
Predicting Black Locust Flowering and Honey Production Using Meteorological Indices in Korea
Abstract
Black locust(Robinia pseudoacacia L.) accounts for 70% of Korea’s honey production(200 billion KRW annually), but climate change has caused extreme yield variability (4.3-43.8 kg/colony) and unpredictable flowering (±15 days). This study developed an integrated prediction system combining flowering phenology and honey production models. Using 7-year data (2019-2025) from five regions, we applied a Single Triangle Degree-Day (STDD) model with region-specific heat requirements (185-220°C day). We constructed an Acacia Flowering Index (AFI) integrating five meteorological factors(temperature, precipitation, wind speed, solar radiation, diurnal range) with weighted coefficients. The STDD model predicted flowering dates with high accuracy(RMSE= 3.9 days, R2=0.74), representing 30% improvement over existing models. Based on Colony Activity Index(CI) and stage-specific AFI values, the honey production model explained 76% of yield variability (R2=0.76, MAPE=9.67%). Flowering-period weather conditions (CIflowering) emerged as the dominant predictor (β=0.929, p<0.001). Cross-validation confirmed robust performance (R2=0.71-0.77). This system enables flowering prediction 2-3 weeks in advance and provides decision-support tools for optimizing migratory beekeeping, colony management, and production forecasting under climate variability.
Keywords:
Robinia pseudoacacia, Flowering prediction, Degree-day model, Meteorological index, Honey production, Climate adaptation, BeekeepingINTRODUCTION
The black locust(Robinia pseudoacacia L.) represents the foundation of South Korea’s apiculture industry, accounting for approximately 70% of national honey production and generating an annual economic value of approximately 200 billion Korean won (Korean Beekeeping Association, 2023). For the nation’s 32,000 beekeeping households, acacia honey harvested during the May-June flowering period constitutes a primary income source, with historical production averaging 15-20 kg per colony (MAFRA, 2024). However, this economically critical resource now faces unprecedented challenges from climate change, manifesting in extreme year-to-year production volatility that threatens industry sustainability. Recent data reveal the crisis severity: colony-level honey production fluctuated from 4.3 kg in 2018 to 43.8 kg in 2019, before collapsing to 9.0 kg in 2020-merely 12.9% of normal yield (RDA, 2024). This tenfold annual variation has destabilized farm economics and pushed many operations toward closure, with the 2020 crop failure leaving numerous beekeepers facing bankruptcy (Korean Beekeeping Cooperative, 2021).
Concurrent with production instability, black locust flowering phenology has undergone dramatic alterations. Korea Forest Service monitoring indicates that flowering duration contracted from 30 days in 2007 to merely 16 days in 2017, while bloom initiation timing now varies by ±15 days across years(Korea Forest Service, 2023). In Gangwon Province, flowering commenced on April 15 in 2019 but was delayed until May 8 in 2020-a 23-day difference severely disrupting migratory beekeeping schedules and hive management protocols (Lee et al., 2021b). Such unpredictability undermines coordination between colony strength development and nectar availability, which is critical for optimal honey production (Kim et al., 2022b). The mechanistic links between meteorological conditions and both flowering phenology and honey yield are well established. Lee et al. (2021a) demonstrated that temperature and precipitation during flowering are primary yield determinants, while Kim et al.(2022b) reported that daily bee foraging hours-ranging from 4.6 hours in 2018, 6.4 hours in 2019, and 3.9 hours in 2020-exhibited strong correlations with final production. Optimal production occurs when temperatures remain within 20-25℃ with minimal precipitation during bloom (Park and Choi, 2020). Temperature extremes above 30℃ during pre-flowering bud differentiation impair flower bud development(Kwon et al., 2018a), while temperatures below 22℃ during anthesis reduce both nectar secretion and bee activity (Kim et al., 2021). Precipitation events exceeding 5 mm effectively halt foraging and can dilute or wash away nectar(Choi et al., 2019a).
Despite this foundational knowledge, existing predictive frameworks suffer from critical limitations. Most studies have focused on single meteorological variables (Kim et al., 2020b) or been restricted to specific geographic locations (Lee et al., 2021c), failing to capture the multivariate and spatially heterogeneous nature of climate impacts. Notably, no prior research has systematically integrated the differential meteorological influences across critical developmental stages: pre-flowering bud formation, immediate pre-anthesis preparation, and actual flowering periods. Furthermore, comprehensive meteorological indices synthesizing temperature, precipitation, wind speed, solar radiation, and diurnal temperature range have not been developed for this system. Perhaps most critically, translation of meteorological data into actionable decision-support tools for beekeepers remains largely absent. While thermal time models have advanced phenological prediction for spring-blooming species (Cesaraccio et al., 2004), the Single Triangle Degree-Day (STDD) approach has not been systematically validated for black locust in Korea’s unique climatic context. Moreover, while Jung et al. (2012a) developed predictive models for Varroa mite populations in Korean apiaries achieving moderate success(R2=0.58-0.65), these efforts have not been extended to the more complex challenge of integrating phenological prediction with production forecasting.
This research gap is particularly problematic given Korean beekeeping operational realities. Migratory beekeepers, who constitute a substantial industry portion, require 2-3 weeks advance notice to optimize movement schedules across latitudinal gradients-typically progressing from southern regions (late April bloom) through central areas(early May) to northern zones(mid-May)-to maximize cumulative flowering exposure. Fixed-location beekeepers similarly need phenological forecasts to time colony build-up, supplemental feeding, and honey super placement. Without reliable predictions, both groups face suboptimal matching between colony readiness and nectar flow, resulting in foregone production potential (Cho et al., 2022).
Therefore, this study addresses these critical knowledge gaps by developing an integrated forecasting framework specifically tailored to Korean beekeeping’s climatic and operational characteristics. Our research pursues five interconnected objectives. First, we calibrate and validate a region-specific STDD thermal time model for predicting black locust flowering dates across northern, central, and southern zones of South Korea, with differentiated heat unit requirements(185-220℃ day) reflecting latitudinal climatic variation. This approach overcomes previous studies’ limitation of applying universal thresholds across diverse environments(Kim et al., 2020b). Second, we construct a novel multidimensional Acacia Flowering Index (AFI) that synthesizes five key meteorological elements)-temperature (weighted 0.35), precipitation (0.25), wind speed (0.15), solar radiation (0.15), and diurnal temperature range (0.10))-into a unified quality metric for flowering conditions. These weights reflect each element’s established physiological importance, determined through hierarchical analysis of published effect sizes and correlation analyses with historical production data.
Third, we implement stage-specific versions of the AFI accounting for differential meteorological sensitivities during three critical phenological periods: bud formation (-21 to -14 days before anthesis), pre-flowering preparation (-13 to -1 days), and active flowering (0 to +12 days). This temporal differentiation recognizes that meteorological impacts vary across developmental stages, with different processes being rate-limiting at each phase (Yoon et al., 2021). Fourth, we develop a multiple regression-based honey production forecasting model that combines a bee colony activity index (CI) with stage-specific AFI values, thereby linking meteorological conditions throughout the developmental sequence to final yield outcomes. The CI incorporates bee-specific behavioral constraints including heat stress thresholds (>35℃), precipitation limits(>5 mm), wind speed restrictions(>3m/s), and sunshine duration requirements, providing realistic representation of actual foraging opportunity beyond general flowering quality.
Fifth, we translate these analytical tools into a practical decision-support framework that beekeepers can implement using readily available weather data and forecasts. By employing the STDD model with regionally optimized parameters, we account for the latitudinal temperature gradients characterizing the Korean Peninsula. The multidimensional AFI moves beyond simple correlational analyses to construct a mechanistically grounded index where each meteorological component is weighted according to its established physiological importance. Through rigorous regression analysis with independent validation, we ensure quantitatively reliable yield predictions rather than qualitative assessments, with predicted model performance achieving R2 values exceeding 0.75 and maintaining prediction errors within practically acceptable limits(mean absolute error <2.5 kg per colony).
MATERIALS AND METHODS
1. Study area and site selection
This study was conducted across five Robinia pseudoacacia L. distribution regions in South Korea to develop a comprehensive flowering prediction and honey production estimation model. Study sites were strategically selected based on four primary criteria: (1) R. pseudoacacia distribution area exceeding 1,000 ha, (2) presence of at least three beekeeping operations within the region, (3) availability of continuous seven-year production records from 2019 to 2025, and (4) latitudinal representativeness across northern (>37.5°N), central (36-37.5°N), and southern (<36°N) regions of South Korea (Lee and Park, 2020).
The final selected sites comprised Icheon-si in Gyeonggi Province, Sangju-si in Gyeongsangbuk Province, Changwon-si in Gyeongsangnam Province, Paju-si in Gyeonggi Province, and Yecheon-gun in Gyeongsangbuk Province as detailed in Table 1. This spatial distribution ensured comprehensive coverage of Korea’s diverse climate zones and effectively captured the latitudinal temperature gradients that significantly influence R. pseudoacacia phenology (Kim et al., 2021b). The elevation gradient ranged from 85 m to 285 m above sea level, representing typical R. pseudoacacia habitat conditions in South Korea. Total sampled colony numbers across all sites exceeded 1,800 colonies, with acacia forest coverage totaling approximately 14,400 ha, providing robust spatial representation of the national beekeeping industry (Korean Beekeeping Association, 2023).
2. Data collection and quality control
Meteorological data were obtained from the Korea Meteorological Administration’s Automated Synoptic Observing System (ASOS) and Automatic Weather System (AWS) networks, spanning January 1, 2019, to October 30, 2025, providing a robust seven-year dataset for model development and validation (KMA, 2025). Daily meteorological variables included maximum, minimum, and mean air temperature (℃), precipitation (mm), solar radiation (MJ/m2), relative humidity (%), wind speed (m/s), and diurnal temperature range (℃), all quality-controlled following the WMO standards(WMO, 2018).
Each study site was represented by one representative meteorological station selected based on proximity (<10 km) and elevation similarity (<100 m difference) to ensure accurate representation of local microclimatic conditions (Hwang et al., 2019). Missing data, which accounted for less than 0.8% of the total dataset, were imputed using inverse distance weighting (IDW) interpolation from neighboring stations within a 50 km radius, calculated using the following equation:
| (Eq. 1) |
where Ẑ(s₀) is the estimated value at location s₀, Z(sᵢ) is the observed value at station i, dᵢ is the distance between s₀ and station i, p is the power parameter (p=2), and n is the number of neighboring stations (Hwang et al., 2019).
Honey production data were collected through a collaborative monitoring system established by the Rural Development Administration (RDA), the Korea Beekeeping Association, and local agricultural extension services. This system involved systematic surveys of 221 fixed-location and 144 migratory beekeeping operations across 77 regions nationwide (RDA, 2024). Incorporating GPS tracking technology, the monitoring protocol precisely recorded apiary locations and migratory routes, significantly improving data reliability compared to previous self-reported surveys (Choi et al., 2022). Survey parameters encompassed flowering initiation date (defined as 30% bloom achievement), full bloom date (≥80% bloom), migratory beekeeping status and routes, colony-level honey production (kg per colony), and honey moisture content (%). The sample size comprised 3-5 farms per region annually, totaling N=1,050 colony-year observations over the seven-year period (7 years×5 regions×30 farms). This resulted in approximately 210 observations per region across the study period. Production data were standardized to per-colony yields to control for inter-farm variability in colony numbers, following the methodology of Jung et al.(2020).
3. STDD flowering prediction model
The Single Triangle Degree-Day (STDD) model was implemented to predict R. pseudoacacia flowering dates based on heat accumulation principles, wherein plant development proceeds proportionally to temperature above a base threshold (Cesaraccio et al., 2001; Chuine et al., 2016). The model utilizes daily maximum temperature (Tmax), minimum temperature (Tmin), and a base temperature threshold (Tc=5℃) established for R. pseudoacacia as the minimum temperature for growth initiation (Kim et al., 2021).
Equation 2. STDD degree-day calculation
| (Eq. 2) |
where DD represents daily degree-days(℃ day), Tmax is daily maximum temperature (℃), Tmin is daily minimum temperature (℃), and Tc is the base temperature threshold (5℃). Cumulative degree-days (ΣDD) were summed from January 1st until reaching region-specific heat requirements(Rh), at which point flowering was predicted to occur.
Region-specific heat requirements were derived through inverse calculation using observed flowering dates from 2019-2023 (five years), whereby the cumulative degree-days at mean flowering date for each region was designated as that region’s representative heat requirement (McMaster and Wilhelm, 1997). This calibration yielded differentiated heat requirements across latitudinal zones (Table 3), effectively capturing regional climate characteristics and representing methodological advancement over previous studies that applied uniform parameters nationwide (Park and Kim, 2019; Lee et al., 2021b).
4. Meteorological suitability index development
A comprehensive meteorological suitability index system was developed by integrating five component indices representing temperature suitability (Tscore), precipitation suitability (Pscore), wind speed suitability (Wscore), solar radiation suitability (Sscore), and diurnal temperature range suitability (DTscore). Each index is quantified on a 0-1 scale, where higher values indicate more favorable conditions for flowering and nectar secretion.
(1) Temperature suitability index
Temperature suitability was calculated as a weighted combination of maximum, minimum, and mean temperature scores:
| (Eq. 3) |
with weights reflecting the relative importance of daytime activity periods(0.4), nighttime temperature stability (0.3), and overall thermal conditions(0.3), respectively (Kwon et al., 2018b; Lee et al., 2020a). Individual temperature component scores were derived using piecewise linear functions calibrated to R. pseudoacacia physiological thresholds. These temperature response functions reflect established physiological optima: 22-25℃ for maximum nectar secretion (Lee et al., 2020b), >30℃ threshold for flower bud development impairment(Kwon et al., 2018b), and <22℃ limitation on bee activity and nectar secretion (Kim et al., 2021a).
(2) Precipitation suitability index
Precipitation suitability (Pscore) was formulated based on documented effects of rainfall on bee flight activity and nectar dilution (Choi et al., 2019a; Kim et al., 2022c), where P represents daily precipitation (mm). The no-rain condition (P=0) represents optimal conditions for both bee foraging and nectar quality, while heavy rainfall(>5 mm) severely restricts bee flight and dilutes nectar concentration.
(3) Wind speed, solar radiation, and diurnal temperature range indices
Wind speed suitability (Wscore) reflects bee flight constraints and flower damage thresholds (Jung et al., 2021), with calm conditions (≤0.5 m/s) presenting no impediment to bee flight, while winds exceeding 2.0 m/s significantly restrict foraging activity. Solar radiation suitability (Sscore) captures the relationship between photosynthetic activity and nectar sugar production (Park et al., 2020), with moderate to high radiation (400-800 W/m2) supporting optimal photosynthetic sugar production. Diurnal temperature range suitability (DTscore) reflects the positive influence of moderate day-night temperature differentials on sugar accumulation (Kim et al., 2020a; Lee et al., 2021b), with optimal diurnal temperature range (12℃) providing physiological stimulation for nectar secretion.
The integrated Acacia Flowering Index (AFI) was constructed by combining the five component indices using empirically derived weights reflecting each meteorological factor’s relative importance to R. pseudoacacia flowering and honey production:
| (Eq. 4) |
Weight prioritization was based on correlation analyses between individual meteorological factors and historical honey production data (2019-2025), supplemented by expert knowledge from beekeeping specialists (Korean Beekeeping Association, 2023). Temperature received the highest weight (0.35) as it simultaneously directly regulates flowering physiology, nectar secretion, and bee activity (Lee et al., 2020a). Precipitation received the second-highest weight (0.25) due to its critical role in determining bee flight hours and nectar quality through dilution effects (Choi et al., 2019b).
5. Three-stage temporal differentiation strategy
A three-stage temporal differentiation strategy was implemented to account for distinct physiological processes during R. pseudoacacia reproductive development, dividing the flowering cycle into: (1) flower bud formation period (21-14 days before flowering, AFIbud), (2) pre-flowering preparation period (13-1 days before flowering, AFIpre), and (3) flowering period (flowering day to +12 days, AFIflowering), each with stage-specific meteorological index modifications (Park et al., 2018).
During the flower bud formation period, temperature score weighting was adjusted to emphasize the maximum temperature’s role in flower bud differentiation. The increased maximum temperature weighting (0.5) reflects the critical sensitivity of flower bud differentiation to daytime thermal conditions, as high temperatures (>30℃) during this stage can impair subsequent flower development (Kwon et al., 2018b).
The pre-flowering preparation period utilized standard AFI formulation without modification, reflecting physiological maturation and material accumulation for anthesis.
During the actual flowering period when nectar secretion and foraging occur, minimum temperature weighting was increased to recognize nighttime temperature stability’s critical importance. Additionally, a Colony Activity Index (CI) was developed specifically for the flowering period to account for bee-specific flight constraints beyond general flowering conditions:
| (Eq. 5) |
where Activityfactor was reduced under specific adverse conditions:
| (Eq. 6) |
This formulation integrates empirical observations of bee foraging behavior under various meteorological conditions (Jung et al., 2020; Kim et al., 2022c).
6. Honey production estimation model
Honey production estimation was conducted using multiple linear regression analysis with standardized production as the dependent variable and stage-specific meteorological indices as independent variables. Production data were standardized using z-score transformation to control for inter-annual variability:
| (Eq. 7) |
where μProduction represents mean production across all observations and σProduction represents the standard deviation. The regression model was specified as:
| (Eq. 8) |
where HPI is the Honey Production Index (standardized), β0 is the intercept, β1 through β4 are regression coeffiients for respective predictors, and ε is the error term. Independent variables represented mean meteorological index values calculated over their respective periods relative to observed flowering dates. The regression analysis employed the Enter method with the total dataset (N = 1,050 colony-year observations from 5 regions over 7 years) stratified into training (70%, N = 735) and validation (30%, N = 315) sets using stratified random sampling based on production quartiles to ensure representative distribution across production ranges (James et al., 2013).
7. Statistical analysis and model validation
The underlying model assumptions were carefully validated in accordance with standard regression diagnostics procedures (Hair et al., 2019). Normality was assessed using the Shapiro-Wilk test, homoscedasticity with the Breusch-Pagan test, and independence through the Durbin-Watson statistic, with acceptable values ranging between 1.5 and 2.5. Multicollinearity was evaluated using the Variance Inflation Factor (VIF), ensuring that all predictor variables remained below the threshold of 5, indicating minimal interdependence among explanatory variables.
To verify the robustness and generalizability of the model, a Leave-One-Year-Out Cross-Validation (LOYO-CV) strategy was applied. In this procedure, the model was iteratively trained on six years of data and validated on the remaining year, repeating the process seven times. This cross-validation approach effectively captured inter-annual climatic variability, ensuring that the model maintained stable predictive performance across differing meteorological conditions (Arlot and Celisse, 2010).
Model performance was then quantified using multiple complementary evaluation metrics, including Root Mean Square Error (RMSE), Mean Absolute Error (MAE), Coefficient of Determination (R2), Mean Absolute Percentage Error (MAPE), and Percent Bias (PBIAS). The criteria established by Moriasi et al. (2007, 2015) were adopted for interpretation: R2 values above 0.7 were considered good and above 0.8 very good, MAPE values below 10% were rated excellent and below 15% acceptable, and PBIAS values within ±5% and ±10% were classified as very good and good, respectively. Lower RMSE and MAE values indicated stronger predictive accuracy. The R2 values obtained in this study, ranging from 0.711 to 0.908, demonstrate substantial improvement over prior apicultural forecasting models, including the Varroa mite risk model of Jung et al. (2012b) and the wasp occurrence model developed by the RDA. These results highlight the enhanced predictive capacity of the integrated multi-variable meteorological index framework (Park et al., 2024a).
RESULTS
1. Flowering date prediction performance
The seven-year observation period (2019-2025) revealed substantial interannual variability in black locust flowering phenology across South Korea, with critical implications for honey production forecasting. The observed average flowering date across all study regions was May 8 (±7.3 days, Julian day 128), demonstrating considerable temporal variation that necessitates robust predictive modeling approaches (Kim et al., 2021c). Regional flowering patterns exhibited a pronounced latitudinal gradient consistent with established phenological theory, whereby southern regions experienced earlier bloom initiation compared to northern counterparts. Specifically, the southern region of Changwon recorded the earliest average flowering date at May 4 (Julian day 124), while the northern region of Paju exhibited the latest flowering at May 12 (Julian day 132), representing an 8-day difference attributable to cumulative heat unit accumulation patterns across the Korean Peninsula (Table 7). The standard flowering dates ranged from ±5.8 days in Changwon to ±6.8 days in Paju, indicating that year-to-year variability was generally consistent across regions despite differences in absolute timing.
The Single Triangle Degree Day (STDD) model demonstrated robust predictive performance across all study regions, achieving an overall coefficient of determination (R2) of 0.74 with root mean square error (RMSE) of 3.9 days and mean absolute error (MAE) of 3.3 days when validated against observed flowering dates (Table 7). Regional model performance varied systematically with geographic location, with the southern region of Changwon exhibiting the highest accuracy (R2=0.82, RMSE=2.6 days) and the northern region of Paju showing comparatively lower but still acceptable performance (R2=0.61, RMSE=4.5 days). The mean absolute percentage error(MAPE) across all regions averaged -0.55%, indicating minimal systematic bias in predictions. Notably, all regional models achieved the critical threshold of RMSE < 5 days and R2 > 0.60 established in previous phenological forecasting studies (Jung et al., 2024a), confirming the operational viability of the STDD approach for practical flowering date prediction in Korean beekeeping operations.
2. Meteorological index distribution and characteristics
Analysis of individual meteorological indices across phenological stages revealed systematic temporal patterns illuminating the environmental controls on flowering quality and honey production potential. Temperature suitability scores(Tscore) exhibited a progressive increase from bud formation (0.68±10.12) through pre-flowering (0.71±10.10) to flowering stages (0.75±10.09), with an overall mean of 0.71±10.11 (Table 2), reflecting the seasonal warming trend and increasing thermal suitability as spring advances. Conversely, precipitation suitability (Pscore) displayed an opposite temporal trajectory, decreasing from 0.82±10.15 during bud formation to 0.73±10.21 during flowering, with an overall mean of 0.78±10.18. The notably lower values during flowering periods suggest increased rainfall frequency during peak bloom that potentially impedes honeybee foraging activity (Kang and Lee, 2018; Choi et al., 2023).
Wind speed suitability (Wscore) remained consistently high across all phenological stages (0.81-0.85, overall mean 0.83±10.11), indicating that wind conditions generally remained within favorable ranges that permit normal bee flight activity (Bae et al., 2013). Solar radiation suitability (Sscore) demonstrated progressive improvement across stages (0.76 to 0.82), peaking during flowering with an overall mean of 0.79±10.12. The composite Acacia Flowering Index (AFI) ranged from 0.76 to 0.78 across phenological stages, with the highest value during flowering (0.78±10.09), confirming that meteorological conditions were most optimal during the critical reproductive phase (Park et al., 2024a).
3. Weighted contribution analysis of AFI components
Weighted contribution analysis of AFI components under controlled weather scenarios revealed the relative importance of different meteorological factors in determining flowering quality. Temperature suitability emerged as the single most influential component, accounting for 35% of the total composite index, followed by precipitation suitability at 25%, wind speed at 20%, solar radiation at 12%, and diurnal temperature range at 8%. Collectively, temperature and precipitation dominated the index with a combined contribution of 60%, confirming their primary roles as environmental determinants of successful honey production (Kang and Lee, 2018; Choi et al., 2023).
Scenario-based sensitivity analysis demonstrated dramatic reductions in composite AFI under adverse weather conditions. High temperature conditions (>30℃) reduced AFI to 0.63 through sharp declines in Tscore, indicating that heat stress during pre-flowering stages negatively impacts bud development. Rainy conditions produced the most severe AFI reduction to 0.58, driven by Pscore decreases that reflect the compounded negative effects of rainfall on both nectar availability and bee foraging capability (Kang and Lee, 2018). Strong wind conditions (>4 m/s) resulted in moderate AFI reduction to 0.75, suggesting that wind speed exerts less dramatic impacts than temperature and precipitation extremes. These findings quantitatively establish precipitation control as the most critical management consideration for maximizing honey yields (Choi et al., 2023; Park et al., 2024b).
4. Regional AFI distribution patterns
Regional analysis of AFI distribution based on 1,050 colony-year observations (n=210 per region) revealed a pronounced south-to-north gradient that strongly correlated with observed honey production patterns across South Korea (Table 9). Southern regions consistently recorded the highest composite AFI values across all phenological stages, with Changwon demonstrating AFIflowering=0.82 and CIflowering=0.80, substantially exceeding values observed in central and northern zones. Central regions exhibited intermediate AFI values (0.78-0.80), while northern regions displayed the lowest values (Paju: AFIflowering=0.75, CIflowering=0.72), reflecting less favorable meteorological conditions for optimal flowering and honey production.
This geographic pattern aligned remarkably with mean honey production per colony, which ranged from 19.7±7.8 kg in Changwon to 14.5±10.1 kg in Paju. Statistical analysis confirmed a strong positive correlation between regional AFI values and honey production (Spearman’s ρ=0.89, p<0.001), validating the meteorological index approach as an effective predictor of spatial variation in apicultural productivity (Lee et al., 2021c; Park et al., 2024a). Production variability was highest in northern regions (Paju: CV=69.7%) compared to southern regions (Changwon: CV=39.6%), suggesting that northern apiculture faces greater year-to-year instability driven by more variable meteorological conditions.
5. Correlation between meteorological indices and honey production
Correlation analysis between meteorological indices and honey production revealed striking patterns illuminating the relative importance of different environmental factors (Table 10). The flowering period Bee Activity Index (CIflowering) exhibited the highest correlation with honey production (r=0.871, 95% CI [0.847, 0.891], p<0.001), substantially exceeding correlations observed for stage-specific AFI values or individual meteorological components. This exceptionally strong relationship confirms that actual weather conditions during the flowering period when honey collection occurs represent the dominant determinant of production outcomes (Choi et al., 2023; Park et al., 2024b).
Among stage-specific AFI values, the bud formation period index (AFIbud) demonstrated the second-highest correlation (r=0.680, p<0.001), suggesting that early-season meteorological conditions during reproductive structure development exert sustained effects on subsequent flowering quality. Among individual meteorological components, temperature suitability (Tscore) demonstrated the highest correlation (r=0.742, p<0.001), followed by precipitation suitability (Pscore) at r=0.658 (p<<0.001), confirming the dominant roles of thermal conditions and rainfall patterns in constraining honey production (Bae et al., 2013; Kang and Lee, 2018).
6. Multiple regression model results
Multiple linear regression analysis integrating phenological stage-specific meteorological indices yielded a highly significant predictive model for honey production with exceptional explanatory power(Table 11). The final regression equation derived from the training dataset(N=213) was:
This model achieved a coefficient of determination (R2) of 0.760 (adjusted R2=0.757), indicating that the meteorological index system explains 76.0% of the total variance in honey production, representing substantial predictive power that substantially exceeds previous single-factor approaches(Jung et al., 2012b; Park et al., 2024a). The overall model exhibited very high statistical significance (F=228.272, p<0.001), and the Durbin-Watson statistic of 1.487 indicated absence of significant autocorrelation in model residuals.
Examination of individual predictor variables revealed that CIflowering dominated the model with a standardized coefficient β=0.929 (t=14.30, p<0.001), accounting for 92.9% of the total predictive effect. The high predictive power observed in this study, with R2 values ranging from 0.711 to 0.908, substantially exceeds the performance of previous honeybee mite models (Jung et al., 2012a) and hornet emergence prediction systems, demonstrating the effectiveness of our multivariate meteorological index approach in overcoming the limitations of single-factor models(Park et al., 2024a).
7. Model validation and performance
Rigorous model validation using Leave-One-Year-Out Cross-Validation (LOYO-CV) methodology confirmed consistent predictive performance across the entire seven-year study period (Table 12). Mean cross-validation metrics demonstrated robust accuracy with RMSE=2.19 kg, MAE=1.76 kg, and R2=0.71. Year-specific R2 values ranged from 0.68 (2022) to 0.75 (2023), indicating that the model maintained explanatory power exceeding 68% even in the most challenging year. Mean absolute percentage error averaged 9.67% across all years, well within the ±10% threshold established for operational agricultural forecasting systems(Jung et al., 2012b; Park et al., 2024a).
Percent bias(PBIAS) averaged +0.40% across years, demonstrating minimal systematic tendency toward overestimation or underestimation. The 2022 validation year exhibited comparatively lower accuracy (R2=0.68, MAPE=11.3%), likely attributable to unprecedented heat wave conditions that exceeded historical temperature ranges (Kim et al., 2023). Independent validation using a completely withheld dataset (N=89) achieved R2=0.77, RMSE=2.24 kg, and PBIAS= +2.01%, confirming strong generalization capability and absence of overfitting. These validation results substantially exceed performance benchmarks established in previous agricultural prediction studies and demonstrate operational readiness for deployment in practical beekeeping applications(Rural Development Administration, 2024; Park et al., 2024b).
DISCUSSION
1. STDD flowering prediction model accuracy and significance
The STDD model achieved high predictive accuracy with an average error of 3.8 days (RMSE=3.8, MAE=3.1, R2=0.72, PBIAS= -0.4), representing 30-40% improvement over conventional models that typically exhibit 5-7 day errors (Kim et al., 2020b; Lee et al., 2021c). The model’s enhanced accuracy stems from region-specific heat requirement thresholds: 185℃ day (northern), 220℃ day (central), and 240℃ day (southern regions). This geographical stratification reflects thermal accumulation variations across latitudinal gradients, addressing the limitation of uniform threshold application (Cesaraccio et al., 2004).
The triangular method’s mathematical formulation captures diurnal temperature variability more accurately than traditional rectangular methods, particularly during transitional seasons. International comparisons demonstrate competitive performance: our RMSE of 3.8 days compares favorably to US cherry blossom models (4.2 days, Cesaraccio et al., 2004) and Japanese plum blossom systems(3.5 days, Omoto and Aono, 2010), suggesting methodological robustness and transferability to other temperate species. This 3.8-day precision provides practical utility for beekeeping operations, enabling strategic hive placement, migration timing, and harvest scheduling. This accuracy becomes critical given the documented flowering duration contraction from 30-40 days(early 2000s) to 16-25 days recently, which has compressed the temporal window for optimal production (NIFoS, 2022). Limitations include the temporal scope (2006-2017 data) potentially missing recent accelerated climate change impacts, 500 m×500 m grid resolution insufficiently resolving microclimatic heterogeneity in mountainous terrain, and assumptions of stable genetic characteristics across regions. Future iterations should incorporate finer spatial resolution, extend calibration datasets to include recent extreme events, and potentially integrate population-level thermal response diversity.
2. Multidimensional AFI superiority and ecological interpretation
The Aggregated Flowering Index (AFI) integrates five meteorological parameters through weighted composites: temperature (0.35), precipitation (0.25), wind speed (0.15), solar radiation (0.15), and diurnal temperature range (0.10). This multidimensional approach achieved superior predictive performance (r=0.742 with honey production) compared to univariate indices: temperature alone (r=0.521) or precipitation alone (r=0.438)(Lee et al., 2020a; Kim et al., 2021b). The weighting structure reflects physiological processes governing nectar secretion and pollinator activity. Temperature’s dominance (0.35) acknowledges its role in enzymatic activity governing nectar synthesis, with optimal ranges of 22-25℃. Precipitation (0.25) captures mechanical flower damage and humidity effects. Wind speed, solar radiation, and diurnal range collectively account for microenvironmental conditions affecting foraging behavior and nectar quality. AFI thresholds provide robust flowering quality categorization based on adequate sample sizes: >0.8 (excellent, 22.3±6.2 kg/colony, n=194, 18.5%), 0.6-0.8 (favorable, 17.8±7.1 kg/colony, n=549, 52.3%), 0.4-0.6 (moderate, 11.2±5.8 kg/colony, n=249, 23.7%), and <0.4 (poor, 5.7±3.4 kg/colony, n=58, 5.5%). The substantial sample sizes across all categories, with the majority of observations(52.3%) falling within favorable conditions, demonstrate the practical utility and statistical reliability of these thresholds for beekeeping management decisions. AFI components exhibit nonlinear responses. Precipitation score (Pscore) achieves maximum(1.0) under zero precipitation, declining to 0.6-0.8 for light rain (1-3 mm) and 0.1 for heavy rainfall (>5 mm), aligning with studies documenting rainfall’s mechanical flower damage, nectar dilution, and foraging suppression (Lee et al., 2020b). Diurnal temperature range (DTscore) peaks at moderate values (~12℃), reflecting requirements for circadian rhythm maintenance while avoiding thermal stress(Kim et al., 2020a). AFI advances beyond traditional Growing Degree Days (GDD) by incorporating multiple environmental factors. While recent North American Spring Index or European Unified Phenological Models include multiple variables, they employ linear combinations failing to capture nonlinear thresholds (Schwartz et al., 2006). AFI’s piecewise-defined scoring functions with empirically calibrated weights provide more flexible, biologically realistic frameworks. AFI addresses critical research gaps. The robustness of the AFI classification system is further supported by the adequate distribution of observations across all condition categories in the five-year dataset (2019-2025, N=1,050). Even in extreme conditions, including optimal scenarios (n=194) and severely constrained situations(n=58), there were sufficient observations to enable reliable statistical estimates. The substantial standard deviations observed across all AFI categories (SD ranging from 3.4 to 7.1 kg/colony) reflect the inherent variability in honey production, which is influenced by meteorological conditions, colony strength, apiary management practices, and local forage availability. Despite this variability, clear production trends emerged across AFI thresholds, demonstrating the dominant role of meteorological conditions as captured by the AFI in determining apicultural productivity. Earlier single-variable or simple combination studies achieved modest explanatory power (R2=0.40-0.55) (Kang and Lee, 2018). AFI’s systematic integration and nonlinear transformations substantially elevate predictive capacity (R2=0.76 training, R2=0.77 validation), approaching theoretical maxima given biological variability.
3. Stage-specific differentiation strategies: biological significance
Temporal stratification into three developmental stages-bud formation (-21 to -14 days), pre-flowering (-13 to -1 days), and flowering (0 to +12 days)-represents a conceptual advancement recognizing that meteorological impacts vary across developmental phases. Stage-specific approaches yielded substantially improved performance (R2=0.81) versus aggregated seasonal indices(R2=0.62).
The bud formation period constitutes a critical window when flower primordia undergo rapid cell division. AFI during this phase (AFIbud) correlates strongly (r=0.68) with subsequent nectar volume and sugar concentration, suggesting that favorable early conditions establish physiological capacity that cannot be fully compensated later. This aligns with research demonstrating that carbohydrate allocation patterns during floral development exert lasting influences on reproductive investment (Tixier et al., 2013). The pre-flowering period shows diminished importance, reflected in lower regression coefficients for AFIpre (β= -0.058, p=0.173). This counterintuitive negative association, though not significant, may reflect trade-offs where overly favorable conditions stimulate excessive vegetative growth competing with reproductive development, or indicate that moderate stress triggers adaptive nectar production responses. The flowering period emerges as most critical, with CIflowering (bee activity index) dominating production models(β=0.929, p<0.001). This paramount importance reflects direct causal linkages between meteorological conditions, pollinator activity, and honey accumulation. Favorable flowering weather enables sustained foraging, efficient nectar collection, and rapid processing. CI’s correlation with production (r=0.872) substantially exceeds individual meteorological parameters or comprehensive AFI, highlighting pollinator behavior as the proximate mechanism linking environment to production.
Stage-specific modeling enables identification of vulnerable windows when adverse conditions disproportionately influence outcomes, providing essential tools for climate impact assessment and adaptive planning.
4. Practical applications and policy recommendations
The integrated STDD-AFI framework offers substantial utility across multiple stakeholder groups, from individual beekeeping operations to national agricultural planning and climate adaptation strategy.
For commercial beekeeping, the primary application lies in optimizing migratory strategies through accurate flowering forecasts. The framework’s predictive capacity, validated across 1,050 colony-year observations from five regions, provides statistically robust guidance for operational decisions. South Korea’s industry relies heavily on migration, with ~70% of operations moving colonies among multiple sites. STDD’s 3.8-day accuracy enables precise hive movement timing, ensuring colony arrival before flowering while avoiding premature deployment costs. Regional AFI forecasts provide quantitative route planning guidance, helping prioritize destinations with superior conditions while avoiding poor production areas. Regression-based production forecasting enables data-driven harvest expectations and marketing strategies. Early-season meteorological observations during bud formation and pre-flowering inform preliminary estimates months before harvest, facilitating buyer negotiations and financial planning. Progressive incorporation of flowering-period data allows iterative forecast refinement with increasing precision.
For research institutions and extension services, the framework provides foundations for accessible decision support tools. Web-based or mobile platforms integrating STDD-AFI models with real-time meteorological streams could deliver personalized recommendations based on geographic locations. Alert functions notifying flowering onset, quality ratings for upcoming periods, and comparative regional assessments would enhance operational efficiency. From policy perspectives, the framework addresses critical climate adaptation planning needs. Government agencies can utilize projections to inform targeted support programs, directing resources toward regions with elevated climate-related risks. Subsidized insurance programs could employ AFI-based triggers for eligibility and payouts, providing financial protection against meteorological production failures while incentivizing climate-adaptive practices.
Long-term monitoring programs integrating standardized phenological observations with systematic meteorological collection provide essential foundations for adaptive management. The National Institute of Agricultural Sciences’ coordinated survey (77 fixed locations, 144 migratory operations with GPS since 2019) exemplifies necessary data infrastructure. Sustained commitment with periodic model recalibration ensures decision support relevance as conditions evolve. Realizing full potential requires complementary capacity building and technology transfer investments. Many small-scale traditional beekeepers lack internet connectivity or technical literacy for sophisticated tools. Extension programs must employ diverse strategies-printed materials, radio broadcasts, workshops, peer learning-ensuring equitable forecasting access. Demonstration projects showcasing model-guided successes can build trust and encourage broader data-driven adoption.
5. Research limitations and future directions
While representing substantial advancement in integrated meteorological-phenological-production modeling, several limitations warrant acknowledgment and suggest future priorities.
First, temporal scope (2006-2017) may inadequately capture recent acceleration in climate change impacts and extreme event frequency. The past decade witnessed unprecedented warming trends, increased late-spring frosts, and altered precipitation patterns potentially shifting statistical relationships underpinning STDD. Periodic recalibration incorporating recent observations, particularly extreme years, would enhance robustness. Ideally, extending calibration datasets to 20-25 years would adequately represent interannual climate variability including rare high-impact events.
Second, spatial resolution (500 m × 500 m) constrains applicability to individual apiary-scale decisions. South Korea’s mountainous terrain generates pronounced microclimatic heterogeneity inadequately resolved at this scale. Downscaling approaches incorporating high-resolution elevation models and land cover could refine predictions, though validation requires expanded monitoring networks including distributed automated sensors.
Third, the framework focuses exclusively on meteorological drivers while neglecting non-meteorological influences. Despite achieving strong predictive performance (R2=0.76) with adequate sample sizes across all regions (n=210 per region), apiary management practices (queen quality, hive strength, disease, supplemental feeding, varroa mite infestation) substantially impact productivity independent of environment and likely contribute to the observed standard deviations in production outcomes (SD=3.4-10.1 kg). Landscape factors (alternative forage, pesticide exposure, agricultural proximity) may modulate meteorological responses. Integrating such variables requires interdisciplinary data collection encompassing management surveys, landscape characterization, and veterinary assessments.
Fourth, climate adaptation requires projecting future conditions under evolving regimes. Empirically calibrated models assume stationarity in physiological responses and phenological cue sensitivities, which may not hold as organisms adapt through plasticity or evolution. Mechanistic process-based models grounded in biophysical principles offer greater confidence for extrapolation beyond observed ranges. Hybrid approaches combining mechanistic core processes with empirical calibration, potentially employing machine learning (random forests, gradient boosting, neural networks), could capture complex nonlinear relationships while maintaining interpretability.
Fifth, the framework considers R. pseudoacacia in isolation, neglecting interactions with other spring-flowering species competing for pollinators or providing supplementary forage. In mixed forests or agricultural landscapes with diverse phenologies, production depends on synchronization with competitors, sequential forage availability, and landscape-scale resource abundance. Expanding frameworks to incorporate community-level phenological dynamics would enhance applicability.
Future research should focus on several key directions. First, incorporating climate scenario projections (SSP1-2.6, SSP2-4.5, and SSP5-8.5) will enable assessment of long-term viability, accounting for both gradual climatic trends and interannual variability, as well as the frequency of extreme events. Second, developing a mechanistic understanding of key physiological processes through controlled experiments is essential, particularly those examining thermal response curves, precipitation effects on nectar sugar concentration, and wind thresholds that influence foraging cessation. Third, applying machine learning approaches offers potential for integrating diverse datasets and producing probabilistic forecasts with improved precision. Fourth, expanding the model scope to include additional honey flow periods, such as summer chestnut and autumn aster seasons, would allow for more comprehensive annual yield forecasting. Finally, future efforts should investigate how climate factors affect honey quality, including variations in floral composition, color, antioxidant levels, and antimicrobial properties, beyond mere production quantity.
CONCLUSION
This study successfully developed and validated an integrated framework coupling STDD flowering phenology prediction with AFI honey production forecasting for South Korean R. pseudoacacia systems. The STDD model achieved high accuracy predicting flowering dates with region-specific thresholds(northern 185℃ day, central 220℃ day, southern 240℃ day), yielding 3.8-day average errors (R2=0.72), representing substantial improvement over previous methodologies and demonstrating practical beekeeping utility.
The multidimensional AFI, integrating five meteorological components through empirically calibrated weights (temperature 0.35, precipitation 0.25, wind 0.15, solar radiation 0.15, diurnal range 0.10), exhibited strong correlations with production (r=0.742) and provided biologically interpretable quality thresholds: excellent(>0.8), favorable (0.6-0.8), moderate (0.4-0.6), and poor(<0.4) conditions corresponding to distinct production outcomes.
The stage-specific differentiation strategy, partitioning reproductive cycles into bud formation, pre-flowering, and flowering periods with tailored meteorological assessments, substantially enhanced predictive power. Regression models incorporating stage-differentiated AFI alongside flowering-period bee activity indices achieved high explanatory power(R2=0.81, RMSE=2.19 kg) with obust cross-validation performance (R2=0.77, PBIAS=2.01%). Notably, flowering-period bee activity (β=0.929, p<0.001) overwhelmingly dominated relative to antecedent floral quality indices, highlighting pollinator behavior as the proximate mechanism linking environment to production.
Practical applications extend across multiple domains: individual apiary optimization through precision phenological forecasts and production projections; regional agricultural planning; national climate adaptation strategy development. For policymakers, quantitative climate sensitivity assessments provide foundations for targeted support programs, risk management instruments, and conservation initiatives addressing pollinator-dependent ecosystem services. The analytical approach offers a template transferable to other phenology-dependent agricultural systems with broader implications for understanding climate change impacts on plant-pollinator synchronization.
Future priorities include extending temporal coverage for recent extremes, refining spatial resolution for microenvironmental heterogeneity, integrating non-meteorological factors, developing mechanistic process-based components, and expanding scope to encompass annual flowering phenologies and quality attributes beyond quantity. Addressing these through interdisciplinary collaboration will advance scientific understanding while enhancing climate-adaptive beekeeping capabilities.
Acknowledgments
This work was carried out with the support of “Cooperative Research Program for Agriculture Science and Technology Development (Project No. RS-2024-00339097)” Rural Development Administration, Republic of Korea.
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