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Article

Processing Tomato Crop Benefits from Flowering Plants in Field Margins That Support Pollinators and Natural Enemies

by
Vaya Kati
1,*,†,
Theodoros Stathakis
2,*,
Leonidas Economou
2,
Philippos Mylonas
2,
Myrto Barda
2,
Theodoros Angelioudakis
3,
Athanasia Bratidou Parlapani
1,
Ilias Tsamis
4 and
Filitsa Karamaouna
2
1
Laboratory of Weed Science, Scientific Directorate of Pesticides’ Control and Phyto-Pharmacy, Benaki Phytopathological Institute, 8 Stefanou Delta Str., GR-145 61 Kifissia, Attica, Greece
2
Laboratory of Efficacy Control of Pesticides, Scientific Directorate of Pesticides’ Control and Phytophamacy, Benaki Phytopathological Institute, 8 Stefanou Delta Str., GR-145 61 Kifissia, Attica, Greece
3
Laboratory of Agricultural Zoology and Entomology, Faculty of Crop Science, Agricultural University of Athens, 75 Iera Odos Str., GR-118 55 Athens, Greece
4
D. Nomikos S.A., 32 Kifissias Ave., GR-151 25 Maroussi, Attica, Greece
*
Authors to whom correspondence should be addressed.
†
Current address: Laboratory of Agronomy, Faculty of Agriculture, Aristotle University of Thessaloniki, GR-541 24 Thessaloniki, Greece.
Agronomy 2025, 15(7), 1558; https://doi.org/10.3390/agronomy15071558
Submission received: 1 June 2025 / Revised: 19 June 2025 / Accepted: 24 June 2025 / Published: 26 June 2025
(This article belongs to the Special Issue Pests, Pesticides, Pollinators and Sustainable Farming)
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Abstract

In a two-year experiment, we examined whether increasing plant diversity in the margins of processing tomato fields could attract pollinators and natural enemies of pests compared to weed flora, and questioned the effect on crop yield. Two plant mixtures sown in winter (WM) and spring (SM) were compared with weed vegetation along a tomato crop (CT) and an adjacent irrigation channel (CC). Flower cover was higher in the sown mixtures than the weedy margins, and brought in more visits of pollinating bees (including potential tomato pollinators) than the latter. Flowering species were mainly Eruca vesicaria (WM, SM), Coriandrum sativum and Lathyrus sativus (WM), Fagopyron esculentum and Phacelia tanacetifolia (SM), and Ammi majus, Rapistrum rugosum (CC, CT). Parasitoids (Eulophidae, Braconidae, Scelionidae) were more abundant in the sown and CC margins compared to the CT margin, while the abundance of predators (Aeolothripidae, Orius sp., Thomisidae) was similar among all types of margins. Fruit weight was higher in the field with the sown margins, while pest incidence in the crop was not affected by the margin type. Our findings provide new insights into the contribution of managed and existing field margins in attracting beneficial arthropods, and their implications on yield.

1. Introduction

Increasing intensification of agriculture to meet the growing demand in food production could dramatically affect farmland biodiversity [1]. Moreover, additional stressors resulting from the ongoing climatic change are reported to negatively affect plant diversity and alter the onset of flowering [2], challenging the ecosystem services of pollination [3,4,5] and biological control [6,7]. Both services are valued at billions of euros worldwide [8,9]. Furthermore, pollination is directly affecting food security since it is vital for 75% of global food crops [10].
Sustainable management strategies in agricultural landscapes are emerging as the way forward to halt the functional biodiversity loss associated with agricultural intensification [11,12]. In this respect, the European Union (EU) has launched mandatory or voluntary agro-environmental schemes specifically designed to promote ecological balance and sustainability in agroecosystems as part of the new Common Agricultural Policy (CAP) program and European Green Deal [13]. Such agro-ecological measures include, among other, universally accepted field management practices to enhance the ecosystem services of pollination and biological control based on the improvement of plant diversity in degraded field margins with sown mixtures of selected flowering plants [14]. Extensive research over the past few decades has demonstrated the positive relationship between plant diversity along field margins, and beneficial arthropod abundance and species richness in various crops [15,16,17,18,19,20,21,22,23,24]. However, a meta-analysis on the impact of field margins on biological pest control identified knowledge gaps regarding their effect on crop yield [25]. Indeed, an earlier quantitative synthesis of data from 35 studies by Albrecht et al. [26] indicated that flower strips along field margins had variable effectiveness on crop yield. What is more evident is a clear advantage regarding crop productivity on smaller farms (<2 ha) gained through higher flower visitor density, while larger fields required higher flower visitor richness for a measurable benefit [27]. It is thus obvious that plant selection and synthesis in mixtures are crucial for the desired impact, depending on crop species, field size and landscape complexity, as some species may also support herbivorous pests, potentially harming the crops.
Field grown tomatoes (Solanum lycopersicum L.) are primarily self-pollinating but can benefit from insect-mediated pollination [10]. Although our knowledge on insect pollinators of tomato remains limited, species like Bombus and Anthophora are known to improve tomato fruit set, especially under high temperatures that reduce pollen viability [28]. Bombus pascuorum (Scopoli) and B. terrestris (L.), Megachile willughbiella (Kirby), Hylaeus gibbus Saunders, and buzzing Lasioglossum species are reported to benefit the pollination of tomato in Central Europe due to their ‘buzzing’ vibration effect on the flowers [29]. A recent study in Greece showed that selected native non-crop flowering plants sown along the margins of a processing tomato field could increase pollinator and beneficial arthropod abundance compared to locally occurring weeds, but did not elucidate the effect on crop pollination and yield [19]. Following on that study and considering the scarcity of seeds from non-crop flowering plants in Greece, here, we studied the effect of field margin management with mixtures of selected flowering cultivated plant species that have available seeds in the Greek market. Our specific objectives were to evaluate the effectiveness of these plant species sown in mixtures along the field margins of processing tomato crops in attracting pollinators and natural enemies compared to weed flora, and whether such species could be a suitable alternative to non-crop flowering plants. Secondly, in addition to the previous study [19], here, we aimed to quantitatively assess the effect of this field margin management practice on crop yield. The study aligns with the EU’s sustainable agriculture policies, aiming to protect and enhance biodiversity within agroecosystems and is conducted as part of the Operation Pollinator biodiversity project in Greece, which has been implemented in various crops since 2010 [18].

2. Materials and Methods

2.1. Selection of Plant Species and Mixture Composition

Two flowering plant mixtures were composed for sowing in winter (WM) and spring (SM) (Table 1). The species selection and their proportion in the mixtures was based on previous experience obtained during implementation in processing tomato and other crops in various areas of Greece [18,19,20,30]. Briefly, the general criteria included the selection of annual species known to exist in the area, which do not pose a threat as potential noxious weeds, or act as specialized hosts for tomato pests, are attractive to pollinators and natural enemies and belong to a diverse range of families. Seed availability defined the final species selection and mixture composition.

2.2. Seed Rate Calculation

The calculation of the seed rate per species was based on the following parameters: target plant number/m2, percentage of each species in the mixture, thousand grain weight per species, seed germination capacity based on growth chamber germination assays, and estimated plant survival rate in the field. Based on the above, the seed weight in grams for each species in a mixture (Ws) is the output of the equation
Ws = Ps × (1/ESR) × (1/Pg) × Tp × A × (TGW/1000)
where Ps = % of each species (Table 1), Tp = total target number of plants/m2 (set to be 80 plants/m2), A = area to be sown per field (84 m2/per plant mixture), TGW = thousand grain weight/species (g), ESR = estimated survival rate of germinated seeds (set to be 65% for all dicots and 80% for wheat), and Pg = seed germination percentage based on petri-dish assay results per species.

2.3. Experimentation Site

Two-year field experiments (2021–2022) were established in the main processing tomato area in Larissa, (Thessaly, Greece) at a different location each year (Figure 1A). The two plant mixtures (WM and SM) were sown along one of the field margins, as shown in Figure 1B. The plot size was 14 m2 (7 m long × 2 m wide), with six plots per plant mixture (total sown area per mixture: 84 m2). Two sets of control plots consisting of weed flora were assigned along the margin next to the crop (CT) or along an adjacent irrigation channel (CC), to cater for the different arable weed communities present around the field (Figure 1B). Each control site had six plots (total area per control site: 84 m2).
Field preparation of the sown margins included shallow soil cultivation to control weeds, and hand raking to smooth the surface. Sowing was performed by hand. A bulking material (corn meal at 200 cm3/m2) was used to augment the seed quantity and enable its even distribution on soil surface. After sowing, the seeds were covered by hand raking, and the soil surface was rolled to ensure good seed/soil contact, using a manually operated seed roller. Sowing of the WM was performed in early February of each year while the SM was sown in early April. The H1015 Heinz jointless processing tomato hybrid (2nd early, 105–114 days to maturity) was transplanted mid-March of each year.

2.4. Arthropod Measurements

Measurements of flowering and attracted beneficial arthropods were performed during the main flowering period of the sown plants, from mid-May to late-June in 2021, and from early-June to mid-July in 2022.
The total plant cover and flower cover (total and per species) were visually estimated and expressed as percentage cover/plot in all plots of the sown mixtures (WM, SM) and the two control sites (CC, CT). Plant species were identified in situ or, when necessary, in the lab using the botanical identification key Flora Europaea [31].
Hymenoptera pollinator (Apis mellifera L., wild bees) visits on the flowers of the sown margins and the control plots were recorded with visual observation of landings for 4′/plot between 10:30 and 14:30 h. The observations and corresponding counts refer to the foraging visits of pollinators and not their absolute numbers which could not be accurate due to their high abundance and their mobility from flower to flower during observation time. All the observations throughout the sampling period were made by the same observer to eliminate potential bias between different observers. Measurements were conducted in three of the six plots (1st, 3rd, and 5th) of each treatment (WM and SM), as well as for the control sites (CT and CC).
Pollinator visits were also recorded on the crop flowers with visual observation as before, at three sites based on the distance from the sown field margin, starting from the first two twin crop rows next to the margin and moving infield to the 20th–21st and 40th–41st twin rows. Each sampling site had three replications of approximately 14 m2 (2 twin rows, 7 m long).
Wild bee specimens that required identification after the visual observation measurements were captured with a sweeping net and were identified later in the lab. The identification of pollinators was based on identification keys [32,33].
Beneficial arthropods (parasitoids, predators) were recorded with suction sampling (16″/plot) using a modified leaf blower (Echo ES-2400, 24 cm3, Kioritz Corporation, Tokyo, Japan), adapted as described in [34]. Measurements were conducted in the other three plots (2nd, 4th, and 6th) of the sown margins and the control sites, and in the three corresponding plots of the first two twin crop rows.
Additionally, the herbivore insects collected via suction sampling were counted. Only individuals belonging to groups that could be potentially harmful for tomato crop were taken into consideration, i.e., aphids (Aphidoidea), leafhoppers (Auchenorrhyncha), whiteflies (Aleyrodidae), stink bugs (Pentatomidae), and thrips (Thysanoptera).
The collected arthropod samples were kept in the freezer (−18 °C) and sorted according to family, genus, and species (where possible) under a stereomicroscope.

2.5. Crop Yield Parameters

The effect of the sown flower margin intervention on the quality characteristics of tomato fruit was assessed based on the following parameters: fruit weight (digital laboratory scale), BRIX [digital refractometer Maselli UR 24 (Maselli Misure S.P.A., Parma, Italy)], pH (pH-meter Metrohm 713), and color, including brightness (L*), redness or greenness (a*), yellowness or blueness (b*) and the ratio a/b [colorimeter Hunterlab DP-9000 D25A (Hunter associates laboratory, Reston, VA, USA)]. All analyses were performed based on in-house protocols at the premises of D. Nomikos S.A., Agricultural Department, Panagia, 35010, Domokos, Greece. Samples of 20 fruits/plot were collected from the fields with the sown margins and the control field with the weed flora, and were transported immediately for analysis. The samples were collected from twin crop rows aligned with the 1st, 3rd, and 5th sown or weed plots, at three field sites (i.e., 1st–2nd next to the margin, the 20th–21st row and 40th–41st row).

2.6. Statistical Analysis

Generalized linear mixed models (GLMM) were conducted to examine the effect of field margin management (treatment), plant cover, flower cover, and number of plant species in bloom, as fixed factors, on the abundance of the main arthropod functional groups (honeybees, wild bees, parasitoids, predators and insect pests) in the field margins. Experimental year and sampling date were treated as random factors. The models used a negative binomial distribution with a log link function. Fisher’s LSD method was used for pairwise comparisons. Kruskal–Wallis ANOVA was used for comparison of plant and flower cover among field margin treatments, and for examination of the effect of field margin management (treatment) on the abundance of parasitoids, predators, and insect pests in the processing tomato crop. A one-way ANOVA (α = 0.05) was performed to examine the effect of field margin management (treatment) on crop yield parameters. The means were separated using Tukey’s HSD test. Statistical analysis was carried out using SPSS version 21.0 for Windows software (IBM Corp; Armonk, NY, USA).

3. Results

3.1. Plant Cover

Overall, the WM margin provided complete plant cover that differed significantly compared to all other treatments (χ2 = 88.565, df = 3, p < 0.001) (Figure 2). In general, all plant species in the mixtures sown in winter (WM) or spring (SM) along the margin of the selected processing tomato field, emerged and provided a dense plant cover both years (98–100% of the plot area). Exceptions were P. tanacetifolia in 2021 and F. esculentum in 2022 in the SM, with very low corresponding densities. This was attributed to the low germination ability of P. tanacetifolia in 2021 and to a late frost that reduced the survival of F. esculentum in 2022. However, the remaining sown species resulted in a high plant cover, which was mainly due to E. vesicaria and secondly to L. sativus.
The two control sites with weed flora, either next to the irrigation channel (CC) or next to the crop (CT), had comparable plant cover in 2021. However, in 2022 from mid- and late June because of an accidental glyphosate application for weed control on that site by field workers. The plant cover in that margin was partially recovered only around mid-July (Supplementary Figure S1).

3.2. Flower Cover

The flower cover was the highest for the sown mixtures (WM, SM), while from the control margins the CC flower cover was comparable to the SM margin and the CT was significantly lower from all other treatments (χ2 = 56.088, df = 3, p < 0.001) (Figure 2). In 2021, the sown mixtures combined provided continuous floral resources from mid-May to mid-June. The mean flower cover of the WM mixture was higher (68–100% of the plot area) compared to the other sites, until late-May. In early-June, flower cover was similar in the sown mixtures (WM and SM) and the CC weed margin site, which maintained an intermediate flower cover (28–61% of plot area) throughout the experiment. In mid-June the highest flower cover was provided by the SM (92% of the plot area). The CT weed margin had the lowest flower cover throughout the season (>10% of the plot area). In 2022, the combined flowering of the two sown mixtures lasted from June to mid-July. Early in the season (early to mid-June) the flower cover of the sown mixtures was comparable to that of the weed flora (18–32%). The flower cover of the sown mixtures was higher compared to the weed flora later in the season, with a peak in late-June (63% and 33% of the plot area for the sown mixtures and weed flora, respectively) (Supplementary Figure S2).
The main flowering species in each sown mixture varied over time during both years (Figure 3). In 2021, the order of flowering species in the WM was E. vesicaria, L. sativus, P. sativum, C. sativum, A. graveolens, and in the SM, it was L. sativus, F. esculentum, P. tanacetifolia, and E. vesicaria. The main dicotyledonous weed species that emerged and reached flowering together with the sown species was Rapistrum rugosum (L.) All. in the WM and Heliotropium europaeum L. in the SM. Other weeds that emerged included Solanum nigrum L., Chenopodium album L., and Sorghum halepense (L.) Pers. Overall, weeds emerged in low numbers and did not affect the establishment and flowering capacity of the sown mixtures. That year, the CC margin was dominated by the Brassicaceae R. rugosum mid-May, and later by Apiaceae species. In the CT margin, flower cover was provided by a few R. rugosum and Sisymbrium sp. plants (Figure 3 and Supplementary Figure S3).
In 2022, E. vesicaria was more dominant in the SM due to the low emergence of P. tanacetifolia and the absence of Fagopyrum esculentum that was lost after the late frost. The order of flowering among the sown species in the WM was the same as the previous year: E. vesicaria > L. sativus > P. sativum > C. sativum > A. graveolens, while in the SM, the order was L. sativus ≥ P. tanacetifolia ≥ E. vesicaria. Several flowering weeds emerged in low numbers in the sown mixtures and had a very low contribution to the flower cover. These were, in WM: Convolvulus arvensis L., Ecballium elaterium (L.) A.Rich., S. nigrum, Fumaria officinalis L., Portulaca oleracea L., Sinapis arvensis L., R. rugosum, and in SM: C. arvensis, E. elaterium, S. nigrum, S. arvensis. As mentioned in the plant cover section, in 2022, flower cover in the CT control site was recorded only in early June and mid-July, with S. arvensis and S. alba L. being the main flowering species. The flower cover in the CC site was almost solely provided by Ammi majus L. throughout the season (Figure 3 and Supplementary Figures S4 and S5).

3.3. Hymenoptera Pollinators

The total abundance of honey bees was significantly higher in the sown margins (WM and SM) compared to the weed flora along the irrigation channel (CC) and the field margin of the tomato crop (CT) (F3,91 = 4.190, p = 0.008). The flower cover percentage had a significant effect on honey bee abundance (F1,91 = 17.709, p = 0.000), while the number of plant species in bloom did not have any significant effect (F1,91 = 1.817, p = 0.181). On the contrary, the abundance of wild bees did not differ statistically among treatments (F3,91 = 2.106, p = 0.105) (Figure 4 and Supplementary Table S1). On a date-to-date examination, it is obvious that honey bees had the highest abundance in late June (27 June 2022) in SM, when the available floral resources of E. vesicaria reached the highest flower cover percentage (Supplementary Figure S6). Sporadic visits of wild bees (1–2 individuals/plot/4 min of observation time) were recorded on tomato flowers regardless of the row distance from the field margins throughout the experiment. These wild bees belonged to the genera Andrena, Lasioglossum and Nomiapis. The wild bee genera recorded on flowers of the sown mixtures, weed flora in the control sites, and tomato flowers are shown in Table 2 (Supplementary Figures S7 and S8).

3.4. Natural Enemies

The total abundance of parasitoid wasps was significantly higher in the sown margins (WM and SM) and weed flora along the irrigation channel (CC) compared to the weedy margin next to the tomato crop (CT) (F3,89 = 4.214, p = 0.008). The plant cover, flower cover percentage and the number of plant species in bloom did not have a significant effect on parasitoid abundance (F1,89 = 0.942, p = 0.334, F1,89 = 0.050, p = 0.823 and F1,89 = 0.073, p = 0.787, respectively). The abundance of predatory arthropods did not differ statistically among treatments (F3,89 = 0.930, p = 0.430); however, the SM margin harbored higher numbers of predators. The flower cover percentage and number of plant species in bloom had a positive effect on the abundance of predatory arthropods (F1,89 = 4.323, p = 0.040 and F1,89 = 5.224, p = 0.025) (Figure 5A and Supplementary Table S2).
The total abundance of natural enemies in the tomato crop followed a similar pattern to that of the field margins. Higher numbers of parasitoid Hymenoptera were observed in the tomato fields with sown margins compared to the control fields (χ2 = 15.566, df = 2, p < 0.001), while the abundance of predators did not differ significantly among treated tomato fields (χ2 = 0.902, df = 2, p = 0.637) (Figure 5B).
Looking at their temporal abundance fluctuation of parasitoid Hymenoptera, the CC control margin was superior to the WM sown margin at the beginning of May 2021 (54 ± 5.3 and 39 ± 17.1, respectively). The WM sown margin had higher numbers of parasitoid wasps compared to both control margins during the rest of the sampling period of both experimental years. The SM margin reached peak levels during early June 2021 (36.7 ± 11) and mid-June 2022 (54 ± 10.4). The highest numbers of predators were observed in SM sown margin from the beginning until mid-June on both sampling periods (Supplementary Figure S9).
In the tomato crop, parasitoids in the fields with sown margins had increased numbers from mid to late May, and mid-June during the first sampling period (2021), while predators had generally low populations. In 2022, predatory arthropods were more abundant reaching peak levels during mid-July in the tomato fields with sown margins (10.7 ± 0.3 for WM and 10 ± 1.2 for SM). At the same time, the highest numbers of parasitoid wasps were observed (8.7 ± 1.3 for WM and 14.3 ± 2.8 for SM) (Supplementary Figure S10).
Parasitic wasps in the samples belonged to 7 different superfamilies (Ceraphronoidea, Chalcidoidea, Chrysidoidea, Cynipoidea, Diaprioidea, Ichneumonoidea, Platygastroidea) and 24 different families. The samples from the WM belonged to 20 families, while the ones from the SM mixture belonged to 19 families and from the CT and CC sites to 20 and 19 families, respectively. The WM and SM sown margins had similar community structure dominated by Eulophidae (35–36%), Braconidae (17–19%), and Scelionidae (19%), while Encyrtidae held 6.5–8%. The weed flora along the irrigation channel (CC) attracted mostly eulophids (44.7%), while braconids, scelionids and encyrtids held subequal proportions (11–12%). The weed margin next to the crop (CT) harbored mostly Eulophidae (37%), Mymaridae (14.6%) and Braconidae (12.6%). Within the tomato crop, the parasitoid families with the highest proportion were Braconidae (35%) for fields with WM margin, Scelionidae (23%) for fields with SM margin and Mymaridae (19.7%) for the control fields (Figure 6A).
The communities of predatory arthropods hosted by the WM and SM sown margins were also similar; they consisted mostly of predatory thrips (Aeolothripidae) (48.5–54.7%), pirate bags (Anthocoridae) (15–21.5%), and crab spiders (Thomisidae) (8–11%). The proportions of these groups in CC sites were 36.5%, 13.5%, and 24.5%, respectively. Predatory thrips were superdominant (71%) in CT margins (Figure 6B). In tomato fields with the sown margins, several predatory groups were detected such as predatory Heteroptera (Anthocoridae, Miridae, Nabidae), spiders (Linyphiidae, Oxyopidae, Thomisidae), and predatory thrips, without any obvious dominance. The control fields hosted mostly crab spiders (25.7%), predatory mirids (20%), and Aeolothripidae (17.6%) (Figure 6C,D).

3.5. Insect Pests

Regarding the abundance of insect pests, the sown margins and the weedy margin next to the crop harbored significantly more insect pests than the margin along the irrigation channel (F3,89 = 3.950, p = 0.011), while there were no statistical differences between the tomato fields (χ2 = 5.129, df = 2, p = 0.077) (Figure 7).
In the SM sown margins, the most abundant group was aphids (44.6%), in the WM margins and the CC sites, thrips (49% and 51.8%, respectively), and in the CT sites, leafhoppers (51%). Within tomato crops, the control fields (C) and those with the SM margin hosted mainly aphids (51% and 52%, respectively), while the fields with WM margin hosted mostly thrips (33.8%) (Supplementary Figure S11).

3.6. Crop Yield

The yield parameters (means) of processing tomato (weight, BRIX, pH, L, color AB) are shown in Table 3. In both experimental years, the average fruit weight from fields with sown margins was similar to the expected values (75–80 g/fruit) and significantly higher compared to the control fields with weed vegetation (F2,24 = 93.179, p < 0.0001 for 2021, F2,24 = 54.136, p < 0.0001 for 2022). The total soluble solids (TSS) of fruits harvested from WM, SM and control fields were similar to the expected value (5.2 °Bx) in 2021; however, in 2022, the fruits from the control field had significantly higher levels of TSS (6.1 °Bx) compared to those with the sown margins (F2,24 = 21.116, p < 0.0001). The pH values ranged between 4.4 and 4.5 without significant differences. Regarding the fruit color parameters among treatments, a significantly higher L value was observed in the control field in 2021 (F2,24 = 10.377, p = 0.011) and significantly lower a/b value in 2022 (F2,24 = 8.714, p = 0.001) (Table 3 and Supplementary Table S3).

4. Discussion

The establishment of flower strips inside the crop or at the field margins is among the agro-ecological practices outlined in the EU’s recent common agricultural policy [35] to maintain functional biodiversity in agroecosystems [36]. Indigenous wild flora species are generally preferred for the flower strips, to avoid genetic erosion from imported species and to exploit their adaptation to local conditions, ensuring a successful establishment (19). However, as indicated by our results, a feasible and effective alternative for the implementation of this agro-ecological practice could be the use of cultivated flowering plants, such as the Apiaceae (C. sativum and A. graveolens), Brassicaceae (E. vesicaria) and Fabaceae (L. sativus, P. sativum) species tested here, with easily available seeds in the market. Nevertheless, field margins with weed flora adjacent to the experimental tomato crop were also assessed to examine the ability of natural flora to safeguard beneficial arthropods for agriculture in comparison to the sown flowering mixtures, as it has been conducted in previous studies in tomato and other crops [18,19,37]. Assessing field margins for the dual purpose of supporting both pollinators and natural enemies is in line with the Integrated Pest and Pollinator Management (IPPM) framework proposed recently by Egan et al. [38] to enhance IPM compatibility with crop pollination management and enable their coordination towards a unified economic decision making and implementation. However, diversified plant resources provided by crop or wild species should be characterized on their capacity to support functionally important arthropods, such as pollinators and natural enemies, while also considering potential trade-offs such as enhancing crop pests or causing weed related problems [18,39] before employing them at the field scale as an IPPM tool [40].
Our two-year field margin assessment indicated that the sown flowering mixtures attracted more honey bee visits compared to the weed flora, as also reported by other studies [20,41]. Furthermore, the flower cover percentage was positively correlated with the number of attracted bee visits, but not the number of flowering plant species, as also recently reported by Liira and Jürjendal [42]. The latter study provided evidence that areas with low flower diversity are likely to attract higher number of bees, and that agricultural policies for the conservation of pollinators should not focus on floral biodiversity, as it is commonly supported [43,44]. However, there is undisputable evidence that diversity in floral traits has a positive effect on bee species diversity because it caters for their variable nectar and pollen needs, while also ensuring that accessibility requirements are met, which depend on flower and bee anatomy [45].
Most of the bees recorded on the margins of the field were honey bees (A. mellifera). However, honey bees are unable to perform buzz pollination, i.e., to vibrate the tomato flower in the frequency needed to release the pollen from the anthers to the stigma [46]. Nevertheless, wild bee genera (e.g., Amegilla) recorded in lower numbers of visits in our study, include species which use floral buzzing while foraging [47], and are thus considered potential pollinators of tomato flowers [48]. In fact, in our study, only wild bees were recorded to visit tomato flowers (Andrena, Lasioglossum and Nomiapis), although in small numbers. However, upscaling the practice of flowering field margins in tomato crops could possibly enhance foraging on tomato flowers by these pollinators and subsequently contribute to the improvement of the fruit set percentage and fruit characteristics [49]. Similar wild bee genera were attracted on the sown flowering strip of similar experimentation in tomato (the WM included also Glebionis coronaria (L.) Cass. ex Spach), in the southern part of continental Greece (area of Orchomenos), i.e., mainly included Andrena spp., Colletes sp., Halictus sp., Hylaeus spp., Lasioglossum spp., Pseudapis sp. (identified later as Nomiapis sp.; Barda, personal communication), and Sphecodes sp. [19], contributing to tomato pollination with Hylaeus spp. and Lasioglossum spp. [29]. Further experimentation should focus on the long-term effects of the sown flower mixtures to insect pollinators.
Eruca vesicaria, the species which held the higher percentage of the flower cover and longest flowering duration in the mixtures, attracted honey bees and wild bees (Amegilla, Andrena, Ceratina, Eucera, Halictus, Lasioglossum). Indeed, Eruca vesicaria has been proven to attract a wide variety of pollinators [50,51,52]. Coriandrum sativum and A. graveolens also provided a long-lasting flowering throughout the season, which overlapped with that of the tomato crop and lasted until the fruit set, providing an attractive habitat for potential pollinators of the crop. Coriandrum sativum attracted both honey bees and wild bees, including genera recorded on tomato flowers (Andrena, Nomiapis), confirming its favorable profile as a tool to promote the ecosystem services of pollination as well as biological pest control [19,53,54,55]. Records of wild bee attraction by C. sativum include the families Andrenidae, Halictidae and Colletidae (Algeria) [53], and Megachile spp. (Pakistan) [56]. Likewise, the second Apiaceae, A. graveolens, brought in honey bees along with wild bee genera (Andrena, Hylaeus) known to pollinate tomato [29]. Lathyrus sativus attracted early in the season bees of Eucera, as also stated at the similar study by Kati et al. [19], but also of Megachile, which are among the most efficient pollinators for tomato [29], and Osmia.
In the SM mixture, the flowering of F. esculentum, evident only in the first year, attracted both honeybees and wild bees of the genera Andrena and Lasioglossum. In other studies, reported foraging of wild bees on F. esculentum flowers include Bombus spp. [57,58], Xylocopa spp., Halictus spp. in Greece [19], as well as other Halictidae species in Florida [51]. The capacity of F. esculentum to upgrade the floral resources of field margins for the benefit of pollinators and beneficial arthropods is supported by [59,60]. The poor establishment of P. tanacetifolia in the SM mixture did not allow for the full evaluation of its potential for use in flowering mixtures in the study area. Its low flower cover was foraged by both honey bees and wild bees as also reported elsewhere [18,61]. However, Petanidou [61] expressed concern on the suitability of this species for use as a flower resource for pollinating bees in Greece.
Furthermore, the contribution of wild plants already present in the field margins to attracting pollinators should not be overlooked, as long as they are not among species that are likely to infest the crop as weeds. For example, A. majus is a wild plant that is not threatening as a weed in the area of our study. This species is known to be a nectar-bearing plant [62] and has been reported to attract species of Andrena [63]. In our study, it supported wild bee genera (Andrena, Hylaeus, Lasioglossum, Nomiapis), some of which were also recorded on tomato flowers and include species that can buzz-pollinate [29].
Both sown flowering mixtures and the undisturbed natural vegetation along the irrigation channel outcompeted the weed margin along the tomato field (CT) as reservoirs of parasitoid wasps. The positive effect of flower-rich strips on the abundance of parasitoid Hymenoptera has been demonstrated by several studies [19,60,64,65,66]. However, the fact that the total flower cover of the plot area and the number of plant species in bloom did not affect the population density of parasitoids, indicate a complex interplay among different variables that can affect parasitoids’ abundance. Each species has indeed a different potential as an insectary plant regarding the abundance and diversity of parasitoids [67,68], while on a given plant individual, the parasitoids’ abundance may be affected by host abundance and the plant’s flowering state [69].
The dominant parasitoid family in all field margins was Eulophidae. This family includes parasitoid wasps against tomato crop pests, such as leafminers Liriomyza spp. (Diptera: Agromyzidae) and the tomato leafminer Tuta absoluta Meyrick (Lepidoptera: Gelechiidae) [70,71]. Braconidae and Scelionidae were also abundant in the field margins. Braconids are known for their contribution to controlling aphid populations and several species are promising biological control agents of T. absoluta and the cotton bollworm Helicoverpa armigera Hübner (Lepidopetra: Noctuidae) [72,73,74]. Scelionids are mainly egg parasitoids, with a wide range of hosts, which have been proven effective in the biological control of stink bugs (Hemiptera: Pentatomidae) such as Nezara viridula (Linnaeus) and Halyomorpha halys (Stål) [75,76].
The benefits from the sown flowering plants to parasitoids have been demonstrated by several studies. The main flowering species of both sown mixtures, E. vesicaria, has been reported to host a high diversity of potential natural enemies (parasitoids and predators) of aphids and thrips, mainly due to its early and long-lasting blooming [68]. Flowers of F. esculentum (component of SM) positively affected the longevity of three T. absoluta parasitoids, Necremnus artynes Walker, N. tutae (Ribes & Bernardo) (Eulophidae) and Bracon nigricans Szépligeti (Braconidae) [77,78]. Nectar of A. graveolens, C. sativum (components of WM) and F. esculentum improved the survival of the stink bug parasitoid Trissolcus japonicus (Ashmead) (Scelionidae) [79].
A higher abundance of parasitoid Hymenoptera was observed also within the tomato fields with sown margins compared to the control fields. Samples included mostly braconid wasps (mainly Aphidiinae) associated with aphids’ abundance; scelionids and eulophids were also present. Fairyflies (Mymaridae) had increased proportions, probably due to the presence of leafhoppers, as they are common egg-parasitoids of Auchenorrhyncha (Hemiptera) [80].
The SM sown margin surpassed numerically the other field margins in predatory arthropods, with the most abundant groups in all cases being Aeolothripidae, Anthocoridae and Thomisidae. All the Aeolothripidae individuals were assigned to the species Aeolothrips intermedius Bagnallis which is considered the primary native predator of Thrips tabaci and other species of Thysanoptera in Europe [81] and has been reported as a very effective predator in the biological control of thrips [82,83]. Minute pirate bugs (Anthocoridae), especially Orius spp., are generalist predators that have proved successful predators of the western flower thrips, Frankliniella occidentalis (Pergande) and other thrips [84], but they can also prey on other key agricultural pests, such as whiteflies, aphids and spider mites [85,86,87]. Both Aeolothripidae and Anthocoridae are omnivorous predators that also feed on different plant resources such as pollen and nectar [88,89] and the increasing plant diversity allows them to optimize their fitness by exploiting various plant-based resources such as nutrition and oviposition sites [81,90,91,92]. Of the two Thomisidae spider species, Thomisus onustus Walckenaer and Runcinia grammica (C. L. Koch) found in the samples, R. grammica, which was most abundant, is a polyphagous predator feeding on a wide range of arthropods (Diptera, Hymenoptera, Lepidoptera) [93]. Crab spiders (Thomisidae) are ambush predators exploiting flowers’ ability to attract insect visitors by sitting on inflorescences and then attacking and consuming insect prey [94,95]. These ecological strategies (omnivory and ambush hunting) explain the positive effect of flower cover and the number of plant species in bloom on the abundance of predators in our study.
Zoophytophagous mirids, Deraeocoris ruber (L.) and Macrolophus pygmaeus (Rambur) were present in the field margins and the tomato crop. Macrolophus pygmaeus is a key predator for biological control of T. absoluta, whiteflies, aphids and other pests on tomato crop [96]. Coccinellid species (Coccinella septempunctata L., Hippodamia variegata Goeze, Propylea quatuordecimpunetata L., Scymnus auritus (Thunberg) and S. suturalis Thunberg) were sampled only from the field margins and they are mainly aphidophagous and coccidophagous [97,98]. Apart from Thomisidae, several spider species of other families were collected both from the field margins and the tomato crop, i.e., Argiope bruennichi (Scopoli), Hypsosinga pygmaea (Sundevall), H. sanguinea (C. L. Koch), Mangora acalypha (Walckenaer), Neoscona adianta (Walckenaer) (Araneidae), Aphantaulax cincta (L. Koch) (Gnaphosidae), Agyneta pseudorurestris Wunderlich, Microlinyphia pusilla (Sundevall) (Linyphiidae), Oxyopes heterophthalmus (Latreille), O. lineatus Latreille (Oxyopidae), Thanatus atratus Simon, Tibellus oblongus (Walckenaer) (Philodromidae), Phylloneta impressa (L. Koch), Theridion cinereum Thorell (Theridiidae), and Trachelas minor O. Pickard-Cambridge (Trachelidae). The web-building families Araneidae, Linyphiidae, and Theridiidae are widely abundant in agricultural landscapes and trap mostly dipteran and homopteran pests on their webs, while the active hunters, Oxyopidae and Philodromidae, are wandering spiders that prey on a wide variety of insect pests [99].
Overall, the good establishment of E. vesicaria and C. sativum in combination with their ability to attract pollinators and other beneficial insects makes them superior candidate plants for agro-ecological practices that utilize annual plants for the conservation of functional groups in agroecosystems in the studied region. Potential tomato pests such as aphids, thrips and leafhoppers were detected in suction samples of all field margins, which were less abundant in the control site near the irrigation channel (CC), probably due to the composition of the flower species. Although no direct monitoring of tomato pests was conducted on the crop, the total pest abundance in the suction samples was comparable across the field margins. Moreover, no outbreak or damage by these or other tomato pests was reported by the farmers in both cultivation periods, supporting on one hand, the lack of an adverse effect of the sown mixtures on the crop and, on the other, the potential of this plant synthesis to function as a banker plant mixture building up reservoirs of natural enemies throughout the crop season [30,100].
Our results provide evidence that attracting pollinators in tomato fields by increasing the floral resources along the crop margin can have a positive impact on fruit weight. Similar findings were reported in previous research which demonstrated that insect pollination, particularly by native and buzz-pollinating bees, leads to increased fruit set, larger fruit size, and improved physicochemical properties compared to self-pollination or wind pollination in open-field tomato crops [46,101,102]. For example, Bashir et al. [102] reported that open-pollinated tomatoes, which received visits from pollinators and the simultaneous effect of wind, had significantly larger fruit sizes and higher seed weights compared to those subjected only to wind or self-pollination. More specifically, open-pollinated fruits had a mean weight (g/fruit) of 109.76, outperforming wind-pollinated (72.97) and self-pollinated (49.03) fruits. On the other hand, studies associating BRIX levels with various pollination methods in tomato are scarce. The existing evidence suggests that insect pollination can enhance tomato fruit quality, as indicated by higher BRIX values that imply changes in sugar composition, and therefore fruit sweetness [103]. In our study, BRIX levels were similar for the fruits harvested from all fields in 2021, while fruits from the control field had higher values in 2022. Fruits in that field had also the smallest size (57.3 g/fruit) compared to those from the field with the sown margins (76.2–79.3 g/fruit, for SM and WM respectively). Previous studies highlighted the complexity between fruit size and BRIX relationship, and reported that there is often an inverse correlation, particularly in small-fruited tomatoes [104]. Fruit quality parameters associated with the fruit color brightness (L*) and the hue index (a*/b*) were not consistent over the years, but ranged between the expected values for the tomato hybrid of our study. Ultimately, fruit quality is likely to be affected by the interaction of genetic, environmental, and agronomic factors, which requires further investigation.

5. Conclusions

Our results suggest that selected flowering plants can replace undesirable weeds commonly found in disturbed field margins, to serve as habitats for pollinating insects and beneficial arthropods with a potential benefit for functional biodiversity and the tomato fruit quality attributes. Introducing selected plants from the families of Brassicaceae, Fabaceae, and Apiaceae in field margins can benefit the processing tomato by attracting wild bee species (e.g., from the genera Amegilla, Andrena, Hylaeus, Lasioglossum, Nomiapis) that can support the pollination of tomato flowers, and parasitoids (Eulophidae, Braconidae, Scelionidae) and predatory arthropods (Aeolothripidae, Anthocoridae, Thomisidae) that could contribute to the biological control of tomato pests. Future research should elaborate further on the possible effect of insect pollination on tomato crop yield and quantify the potential of the sown margins to support biological control of tomato pests.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/agronomy15071558/s1, Figure S1: Mean plant cover (% of plot area) of the processing tomato field margin sown with plant mixtures and the weed vegetation; Figure S2: Mean flower cover (% of plot area) of the processing tomato field margin sown with plant mixtures and the weed vegetation; Figure S3: Flowering margins in the studied processing tomato fields for different sampling dates in late spring (May–June) 2021; Figure S4: Flowering margins in the studied processing tomato fields for different sampling dates in early summer (June–July) 2022; Figure S5: Close-up images of the flowering margins in the studied processing tomato fields on different sampling dates in early summer (June–July) 2022; Figure S6: Temporal distribution of Hymenoptera pollinators in the processing tomato field margin sown with plant mixtures, and the weed vegetation; Figure S7: Pollinators of tomato flower and various flowering species, either sown in winter (WM) or spring (SM) along the margin of a processing tomato field, or present in the weed vegetation of the margin in the control tomato field (CT) and in the weed vegetation along a nearby irrigation channel (CC) during May–June 2021; Figure S8: Pollinators of tomato flower and various flowering species, either sown in winter (WM) or spring (SM) along the margin of a processing tomato field, or present in the weed vegetation of the margin in the control tomato field (CT) and in the weed vegetation along a nearby irrigation channel (CC) during June-mid July 2022; Figure S9: Temporal distribution of natural enemies in the processing tomato field margin sown with plant mixtures, and the weed vegetation; Figure S10: Temporal distribution of natural enemies in the processing tomato crop with different field margin managements; Figure S11: Community composition of insect pests (A) in field margins; (B) in tomato crop with different field margin management. Table S1: Summary of the statistics for GLMM testing the effect of field margin management (treatment), flower cover, and number of plant species in bloom, on the abundance of Hymenoptera pollinators (honeybees, wild bees) in field margins; Table S2: Summary of the statistics for GLMM testing the effect of field margin management (treatment), plant cover, flower cover, and number of plant species in bloom, on the abundance of natural enemies (parasitoids, predators) and insect pests in field margins; Table S3: Summary of the statistics for a one-way ANOVA testing the effect of field margin management (treatment) on the crop yield parameters of processing tomato fruits.

Author Contributions

Conceptualization, V.K. and F.K.; methodology, V.K., F.K., L.E., P.M., T.S., T.A. and I.T.; formal analysis, T.S. and L.E.; investigation, V.K., F.K., L.E., P.M., M.B., T.S., I.T. and A.B.P.; resources, I.T., V.K. and F.K.; data curation, L.E., T.S., T.A. and P.M.; writing—original draft preparation, V.K., T.S. and F.K.; writing—review and editing, T.S., F.K. and V.K.; visualization, V.K., F.K., T.S., T.A. and M.B.; supervision, V.K. and F.K.; project administration, M.B., A.B.P., V.K. and F.K.; funding acquisition, F.K. and V.K. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by NOMIKOS S.A., Processing Tomato Company, Greece (private grant).

Data Availability Statement

The data presented in this study are available on request from the corresponding authors.

Acknowledgments

This paper is dedicated to the memory of Fotis Andrinopoulos (1979–2025) as a token of our sincere appreciation, respect, and gratitude for his continuous support and collaboration during the biodiversity project Operation Pollinator of Syngenta, a predecessor of the current study. His enthusiasm, creativity, and luminous personality will be greatly missed.

Conflicts of Interest

Author I.T. was employed by the company Nomikos S.A., Processing Tomato Company, Greece. The remaining authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

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Figure 1. (A) Experimental sites in the county of Larissa, Greece, and (B) layout of the sown plant mixtures and the control weed flora, with designated plots for measurements.
Figure 1. (A) Experimental sites in the county of Larissa, Greece, and (B) layout of the sown plant mixtures and the control weed flora, with designated plots for measurements.
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Figure 2. Mean total flower cover % of plot area of the processing tomato field margin with weed flora or sown with plant mixtures. Treatments: CT—control weed margin next to tomato crop; CC—control weed margin along a neighboring irrigation channel; WM—winter sown mixture; SM—spring sown mixture. Identical letters above the error bar indicate no statistically significant differences among treatments.
Figure 2. Mean total flower cover % of plot area of the processing tomato field margin with weed flora or sown with plant mixtures. Treatments: CT—control weed margin next to tomato crop; CC—control weed margin along a neighboring irrigation channel; WM—winter sown mixture; SM—spring sown mixture. Identical letters above the error bar indicate no statistically significant differences among treatments.
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Figure 3. Mean flower cover per species (% of plot area) in the tomato field margin during spring 2021 and 2022 of (A) sown mixtures in winter (WM) or spring (SM), and of (B) weed species in CT (field margin with weed flora along the control tomato crop) and CC sites (weed flora along the irrigation channel next to the control field). Main weed species that emerged in the sown margins are indicated with an asterisk (*). Other flowering species include weeds with a low abundance and a small contribution to the overall flower cover (<2%). In 2021, these included WM: Papaver rhoeas L., SM: S. nigrum, Chrozophora tinctoria (L.) A.Juss., Euphorbia sp., H. europaeum, C. arvensis, R. rugosum, CT: Calendula sp., Cirsium sp., Mantisalca salmantica (L.) Briq. and Cavill., Sonchus sp., P. rhoeas, E. elaterium, F. officinalis, Consolida regalis Gray, CC: Euphorbia sp., Geranium sp., Malva sp., Sonchus sp., and in 2022 WM: C. arvensis, E. elaterium, S. nigrum, F. officinalis, P. oleraceae, S. arvensis, R. rugosum, SM: C. arvensis, E. elaterium, S. nigrum, S. arvensis, CT: C. arvensis, Euphorbia humifusa Willd., Matricaria chamomilla L., Malva sp., P. oleracea, Tribulus terrestris L., and in CC: Lactuca serriola L., M. chamomilla, Malva sp.
Figure 3. Mean flower cover per species (% of plot area) in the tomato field margin during spring 2021 and 2022 of (A) sown mixtures in winter (WM) or spring (SM), and of (B) weed species in CT (field margin with weed flora along the control tomato crop) and CC sites (weed flora along the irrigation channel next to the control field). Main weed species that emerged in the sown margins are indicated with an asterisk (*). Other flowering species include weeds with a low abundance and a small contribution to the overall flower cover (<2%). In 2021, these included WM: Papaver rhoeas L., SM: S. nigrum, Chrozophora tinctoria (L.) A.Juss., Euphorbia sp., H. europaeum, C. arvensis, R. rugosum, CT: Calendula sp., Cirsium sp., Mantisalca salmantica (L.) Briq. and Cavill., Sonchus sp., P. rhoeas, E. elaterium, F. officinalis, Consolida regalis Gray, CC: Euphorbia sp., Geranium sp., Malva sp., Sonchus sp., and in 2022 WM: C. arvensis, E. elaterium, S. nigrum, F. officinalis, P. oleraceae, S. arvensis, R. rugosum, SM: C. arvensis, E. elaterium, S. nigrum, S. arvensis, CT: C. arvensis, Euphorbia humifusa Willd., Matricaria chamomilla L., Malva sp., P. oleracea, Tribulus terrestris L., and in CC: Lactuca serriola L., M. chamomilla, Malva sp.
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Figure 4. Total abundance of Hymenoptera pollinators (mean ± SE) over 2021 and 2022 in sown flower mixtures and weed flora of processing tomato field margins. Treatments: CT—control weed margin next to tomato crop; CC—control weed margin along a neighboring irrigation channel; WM—winter sown mixture; SM—spring sown mixture. Identical letters above the error bar indicate no statistically significant differences among treatments.
Figure 4. Total abundance of Hymenoptera pollinators (mean ± SE) over 2021 and 2022 in sown flower mixtures and weed flora of processing tomato field margins. Treatments: CT—control weed margin next to tomato crop; CC—control weed margin along a neighboring irrigation channel; WM—winter sown mixture; SM—spring sown mixture. Identical letters above the error bar indicate no statistically significant differences among treatments.
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Figure 5. Total abundance (mean ± SE) of Hymenoptera parasitoids and predatory arthropods (A) in field margins and (B) in tomato crop with different field margin management: CT—control weed margin next to tomato crop; CC—control weed margin along the irrigation channel; WM—winter sown mixture; SM—spring sown mixture; C—control tomato field. Identical letters above the error bar indicate no statistically significant differences among treatments.
Figure 5. Total abundance (mean ± SE) of Hymenoptera parasitoids and predatory arthropods (A) in field margins and (B) in tomato crop with different field margin management: CT—control weed margin next to tomato crop; CC—control weed margin along the irrigation channel; WM—winter sown mixture; SM—spring sown mixture; C—control tomato field. Identical letters above the error bar indicate no statistically significant differences among treatments.
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Figure 6. Community composition of natural enemies. In field margins: (A) Hymenoptera parasitoids; (B) predatory arthropods. In tomato crop with different field margin management: (C) Hymenoptera parasitoids; (D) predatory arthropods. CT—weedy margin next to tomato crop; CC—weedy margin along the irrigation channel; WM—winter sown mixture; SM—spring sown mixture; C—control tomato field.
Figure 6. Community composition of natural enemies. In field margins: (A) Hymenoptera parasitoids; (B) predatory arthropods. In tomato crop with different field margin management: (C) Hymenoptera parasitoids; (D) predatory arthropods. CT—weedy margin next to tomato crop; CC—weedy margin along the irrigation channel; WM—winter sown mixture; SM—spring sown mixture; C—control tomato field.
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Figure 7. Total abundance (mean ± SE) of insect pests (A) in field margins and (B) in tomato crop with different field margin management: CT—weedy margin next to tomato crop; CC—weedy margin along the irrigation channel; WM—winter mix; SM—spring mix; C—control tomato field. Identical letters above the error bar indicate no statistically significant differences among treatments.
Figure 7. Total abundance (mean ± SE) of insect pests (A) in field margins and (B) in tomato crop with different field margin management: CT—weedy margin next to tomato crop; CC—weedy margin along the irrigation channel; WM—winter mix; SM—spring mix; C—control tomato field. Identical letters above the error bar indicate no statistically significant differences among treatments.
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Table 1. Plant species selected for the mixtures sown in winter (WM) or spring (SM) along the margin of a processing tomato field, and the corresponding percentage.
Table 1. Plant species selected for the mixtures sown in winter (WM) or spring (SM) along the margin of a processing tomato field, and the corresponding percentage.
(%)g/100 m2
FamilySpeciesWMSMWMSM
ApiaceaeAnethum graveolens L.26 6
Coriandrum sativum L.18 32
BrassicaceaeEruca vesicaria (L.) Cav.2326914
FabaceaePisum sativum L.7 368
Lathyrus sativus L.1212301361
PolygonaceaeFagopyrum esculentum Moench 33 214
BoraginaceaePhacelia tanacetifolia Benth. 29 31
PoaceaeTriticum aestivum L.14 150
TOTAL100100866620
Table 2. Wild hymenoptera pollinators and associated plant species of sown mixtures and weed flora in processing tomato field margins: CT—control weed margin next to tomato crop; CC—control weed margin along an irrigation channel; WM—winter sown mixture; SM—spring sown mixture, Larisa plain, Greece, 2021 and 2022.
Table 2. Wild hymenoptera pollinators and associated plant species of sown mixtures and weed flora in processing tomato field margins: CT—control weed margin next to tomato crop; CC—control weed margin along an irrigation channel; WM—winter sown mixture; SM—spring sown mixture, Larisa plain, Greece, 2021 and 2022.
Plant SpeciesBee GenusTreatment
Ammi majusAndrena, Hylaeus, Lasioglossum, NomiapisCC
Anethum graveolensAndrena, HylaeusWM
Centauria sp.HalictusCC
Coriandrum sativumAndrena, Ceratina, NomiapisWM
Ecballium elateriumLasioglossum, CeratinaSM
Eruca vesicariaAmegilla, Andrena, Ceratina, Eucera, Halictus, LasioglossumSM
Fagopyrum esculentumAndrena, LasioglossumSM
Lathyrus sativusEucera, Megachile, OsmiaSM
Phacelia tanacetifoliaCeratinaSM
Rapistrum rugosumAndrena, Eucera, Lasioglossum, NomadaCT
Solanum lycopersicumAndrena, Lasioglossum, NomiapisWM
Table 3. Yield parameters (mean ± SE) of processing tomato hybrid H1015 fruits (weight, BRIX, pH, L, a/b) harvested from fields with different margin management: WM—winter sown mixture; SM—spring sown mixture; C—control tomato field. Identical letters indicate no statistically significant differences among treatments.
Table 3. Yield parameters (mean ± SE) of processing tomato hybrid H1015 fruits (weight, BRIX, pH, L, a/b) harvested from fields with different margin management: WM—winter sown mixture; SM—spring sown mixture; C—control tomato field. Identical letters indicate no statistically significant differences among treatments.
Treatment
20212022
Yield ParameterWMSMCWMSMC
Weight (g/fruit)74.9 ± 0.7 a76.8 ± 0.9 a63.4 ± 0.6 b79.3 ± 1.2 A76.2 ± 2.2 A57.3 ± 1.3 B
BRIX5.2 ± 0.04 a5.2 ± 0.02 a5.2 ± 0.07 a5.2 ± 0.1 B5.3 ± 0.1 B6.1 ± 0.1 A
pH4.4 ± 0.03 a4.5 ± 0.03 a4.5 ± 0.04 a4.5 ± 0.02 A4.5 ± 0.02 A4.5 ± 0.03 A
L27.8 ± 0.1 b28.0 ± 0.3 b29.6 ± 0.4 a28.3 ± 0.3 A28.1 ± 0.3 A28.1 ± 0.6 A
a/b2.5 ± 0.01 a2.5 ± 0.04 a2.4 ± 0.05 a2.5 ± 0.04 A2.5 ± 0.03 A2.3 ± 0.03 B
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MDPI and ACS Style

Kati, V.; Stathakis, T.; Economou, L.; Mylonas, P.; Barda, M.; Angelioudakis, T.; Parlapani, A.B.; Tsamis, I.; Karamaouna, F. Processing Tomato Crop Benefits from Flowering Plants in Field Margins That Support Pollinators and Natural Enemies. Agronomy 2025, 15, 1558. https://doi.org/10.3390/agronomy15071558

AMA Style

Kati V, Stathakis T, Economou L, Mylonas P, Barda M, Angelioudakis T, Parlapani AB, Tsamis I, Karamaouna F. Processing Tomato Crop Benefits from Flowering Plants in Field Margins That Support Pollinators and Natural Enemies. Agronomy. 2025; 15(7):1558. https://doi.org/10.3390/agronomy15071558

Chicago/Turabian Style

Kati, Vaya, Theodoros Stathakis, Leonidas Economou, Philippos Mylonas, Myrto Barda, Theodoros Angelioudakis, Athanasia Bratidou Parlapani, Ilias Tsamis, and Filitsa Karamaouna. 2025. "Processing Tomato Crop Benefits from Flowering Plants in Field Margins That Support Pollinators and Natural Enemies" Agronomy 15, no. 7: 1558. https://doi.org/10.3390/agronomy15071558

APA Style

Kati, V., Stathakis, T., Economou, L., Mylonas, P., Barda, M., Angelioudakis, T., Parlapani, A. B., Tsamis, I., & Karamaouna, F. (2025). Processing Tomato Crop Benefits from Flowering Plants in Field Margins That Support Pollinators and Natural Enemies. Agronomy, 15(7), 1558. https://doi.org/10.3390/agronomy15071558

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