Pest infestation levels are subject to various biotic and abiotic factors that impact on the population dynamics (Khaliq et al. 2014). The coincidence of risk factors in space and time is decisive in either raising or lowering damage potential an insect pest. This coincidence can be favorably exploited in environmental control (Teetes 1991). This study tested the combined effects of three different risk factors, (i) flowering of the host plant, (ii) arrival of first male moths in pheromone monitoring traps and (iii) crop distance, as potential predictors, defining C. nigricana pest status in peas. Low infestation was mainly associated with the co-occurrence of an asynchronous flower onset outside a certain GDD range, late arrival time of the first moths in the actual fields, and an increased distance between the previous and the currently growing pea field, as expressed by MD.
Infestation risk was significantly related to the onset of flowering (BBCH 60). Pea varieties that started flowering specifically in a GDD range of about 900 GDD showed the highest attack levels, exceeding 40% (Fig. 1). This fact also explained the low infestation in the first (2016) and final year (2019) of the study, when all monitored fields started flowering only before or only after this particular GDD timespan. This flowering pattern resulted in a significantly lower mean infestation when compared to 2017 and 2018. During flower development, pea plants are olfactorily more attractive to mated C. nigricana females than during other development stages, for example, when leaves or pods are forming (Thöming et al. 2014). In line with the preference-performance theory for phytophagous insects, female pea moths’ preference reflects offspring performance (Gripenberg et al. 2010). The arrival of pea moths in pea fields during this critical developmental stage is important because pod formation starts shortly thereafter, and pods are the nutrient source for C. nigricana larvae. The importance of flowering time and duration for potential pea-moth damage has been reported by several authors (Hanson and Webster 1936; Nolte and Adam 1962). As an environmental control measure (Pimentel and Goodman 1978; Teetes 1991), early sowings in combination with the use of an early flowering variety was the most recommended approach to prevent pea-moth damage in green vegetable peas (Wright et al. 1951; Thöming et al. 2011). Conversely, a similar effect was reported for sowing dates that were significantly late (Anonymous 1948). Thus, even when only flower onset was included in our model as a single factor, 86% of the fields in high- and low-infestation areas were classified correctly. This finding highlights the importance and value of flowering time as an infestation risk predictor (Table 5, M1).
Furthermore, the study found that the arrival of the first male moth in the currently growing pea fields turned out to be a further indicator for pest-host synchronization. The extent of pea-moth attack decreased exponentially with a later arrival time (Fig. 2). Infestation risk was high only when the male moth arrived early in the season, which occurred in 2017 and 2018. Our findings suggest that fields that have attracted male moths earlier in the season are likely to accumulate more individuals over time, resulting in higher damage. Fields that started flowering significantly early or late were characterised by the late arrival of moths in monitoring traps towards the end of the season. This scenario resulted in a low percentage of damaged grains. Therefore, infestation was highest if the susceptible development stage of the plant coincided with the beginning of the emergence period. For most crops, the time of infestation in a susceptible plant growth stage and yield loss are closely related (Awuni et al. 2015; Bardner and Fletcher 1974). The results of this study confirm that the same applies to C. nigricana: A favourable match in space and time between crop and pest-population development is a prerequisite for high damage.
However, in this study, not all the fields with a high infestation rate were flowering within the GDD of mean early pea-moth emergence. The dispersion of the insect in space is likely another important factor. In contrast to the results of Thöming et al. 2011) and Huusela-Veistola and Jauhiainen 2006), CAI was not significantly correlated with damage in this study. Therefore, CAI was not included in the study’s final model. Minimum distance significantly affected the infestation rate only in the high infestation years, 2017 and 2018 (Fig. 3). In those two years, a negative exponential decrease in infestation rate with increasing distance to previous year’s pea fields was observed. This finding is in accordance with that of Huusela-Veistola and Jauhiainen 2006). No correlation was found in low infestation years, which explains the poor fit overall.
Combining MD and flower time in one model increased the prediction quality significantly. Pea fields closely located to the overwintering sites, and thus characterized by low MDs, were substantially infested only when they were also olfactorily most attractive, which is from bud development until the end of flowering (Thöming and Knudsen 2014). This means that if a pea site is closer than another but not flowering, the directional movement of the moths would be towards the more distant but flowering pea site, resulting in a higher infestation that is independent from MD. In the case of the leaf-defoliating Colorado potato beetle, which is less dependent on a certain plant development stage for offspring survival than the pea moth, host location is less directional and less stage specific.Weisz et al.(1996) were therefore able to model the intensity of attack as a function of only the migratory distance with much greater precision than it was possible in this study. The wide range of flowering patterns in our study region and the related variable olfactorial attraction negatively affected CAI- as well as MD-only based damage predictions. Spatial risk factors might lead to better results in environments with a more synchronized crop phenology, which applies to either early vegetable pea varieties or grain pea cultivars being more synchronized at scandinavian agroclimatic conditions described in the studies ofThöming et al.(2011) and Huusela-Veistola and Jauhiainen, (2006), respectively.
Combining all three of the factors discussed above was essential and yielded the best model results. Information about flowering time, pea-moth arrival and MD makes it possible to discriminate between high- and low-infestation fields with greater precision. However, these results are preliminary and, at this stage, implementation of the model requires data monitoring of the fields, which is time intensive. Farmers do not have the required time to perform such monitoring. Thus, for the practical application of the model, the future development of digital data-acquisition processes is of major importance. In recent years, the technological means of remote sensing have expanded rapidly in agriculture. Crop classification methods using satellite data are already widely studied (Sun et al. 2018; Meng et al. 2021; Wang et al. 2022b). The automatic recognition of pea fields based on satellite imaging would allow for MD calculations without the need to involve farmers. In addition, even without including MD, the model suggested by this study correctly classified 97% of the fields. Schieler et al. (subm. unpublished) developed a temperature- and photoperiod-based model that can predict development stages, including florescence, based on sowing date and pea variety. Furthermore, in the future, the practical implementation of satellite remote sensing in predicting crop phenology can facilitate the recognition of a particular growth stage, like florescence, without requiring actual field visits (Gao and Zhang 2021).
The detection of the first male moth arrival was of great significance for the quality of the proposed model. The manual monitoring of pheromone traps would require at least weekly control of all currently growing pea sites subsequent to moth emergence in previous pea sites. A temperature-based model that predicts pea-moth emergence already exists and can help to restrict the time span needed for placing and maintaining pheromone monitoring traps in currently growing pea sites (Riemer et al. 2021). Automatic pest-counting insect traps using deep-learning techniques (Bjerge et al. 2021; Hong et al. 2021tő 2021; Wang et al. 2022a) would be a substantial improvement on these traps, but they are not market ready yet.
Limitations and recommendation for future research
This study faced certain limitations. Although weather data was interpolated from a height 2 m, reproducible results were achieved. However, further microclimatic refinements would produce greater precision. Data collected over only four years was included in the model, and the number of fields was reduced by almost half due to the exclusion of fields that received insecticidal treatment. The limited data can influence the precision of the model, which therefore requires validation by independent datasets of successive years. This study’s data is also restricted to the local conditions of one model region. Therefore, the data requires validation before generalizing it to other agroecological pea-cultivation areas. Furthermore, this preliminary model was developed with data from grain peas; the prediction needs to be refined and expanded to other grain-pea and vegetable-pea varieties.
Regarding the factor ‘first male moth trapped’, two aspects must be pointed out: First, even detecting just the male fraction of the pest population resulted in reproducible predictions. Nevertheless, predictions could be improved in the future by addressing the fact of specifically mated females being more attracted to the floral odor of flowering peas than males (Thöming and Knudsen 2014). Monitoring the females with kairomone-baited monitoring traps could improve the prediction results, in analogy to Cydia pomonella monitoring (Knight 2010; Knight and Light 2005; Light et al. 2001). when this study was conducted, such traps containing pea-plant volatiles (kairomones) were not commercially available yet. Second, Pisum sativum is the preferred and most abundant host plant within the model region in terms of acreage. However, other cultivated and non-cultivated grain legumes, such as Vicia faba, Vicia sepium, Laturus odoratus, Orobus sp. and others, have also been reported as potential host plants (Wright and Geering 1948; Hoffmann and Schmutterer 1999; Kruess and Tscharntke 2000). Although the reproductive potential in alternate hosts can be regarded as comparably low in general (Thöming and Norli 2015), the extent to which alternate host plants may affect the forecasting value of first moth arrival in monitoring traps and the factor MD remains unknown.