), Digitaria sanguinalis (293 individuals), and Pennisetum clandestinum (249 individuals).
The species exhibiting the highest densities were Lolium multiflorum, Dactylis glomerata, Helianthus sp., Rumex obtusifolius, Salvia rosmarinus, Tessaria absinthioides, and Plantago major, all with densities exceeding 45 individuals per 25 m2 (Table 2). The species recorded in the greatest number of quadrats were Digitaria sanguinalis, Trifolium repens, and Rubus urticifolius (11 quadrats each), followed by Dactylis glomerata (10 quadrats), Lolium multiflorum and Rumex obtusifolius (9 quadrats each), Pennisetum clandestinum (8 quadrats), Bidens pilosa, Baccharis salicifolia, Baccharis latifolia, and Philoglossa mimuloides (7 quadrats each), and Ranunculus repens (6 quadrats). All remaining species occurred in five or fewer quadrats.
Table 2.
Bee forage species recorded in the study area, including vegetation stratum, floral reward type, abundance, and density.
Regarding the vegetation strata of bee forage species, most species (76 species; 84%) belonged to the low stratum, whereas the medium stratum (10 species) accounted for approximately 11% and the high stratum (4 species) represented only 5% of the total (Figure 3a). Concerning floral resource type, 77% of the species (69 species) provided both nectar and pollen, whereas 23% (21 species) were classified as species classified as pollen sources for honey bees (Figure 3b).
Figure 3.
Distribution of bee forage species according to vegetation stratum (a), floral resource type (b), and generic composition (c).
Regarding the bee forage species, the most representative genera were Lolium (13%), Dactylis (11%), Rumex (9%), Digitaria (7%), Galinsoga (6%), Pennisetum (5%), Baccharis (4%), Bidens (4%), Plantago (3%), and Trifolium (3%). In contrast, Helianthus and Conyza each accounted for only 2% of the recorded species. The remaining genera collectively represented 40% of the total, with each individual genus contributing less than 2% (Figure 3c).
Among the bee forage species recorded in cropland, the most representative were Lolium multiflorum (15%), Dactylis glomerata (11%), Rumex obtusifolius (11%), Digitaria sanguinalis (7%), and Pennisetum clandestinum (7%). Other relevant species included Galinsoga parviflora (5%), Bidens pilosa (4%), Trifolium repens (3%), Galinsoga ciliata (3%), Plantago major (3%), and Conyza bonariensis (2%). Species with lower frequencies (<2% each) collectively accounted for 29% of the bee forage species recorded in cropland (Figure 4a).
Figure 4.
Relative distribution of the main bee forage species recorded in cropland (a) and forest areas (b).
In contrast, species with low representation (<3% each) collectively accounted for 49% of the bee forage species recorded in forest areas. The most representative species were Dactylis glomerata (8%), whereas Lolium multiflorum, Rubus urticifolius, and Baccharis latifolia each accounted for 5% of the records. Digitaria sanguinalis, Gnaphalium americanum, Drymaria cordata, and Geranium sessiliflorum each represented 4%, while Plantago major, Miconia papillosa, Oxalis corniculata, and Ageratum conyzoides each accounted for 3% of the recorded species (Figure 4b).
3.3. Species Accumulation Curves
The species accumulation curve for the entire general floristic composition showed that more than 110 species were recorded within the first 10 quadrats. Thereafter, the rate of species accumulation decreased substantially, with only 14 additional species being recorded in the remaining quadrats, resulting in a total of 124 species across the 24 sampled quadrats (Figure 5a). According to the Bootstrap richness estimator, the expected total species richness was 147 species. Therefore, the sampling effort captured approximately 84.4% of the estimated total species richness in the study area (124/147 × 100).
Figure 5.
Rarefaction-based species accumulation curves for total plant species (a) and bee forage species (b). Shaded areas indicate the 95% confidence intervals.
Regarding bee forage species, more than 75 species were recorded within the first 10 quadrats. Similar to the pattern observed for the general floristic composition, the rate of species accumulation decreased thereafter, with only 15 additional species being recorded in the remaining quadrats, resulting in a total of 90 species across the 24 sampled quadrats (Figure 5b). According to the Bootstrap richness estimator, the expected richness of bee forage species was 106 species. Therefore, the sampling effort captured approximately 84.9% of the estimated bee forage species richness in the study area (90/106 × 100).
3.4. Richness, Diversity, Evenness, and Dominance
No significant differences (p > 0.05) were observed in species richness or in the diversity, evenness, and dominance indices between land-cover types (forest and cropland), distances from the apiary, or transect orientations. The only exception was the Berger–Parker dominance index, which differed significantly among distances (p < 0.05). Dominance was highest in areas closest to the apiary (0.5 km), with a value of 0.46, whereas at greater distances (2–3 km) dominance values ranged from 0.23 to 0.25 (Table 3).
Table 3.
Mean (±SD) values of species richness, diversity, evenness, and dominance indices according to land-cover type, distance from the apiary, and transect orientation.
Species richness, diversity, evenness, and dominance indices of bee forage species were not significantly affected (p > 0.05) by land-cover type (cropland versus forest), distance from the apiary, or transect orientation (Table 4).
Table 4.
Mean (± SD) values of species richness, diversity, evenness, and dominance indices of bee forage species according to land-cover type, distance from the apiary, and transect orientation.
3.5. Species Composition Patterns Revealed by PERMANOVA and NMDS
PERMANOVA revealed that land-cover type and transect orientation significantly influenced the composition of both the general floristic composition and the bee forage species assemblage (p < 0.05). In contrast, distance from the apiary did not have a significant effect on species composition (p > 0.05) (Table 5).
Table 5.
PERMANOVA results for the general floristic composition and bee forage species, including F-statistics, coefficients of determination (R2), and p-values.
Post hoc pairwise PERMANOVA comparisons revealed that species composition differed significantly between cropland and forest for both the general floristic composition and the bee forage species assemblage (p < 0.05; Table 6).
Table 6.
Results of post hoc pairwise PERMANOVA comparisons between land-cover types for the general floristic composition and bee forage species.
Post hoc pairwise comparisons revealed significant differences in species composition among transects for both the general floristic composition and the bee forage species assemblage (p < 0.05). The only exception was the comparison between the south and east transects, which showed no significant differences for either dataset (p > 0.05) (Table 7).
Table 7.
Results of post hoc pairwise PERMANOVA comparisons of species composition among transects for the general floristic composition and bee forage species.
The NMDS ordination for the general floristic composition showed an acceptable goodness-of-fit, with a stress value of 0.1533, indicating a reasonable representation of multidimensional relationships in the two-dimensional configuration. Clear differences in species composition were observed between cropland and forest, although substantial within-group variability was also evident (Figure 6a). The north transect exhibited the most distinct species composition compared with the other transects. In contrast, the south and west transects showed partial separation, whereas the east transect was more similar to the south and west transects (Figure 6b). No clear differences in species composition were observed among distances from the apiary (Figure 6c).
Figure 6.
Non-metric multidimensional scaling (NMDS) ordination plots of the general floristic composition according to land-cover type (a), transect orientation (b), and distance from the apiary (c).
The NMDS ordination for bee forage species showed an acceptable goodness-of-fit, with a stress value of 0.1584, indicating a reasonable representation of multidimensional relationships in the two-dimensional configuration. Clear differences in species composition were observed between cropland and forest, although substantial within-group variability was evident (Figure 7a). The north transect exhibited a distinct species composition compared with the other transects. The west and south transects showed slight separation from one another, whereas the east transect was more similar to both the west and south transects (Figure 7b). No clear separation in species composition was observed among distances from the apiary (Figure 7c).
Figure 7.
Non-metric multidimensional scaling (NMDS) ordination plots of bee forage species according to land-cover type (a), transect orientation (b), and distance from the apiary (c).
4. Discussion
4.1. General Floristic Composition
The results revealed an interesting floristic richness in the study area (124 species), reflecting the structural complexity and heterogeneity of the landscape. This pattern is particularly relevant from a beekeeping perspective due to the high proportion of species identified as potential bee forage resources (approximately 73%) [119,120]. These findings indicate that the landscape is not only diverse but also functionally important for providing floral resources to honey bees.
The predominance of the low vegetation stratum, which accounted for approximately 86% of the recorded species, indicates that herbaceous plants represent a major component of the vegetation with potential relevance as bee forage resources in the study area. However, the contribution of these resources to the seasonal availability of forage cannot be determined from the present study, as flowering phenology was not monitored throughout the year [121]. Such vegetation is often associated with disturbed or managed landscapes and can enhance the temporal availability of floral resources for foraging honey bees [122].
From both ecological and productive perspectives, flowering plant diversity is closely associated with the availability of nectar and pollen, which are essential resources for the maintenance and development of A. mellifera colonies [123]. Landscapes with high floristic diversity generally support greater abundance and stability of pollinator communities [5]. Furthermore, the occurrence of numerous species in both cropland and forest indicates a certain degree of floristic connectivity within the landscape. Such connectivity may facilitate pollinator movement among habitat types and promote more efficient use of available floral resources, thereby contributing to the ecological stability of pollinator-dependent systems under changing environmental conditions [124].
4.2. Bee Forage Species
The identification of 90 plant species with literature-supported bee forage potential indicates that the study area contains a diverse pool of potentially available floral resources. However, their local use and relative importance to A. mellifera were not directly evaluated.
The high proportion of species providing both nectar and pollen (approximately 77%) represents an important functional advantage, as these species may contribute to a mzore balanced diet for honey bee colonies. The simultaneous availability of nectar and pollen supports larval development [125], colony energy requirements [126], and may help mitigate the effects of environmental stressors on colony performance [127]. These findings should be interpreted within the scope of the adopted classification, which considered nectar and pollen as the main floral rewards relevant to A. mellifera. Consequently, plant species providing other resources, particularly resins used for propolis production, were not included in this classification.
The high abundance and spatial occurrence of species such as Lolium multiflorum, Dactylis glomerata, Rumex obtusifolius, and Digitaria sanguinalis identify them as potentially important components of the local forage landscape. Given their abundance and frequency, these species may contribute substantially to the local availability of floral resources, particularly because several of them are documented pollen sources [128]. However, their actual contribution to honey bee nutrition and their relative use by A. mellifera were not directly assessed. Therefore, abundance within vegetation quadrats should not be interpreted as evidence of honey bee preference or as a proxy for the proportion of pollen collected from each species.
However, an excessive reliance on a limited number of dominant species may reduce the nutritional diversity available to honeybees [129]. In this context, the coexistence of less abundant yet functionally complementary species is essential to ensure a diverse and balanced diet [130]. Therefore, the conservation of nectar-producing species within the genus Baccharis is of particular importance, as they diversify the availability of floral resources and enhance the nutritional resilience of honey bee colonies.
Differences in species composition between cropland and forest indicate that both land-cover types contribute complementary floral resources to the landscape. Cropland tended to be characterized by a higher abundance of a few dominant species, whereas forest contributed additional bee forage species that increased landscape-level heterogeneity [131,132]. This pattern highlights the importance of maintaining both components of the landscape to sustain the diversity and temporal stability of floral resources available to honey bees. Consequently, identifying key bee forage species provides a useful basis for ecological restoration programs, the design of pollinator-friendly productive landscapes, and beekeeping planning strategies [133].
4.3. Species Accumulation Curves
The species accumulation curves indicate that the sampling effort was sufficient to capture most of the floristic richness present in the study area. The tendency of both the total plant species and bee forage species curves to approach an asymptote suggests that additional sampling would likely yield only a limited number of new species [134]. The pattern of rapid initial species accumulation followed by a gradual stabilization is characteristic of well-sampled communities and has been widely reported in biodiversity studies [135,136,137]. In the present study, the similarity between the total plant species and bee forage species accumulation curves further suggests that a large proportion of the local flora contributes to the floral resources available to honey bees.
These results confirm the representativeness of the sampling effort and provide a robust basis for interpreting patterns of species richness, diversity, and composition observed in the study area. Furthermore, the similarity between the total plant species and bee forage species curves indicates that a substantial proportion of the flora is associated with resources used by honey bees [138,139]. In this context, the ecosystem appears to function not only as a reservoir of plant biodiversity but also as an important provider of floral resources that support pollinator communities and beekeeping activities [140].
4.4. Richness and Diversity
The absence of significant differences in species richness and diversity between cropland and forest indicates that both land-cover types contribute similarly to the maintenance of floral resources within the landscape. From a beekeeping perspective, this finding suggests that agricultural systems, when characterized by a certain degree of heterogeneity, may provide floral resource diversity comparable to that of forest [141,142].
The relatively homogeneous distribution of species diversity throughout the 3 km foraging radius represents another important finding, as it suggests that honey bee colonies have access to similar levels of floral diversity across the landscape [143]. Such spatial consistency in floral resources may contribute to more stable foraging opportunities and reduce the likelihood of localized resource shortages [144].
The higher Berger–Parker dominance values observed near the apiary may indicate areas where floral resources are concentrated in a few highly abundant species, potentially influencing honey bee foraging patterns [145]. This information may be useful for optimizing apiary placement and improving landscape management strategies based on the spatial distribution of bee forage species. Overall, these findings suggest that both species diversity and the spatial arrangement of dominant bee forage species are important components of landscape functionality for beekeeping systems [146,147].
4.5. Species Composition Based on PERMANOVA and NMDS
The multivariate analyses revealed significant differences in species composition among land-cover types and transect orientations. Although species richness and diversity were similar across the landscape, the identity of species varied spatially. From a beekeeping perspective, this spatial variation may be advantageous because it increases the functional diversity of floral resources available to honey bees [148].
The floristic differentiation observed between cropland and forest suggests that each land-cover type contributes a distinct set of species to the landscape, thereby increasing resource complementarity. Such compositional diversity may promote a broader range of floral resources for honey bees and other pollinators and has been associated with greater ecological stability and resilience of pollinator communities under environmental stress [149].
The absence of a significant effect of distance from the apiary further supports the idea that floral resources are widely distributed throughout the landscape. This pattern suggests that potential bee forage species occur across the entire 3 km study radius rather than being concentrated in specific zones [150]. In contrast, the significant differences among transects indicate that local environmental conditions and landscape characteristics influence the spatial distribution of plant communities.
The conservation of heterogeneous landscapes that integrate both productive areas and forest is therefore of considerable importance [151]. Such landscape configurations can enhance plant biodiversity and increase the diversity of floral resources available to honey bees [152]. Consequently, detailed characterization of floristic composition provides valuable information for identifying key species and spatial patterns, representing an important tool for the development of management and conservation strategies aimed at promoting sustainable beekeeping systems [153].
4.6. Study Limitations and Future Research
Despite these findings, some limitations should be considered when interpreting the results. This study provides a floristic and spatial characterization of plant species that represent potential floral resources for A. mellifera within the landscape surrounding a managed apiary. The classification of species as potential pollen, nectar, or mixed resources was based on published literature rather than on direct validation through field observations of honey bee visitation or analyses of corbicular pollen, bee bread, or honey pollen. Therefore, plant abundance and spatial distribution should be interpreted as indicators of the potential availability of floral resources rather than as evidence of honey bee preference or actual resource use by local colonies.
In addition, vegetation was surveyed only during April and May 2025, precluding the assessment of flowering phenology, seasonal variation in resource availability, and potential forage gaps throughout the year. Likewise, resin-producing plant species were not assessed, which may have resulted in an underestimation of the diversity of resources potentially available to honey bee colonies. Furthermore, the study was conducted within the landscape surrounding a single apiary in Levanto, Chachapoyas. Although vegetation sampling was systematic and covered different distances and land-cover types, the results are specific to the study landscape and should not be extrapolated beyond the area evaluated.
Future studies should include multiple apiaries, year-round phenological monitoring, direct observations of honey bee foraging, and palynological or molecular approaches to validate plant use under local conditions. These approaches would help determine which plant species are actually visited and exploited by A. mellifera, characterize seasonal patterns of resource availability, and provide a more comprehensive assessment of the floral and resin resources available to honey bee colonies within the studied landscape.
5. Conclusions
The study area exhibited high floristic richness and a diverse assemblage of plant species reported in the literature as potential floral resources for Apis mellifera. Although species richness, diversity, evenness, and dominance did not differ significantly among land-cover types, distances, or transect orientations, PERMANOVA and NMDS analyses revealed significant spatial variation in species composition between cropland and forest and among transects. These findings indicate that the heterogeneous landscape provides a spatially diverse pool of potential floral resources for honey bees. Because this study was based on vegetation surveys, the results represent the potential availability of floral resources rather than direct evidence of plant use by Apis mellifera. Nevertheless, this work provides a valuable baseline for understanding the floristic composition surrounding managed apiaries in the Peruvian Amazon. Future studies incorporating direct observations of honey bee visitation, pollen-load or melissopalynological analyses, year-round phenological monitoring, and the assessment of resin-producing plants will be essential to validate local resource use and support evidence-based beekeeping management.
Author Contributions
Conceptualization, methodology, writing—review and editing, supervision, Y.M.B.-M. and A.E.-C.; data collection, methodology, investigation, K.G.-H. and L.T.-G.; formal analysis, validation, review J.A.S.-U.; J.L.M.-Q. and L.M.M.-C.; data curation, statistics, Y.P.; conceptualization, writing—review and editing, supervision, I.S.C.-C. All authors have read and agreed to the published version of the manuscript.
Funding
The APC was funded by the Vicerrectorado de Investigación—Universidad Nacional Toribio Rodríguez de Mendoza de Amazonas.
Institutional Review Board Statement
Not applicable.
Data Availability Statement
The original contributions presented in this study are included in the article. Further inquiries can be directed to the corresponding author.
Acknowledgments
The authors like to thank the Doctoral Program in Sciences for Sustainable Development of the Universidad Nacional Toribio Rodríguez de Mendoza de Amazonas. Also to the Consejo Nacional de Ciencia, Tecnología e Innovación Tecnológica (CONCYTEC) and the Programa Nacional de Investigación Científica y Estudios Avanzados (PROCIENCIA) within the framework of the call E033-2023-01-BM “Interinstitutional Alliances for Doctoral Programs”, under grant number (PE501084305-2023).
Conflicts of Interest
The authors declare no conflicts of interest.
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