Abstract
Pollinator assemblages of different habitats are widely studied, but sampling methodologies and covering of lesser-known pollinators and other flower-visitor taxa (e.g., floriphagous species) need improvement to obtain reliable data on the structure and functioning of these communities. We assessed flower-visiting insect assemblages in arable lands and apple orchards of the eastern Pannonian Lowland using phenylacetaldehyde-based volatile traps originally developed for pest monitoring. In seven sites, 27,539 individuals belonging to 123 species and seven insect orders were recorded. On average, over 90% of the captured insects were potential pollinators, with a higher proportion in orchards (98%) than in arable lands (85%). In orchards, the dominant groups were Hymenoptera, Diptera, Lepidoptera and Neuroptera, while in arable lands, Lepidoptera were most abundant. Floriphagous and pest species were more numerous in arable lands. These results demonstrate that phenylacetaldehyde-baited traps are easy to use, standardized, and sufficiently efficient for assessing of flower-visiting assemblages. Beyond their application in pest monitoring, they can provide reliable data on the flower-visiting and pollinator assemblages in different natural and agricultural habitats.
Keywords: phenylacetaldehyde, pollinator assemblages, alfalfa field, apple orchard, agroecosystem, pest monitoring, floral scent
Introduction
Insect pollination is an essential ecological service, that plays a key role in global food production and the maintenance of biodiversity (Kevan et al. 1990, Klein et al. 2007, Aizen et al. 2009). Consequently, numerous studies have investigated pollinators in both natural and agricultural habitats (Brunet and Stewart 2010, Ambaw and Workiye 2020). However, most of these surveys primarily focus on major pollinator groups – particularly bees – while other flower-visiting insect taxa remained comparatively understudied (Cecen et al. 2008). Nevertheless, a wide range of less conspicuous or less well-known flower-visitor insect groups (e.g., lepidopterans, flies, beetles, etc.) also contribute considerably to plant pollination (Steffan-Dewenter et al. 2005, Garibaldi et al. 2013, Rauf et al. 2021, Rácz et al. 2023). In addition, taxa belong to other flower-visiting functional groups such as species feeding on nectar, pollen and/or different parts of the flowers and buds, and even their predators and parasites are also less known, and their interactions are also not revealed. Therefore, comprehensive surveys that encompass the wide spectrum of flower-visiting assemblages are essential to advance our understanding of insects-plant interactions (Biesmeijer et al. 2006, O'Connor et al. 2019).
Moreover, the methodological heterogeneity among studies often limits the comparability of results across different habitats (Thompson et al. 2021). Some sampling methods focus exclusively on particular flower-visiting taxa – such as pan traps or insect nets (Eeraerts and Meeus 2025) – while others rely on labour-intensive and frequently subjective sampling approaches, including visual observations, transect walks, or focal plant surveys (Sarma et al. 2025). These observer-based techniques require substantial field effort, depend heavily on the observer's expertise, and can underrepresent small, fast-moving, or nocturnal species (Roy et al. 2016). Their effectiveness is also stronly influenced by environmental conditions, leading to inconsistent detectability and limited comparatibility across studies (Sarma et al. 2025). This highlights the need for standardized methods that can be applied broadly in diverse ecological contexts (Kearns C. A. and Inouye D. W. 1993, O'Connor et al. 2019).
Volatile-baited traps, originally developed for monitoring agricultural pests, have undergone considerable development and methodological advancement in the last decades (Eby et al. 2013, Landolt et al. 2013, Szanyi et al. 2024). Due to their efficiency in the case of a broad spectrum of insect taxa, these traps are increasingly used in faunistic and ecological studies as standardized sampling tools (Henning et al. 1992, Andrews et al. 2007, Nagy et al. 2023a, Nagy et al. 2023b, Szanyi et al. 2024). Their advantages include continuous sampling independent of observer bias, high efficiency in capturing both dominant and less conspicuous taxa, and applicability across various habitats.
The efficiency of volatile baited traps for attraction of lepidopteran and hemipteran pests was previously studied by our research group. During these experiments, the phenylacetaldehyde-based lures – imitating odors of flowers – proved effective not only for their target pest taxa (Szanyi et al. 2019, Szalárdi et al. 2021), but also for several non-target flower-visiting groups, including non-pest Lepidoptera (Szanyi et al. 2019, Szanyi et al. 2024), Diptera (Nagy et al. 2023b), Orthoptera (Nagy et al. 2023a), Hymenoptera (Segers et al. 2023) and Neuroptera (Szanyi et al. 2025).
In the present study, we provide data on the flower-visiting insect assemblages of agricultural habitats of the northeastern Pannonian Lowland based on the non-target catches of phenylacetaldehyde-baited traps. Additionally, we compare the composition and diversity of these assemblages living in two different agricultural habitat types – arable lands and orchards – during different flowering periods.
Material and methods
Study sites and sampling
We studied the efficiency of different phenylacetaldehyde-based lures between 2019 and 2022 at seven sites including four alfalfa fields and three commercial apple orchards in the Pannonian Lowland of Hungary (Fig. 1).
Figure 1.
Locations of the sampling sites. Red circles=arable lands, yellow triangles=orchards.
Experiments in alfalfa fields:
Püspökladány, Hajdú-Bihar County (GPS: 47.1814N, 21.0913E). Baits: phenylacetaldehyde, 1-phenylacetaldehyde, 4-methoxycinnamaldehyde, phenylacetaldehyde + 4-methoxycinnamaldehyde, phenylacetaldehyde + 1-phenylacetaldehyde + 4-methoxycinnamaldehyde + 1-nonanol, phenylacetaldehyde + 1,4-dimethoxybenzene, phenylacetaldehyde + benzyl alcohol + 6-methyl-5-hepten-2-one, phenylacetaldehyde + 1,4-dimethoxybenzene + benzyl alcohol + 6-methyl-5-hepten-2-one, and unbaited traps. Number of repetitions: 10. Date: from 03/07 to 31/08, 2019.
Gégény, Szabolcs-Szatmár Bereg County (GPS: 48.1608218N, 21.9350966E). Baits: phenylacetaldehyde, phenylacetaldehyde + anetol, phenylacetaldehyde + anisacet, phenylacetaldehyde + betion, phenylacetaldehyde + acetacid, unbaited control traps. Number of repetitions: 6. Date: from 09/07 to 31/08, 2020.
Nyírtelek, Szabolcs-Szatmár Bereg County (GPS: 48.000133N, 21.600674E). Baits: phenylacetaldehyde, phenylacetaldehyde + 4-methoxycinnamaldehyde, phenylacetaldehyde + 1,4-dimethoxybenzene, phenylacetaldehyde + 1,4-dimethoxybenzene, phenylacetaldehyde + 4-methoxycinnamaldehyde + 1,4-dimethoxybenzene, unbaited control traps. Number of repetitions: 7. Date: from 22/06 to 31/08, 2021.
Gyomaendrőd, Békés County (GPS: 46.54569N, 20.52989E). Baits: phenylacetaldehyde, phenylacetaldehyde + hexanal, phenylacetaldehyde + (E)-2-hexanal, phenylacetaldehyde + alpha-pyrene, unbaited control traps. Number of repetitions: 5. Date: from 14/07 to 31/08, 2022
Experiments in apple orchards:
Komoró, Szabolcs-Szatmár Bereg County (GPS: 48.292307N, 22.133233E),
Ajak, Szabolcs-Szatmár Bereg County (GPS: 48.163577N, 22.059229E), and
Nyírpazony, Szabolcs-Szatmár Bereg County (GPS: 47.978482N, 21.824846E). Investigated habitat: apple orchard. Baits: phenylacetaldehyde + eugenol + benzyl acetate, unbaited control traps. Number of repetitions: 3. Date: from 23/04 to 06/05, 2022.
The samplings were carried out in the flowering period of the chosen cultures. CSALOMON® VARL+ baited traps (Plant Protection Institute, CAR, Budapest, Hungary), available at www.csalomontraps.com (accessed on 11 May 2025), were used in each sampling sites (Fig. 2.), with phenylacetaldehyde-based synthetic lures that imitate the scent of strongly fragnant, mostly white flowers (Szanyi et al. 2019, Nagy et al. 2022). The tested lures were enclosed in 1.5 × 1.5 cm polyethylene bags with a 0.2 mm wall thickness, allowing the volatiles to diffuse through the bag's walls (Tóth M. et al. 2020). In apple orchards, the traps were placed on branches of trees, with a spacing of 20 m between them at heights of 1.8–2.0 m. In alfalfa fields, traps were placed at the field margins on 1 m height mounted on wooden stakes and spaced 15 m apart. Traps were checked and emptied weekly in the apple orchards, and twice a week in the alfalfa fields. The baits were replaced every four weeks. To kill the trapped insects, we used Vaportape® II pesticide strips designed specifically for insect traps.
Figure 2.

CSALOMON® VARL+ baited trap used to collect the flower visiting insects.
Evaluation of samples and data analysis
Samples were stored at -20 °C in a fridge until identification in a laboratory. The identification of the insects caught was made at species level, when it was possible, based on the keys of Wagner (1952) and Schwartz M. D and Foottit R. G (1998) for heteropterans, Harz (1957), Harz (1960) and Harz (1975) for orthopterans, Aspoeck et al. (1980) and Thierry et al. (1998) for neuropterans, Móczár (1985), Goulet and Hubert (1993) and Bossert (2015) for hymenopterans, Papp and Darvas (2000) and Tóth (2017) for dipterans, and Varga Z. (2010) for lepidopterans.
Relative frequencies (RF%) of insect groups were calculated using all collected specimens, including those identified only at the genus or family level. To evaluate the composition and mean ratios (MR%) of different taxonomical and functional groups, we compared both the total number and mean number of individuals (individuals/sampling sites). Two functional group classifications were applied. The first classification was based on the presumed reason attraction to floral scent: potential pollinators were considered species attracted by floral rewards for the pollination (e.g., nectar, pollen), while floriphagous species (e.g., Miridae, Cerambycidae) were those attracted due to their consumption of floral tissues. The second classification distinguished species by pest status: species capable of causing serious damage to the host plant at any phenological stage were categorized as pests, whereas those not causing such damage were considered non-pests. Data normality and variance homogeneity were assessed using Q-Q plots and Levene’s test, respectively. As assumptions were not met, the non-parametric Kruskal–Wallis test was used. Where significant differences were detected, pairwise comparisons were conducted using the Mann–Whitney U-test. All statistical analyses were performed using SPSS 21.0 (Ketskeméty et al. 2011).
All records collected for the present work were published through GBIF (Ősz et al. 2025).
Results
A total of 30,595 insects were caught using volatile traps baited with phenylacetaldehyde-based lures (Table 1, Suppl. material 1). Of these, 27,539 individuals of 123 flower-visiting species were identified to species level, belonging to eight insect orders. Lepidoptera was notably the most species-rich order, with 98 species, and was dominated by the family Noctuidae, with 52 species of 9 subfamilies. The further most diverse groups were Hymenoptera (S=9) and Hemiptera (S=8), while Diptera, Orthoptera, Coleoptera, and Mecoptera were represented by only a few species.
Table 1.
Checklist by habitat types (A=arable land, O=orchard), including information on the occurence (1=presence), pest status (PE=pest, causes damage to crops at any developmental stage; NPE=non-pest, does not cause serious damage) and pollinator role of the species (PO=known pollinator; PPO=potential pollinator; FLO=floriphagous, feeds on the flowers). Protected species in Hungary are marked with an asterisk (*).
| Order | Family | Species | A | O | PE | PO |
| Coleoptera | Cerambycidae | Plagionotus floralis (Pallas, 1773) | 1 | PE | FLO | |
| Diptera | Bombyliidae | Bombylius major (Linnaeus, 1758) | 1 | NPE | PO | |
| Diptera | Syrphidae | Episyrphus balteatus (De Geer, 1776) | 1 | NPE | PO | |
| Diptera | Syrphidae | Eristalis tenax (Linnaeus, 1758) | 1 | NPE | PO | |
| Diptera | Syrphidae | Melanostoma scalare (Fabricius, 1794) | 1 | NPE | PO | |
| Hemiptera | Miridae | Adelphocoris lineolatus (Goeze, 1778) | 1 | PE | FLO | |
| Hemiptera | Miridae | Adelphocoris quadripunctatus (Fabricius, 1794) | 1 | PE | FLO | |
| Hemiptera | Miridae | Adelphocoris reichelii (Fieber, 1836) | 1 | PE | FLO | |
| Hemiptera | Miridae | Adelphocoris seticornis (Fabricius, 1775) | 1 | PE | FLO | |
| Hemiptera | Miridae | Lygus gemellatus (Herrich-Schaeffer, 1835) | 1 | PE | FLO | |
| Hemiptera | Miridae | Lygus pratensis (Linnaeus, 1758) | 1 | PE | FLO | |
| Hemiptera | Miridae | Lygus rugulipennis (Poppius, 1911) | 1 | PE | FLO | |
| Hemiptera | Miridae | Polymerus vulneratus (Panzer, 1805) | 1 | PE | FLO | |
| Hymenoptera | Apidae | Apis mellifera (Linnaeus, 1758) | 1 | 1 | NPE | PO |
| Hymenoptera | Apidae | Bombus argillaceus (Scopoli, 1763)* | 1 | NPE | PO | |
| Hymenoptera | Apidae | Bombus hortorum (Linnaeus, 1761) | 1 | NPE | PO | |
| Hymenoptera | Apidae | Bombus lapidarius (Linnaeus, 1758) | 1 | NPE | PO | |
| Hymenoptera | Apidae | Bombus pascuorum (Scopoli, 1763) | 1 | NPE | PO | |
| Hymenoptera | Apidae | Bombus ruderatus (Fabricius, 1775)* | 1 | NPE | PO | |
| Hymenoptera | Apidae | Bombus sylvarum (Linnaeus, 1761) | 1 | NPE | PO | |
| Hymenoptera | Apidae | Bombus terrestris (Linnaeus, 1758) | 1 | NPE | PO | |
| Hymenoptera | Vespidea | Vespa crabro (Linnaeus, 1758) | 1 | PE | FLO | |
| Lepidoptera | Crambidae | Cydalima perspectalis (Walker, 1859) | 1 | PE | PPO | |
| Lepidoptera | Crambidae | Haritala ruralis (Scopoli, 1763) | 1 | NPE | PPO | |
| Lepidoptera | Crambidae | Loxostege sticticalis (Linnaeus, 1761) | 1 | PE | PPO | |
| Lepidoptera | Crambidae | Nomophila noctuella (Denis & Schiffermüller, 1775) | 1 | NPE | PPO | |
| Lepidoptera | Crambidae | Nymphula nitidulata (Hufnagel, 1767) | 1 | NPE | PPO | |
| Lepidoptera | Crambidae | Sitochroa verticalis (Linnaeus, 1758) | 1 | NPE | PPO | |
| Lepidoptera | Erebidae | Amata phegea (Linnaeus, 1758) | 1 | NPE | PO | |
| Lepidoptera | Erebidae | Dysauxes ancilla (Linnaeus, 1767) | 1 | NPE | PPO | |
| Lepidoptera | Erebidae | Dysgonia algira (Linné, 1767) | 1 | NPE | PPO | |
| Lepidoptera | Erebidae | Eilema lurideola (Zincken, 1817) | 1 | NPE | PPO | |
| Lepidoptera | Noctuidae | Euclidia glyphica (Linnaeus, 1758) | 1 | NPE | PO | |
| Lepidoptera | Erebidae | Herminia tarsipennalis (Treitschke, 1835) | 1 | NPE | PPO | |
| Lepidoptera | Erebidae | Hypena proboscidalis (Linnaeus, 1758) | 1 | NPE | PPO | |
| Lepidoptera | Erebidae | Hypena rostralis (Linnaeus, 1758) | 1 | NPE | PPO | |
| Lepidoptera | Erebidae | Lygephila craccae (Denis & Schiffermüller, 1775) | 1 | NPE | PPO | |
| Lepidoptera | Erebidae | Lymantria dispar (Linnaeus, 1758) | 1 | PE | PPO | |
| Lepidoptera | Erebidae | Prodotis stolida (Fabricius, 1775) | 1 | NPE | PPO | |
| Lepidoptera | Erebidae | Rivula sericealis (Scopoli, 1763) | 1 | NPE | PPO | |
| Lepidoptera | Erebidae | Scoliopteryx libatrix (Linnaeus, 1758) | 1 | NPE | PPO | |
| Lepidoptera | Gelechiidae | Sitotroga cerealella (Olivier, 1789) | 1 | PE | PPO | |
| Lepidoptera | Gelechiidae | Stomopteryx radicalis (Falkovitsh & Bidzilya, 2003) | 1 | NPE | PPO | |
| Lepidoptera | Gelechiidae | Camptogramma bilineata (Linnaeus, 1758) | 1 | NPE | PPO | |
| Lepidoptera | Geometridae | Chiasmia clathrata (Linnaeus, 1758) | 1 | PE | PO | |
| Lepidoptera | Geometridae | Cidaria fulvata (Forster, 1771) | 1 | NPE | PPO | |
| Lepidoptera | Geometridae | Ematurga atomaria (Linnaeus 1758) | 1 | PE | PPO | |
| Lepidoptera | Geometridae | Epirrhoe alternata (Müller, 1764) | 1 | NPE | PPO | |
| Lepidoptera | Geometridae | Idaea aversata (Linnaeus, 1758) | 1 | NPE | PPO | |
| Lepidoptera | Geometridae | Idaea rusticata (Denis & Schiffermüller, 1775) | 1 | NPE | PPO | |
| Lepidoptera | Geometridae | Ligdia adustata (Denis & Schiffermüler, 1775) | 1 | NPE | PPO | |
| Lepidoptera | Geometridae | Peribatodes rhomboidaria (Denis & Schiffermüller, 1775) | 1 | PE | PPO | |
| Lepidoptera | Geometridae | Peribatodes secundaria (Denis & Schiffermüller, 1775) | 1 | NPE | PPO | |
| Lepidoptera | Geometridae | Plemyria rubiginata (Denis & Schiffermüller, 1775) | 1 | NPE | PPO | |
| Lepidoptera | Geometridae | Pseudopanthera macularia (Linnaeus, 1758) | 1 | NPE | PPO | |
| Lepidoptera | Geometridae | Tephrina arenacearia (Denis & Schiffermüller, 1775) | 1 | PE | PPO | |
| Lepidoptera | Lasiocampidae | Lasiocampa trifolii (Denis & Schiffermüller, 1775) | 1 | NPE | PPO | |
| Lepidoptera | Lycaenidae | Polyommatus icarus (Rottemburg, 1775) | 1 | NPE | PO | |
| Lepidoptera | Noctuidae | Acontia lucida (Hufnagel, 1766) | 1 | NPE | PPO | |
| Lepidoptera | Noctuidae | Acontia trabealis (Scopoli, 1763) | 1 | 1 | NPE | PPO |
| Lepidoptera | Noctuidae | Aedia leucomelas (Linnaeus, 1758) | 1 | NPE | PPO | |
| Lepidoptera | Noctuidae | Abrostola tripartita (Hufnagel, 1766) | 1 | 1 | NPE | PO |
| Lepidoptera | Noctuidae | Abrostola triplasia (Linnaeus, 1758) | 1 | NPE | PO | |
| Lepidoptera | Noctuidae | Agrotis bigramma (Esper, 1790) | 1 | PE | PPO | |
| Lepidoptera | Noctuidae | Agrotis exclamationis (Linnaeus, 1758) | 1 | PE | PPO | |
| Lepidoptera | Noctuidae | Agrotis segetum (Denis & Schiffermüller, 1775) | 1 | PE | PPO | |
| Lepidoptera | Noctuidae | Apamea sordens (Hufnagel, 1766) | 1 | NPE | PPO | |
| Lepidoptera | Noctuidae | Apterogenum ypsillon (Denis & Schiffermüller, 1775) | 1 | NPE | PPO | |
| Lepidoptera | Noctuidae | Autographa gamma (Linnaeus, 1758) | 1 | 1 | PE | PO |
| Lepidoptera | Noctuidae | Axylia putris (Linnaeus, 1761) | 1 | NPE | PPO | |
| Lepidoptera | Noctuidae | Calophasia lunula (Hufnagel, 1766) | 1 | NPE | PPO | |
| Lepidoptera | Noctuidae | Caradrina morpheus (Hufnagel, 1766) | 1 | NPE | PPO | |
| Lepidoptera | Noctuidae | Chrysodeixis chalcites (Esper, 1789) | 1 | PE | PO | |
| Lepidoptera | Noctuidae | Cirrhia icteritia (Hufnagel, 1766) | 1 | NPE | PPO | |
| Lepidoptera | Noctuidae | Cornutiplusia circumflexa (Linnaeus, 1767) | 1 | NPE | PO | |
| Lepidoptera | Noctuidae | Cucullia chamomillae (Denis & Schiffermüller, 1775) | 1 | NPE | PPO | |
| Lepidoptera | Noctuidae | Cucullia umbratica (Linnaeus, 1758) | 1 | NPE | PO | |
| Lepidoptera | Noctuidae | Diachrysia chrysitis (Linnaeus, 1758) | 1 | 1 | NPE | PO |
| Lepidoptera | Noctuidae | Diachrysia stenochrysis (Warren, 1913) | 1 | NPE | PO | |
| Lepidoptera | Noctuidae | Eupsilia transversa (Hufnagel, 1766) | 1 | NPE | PPO | |
| Lepidoptera | Noctuidae | Euxoa temera (Hübner, 1808) | 1 | PE | PPO | |
| Lepidoptera | Noctuidae | Hadena bicruris (Hufnagel, 1766) | 1 | NPE | PPO | |
| Lepidoptera | Noctuidae | Hadula trifolii (Hufnagel, 1766) | 1 | PE | PO | |
| Lepidoptera | Noctuidae | Helicoverpa armigera (Hübner, 1808) | 1 | 1 | PE | PO |
| Lepidoptera | Noctuidae | Heliothis maritima (Draudt, 1938) | 1 | PE | PO | |
| Lepidoptera | Noctuidae | Heliothis viriplaca (Hufnagel, 1766) | 1 | PE | PO | |
| Lepidoptera | Noctuidae | Hoplodrina ambigua (Denis & Schiffermüller, 1775) | 1 | NPE | PPO | |
| Lepidoptera | Noctuidae | Hoplodrina octogeneria (Goeze, 1781) | 1 | NPE | PPO | |
| Lepidoptera | Noctuidae | Hydraecia micacea (Esper, 1789) | 1 | NPE | PPO | |
| Lepidoptera | Noctuidae | Lacanobia oleracea (Linnaeus, 1758) | 1 | PE | PPO | |
| Lepidoptera | Noctuidae | Macdunnoughia confusa (Stephens, 1850) | 1 | 1 | PE | PPO |
| Lepidoptera | Noctuidae | Mamestra brassicae (Linnaeus, 1758) | 1 | PE | PPO | |
| Lepidoptera | Noctuidae | Mesapamea secalis (Linnaeus, 1758) | 1 | PE | PPO | |
| Lepidoptera | Noctuidae | Mesoligia furuncula (Denis & Schiffermüller, 1775) | 1 | NPE | PPO | |
| Lepidoptera | Noctuidae | Mythimna albipuncta (Denis & Schiffermüller, 1775) | 1 | NPE | PPO | |
| Lepidoptera | Noctuidae | Mythimna l-album (Linnaeus, 1767) | 1 | NPE | PPO | |
| Lepidoptera | Noctuidae | Mythimna pallens (Linnaeus, 1758) | 1 | PE | PPO | |
| Lepidoptera | Noctuidae | Mythimna vitellina (Hübner, 1808) | 1 | PE | PPO | |
| Lepidoptera | Noctuidae | Noctua interposita (Hübner, 1790) | 1 | PE | PPO | |
| Lepidoptera | Noctuidae | Noctua janthina (Borkhausen, 1792) | 1 | PE | PPO | |
| Lepidoptera | Noctuidae | Noctua pronuba (Linnaeus, 1758) | 1 | PE | PPO | |
| Lepidoptera | Noctuidae | Orthosia cruda (Denis & Schiffermüller, 1775) | 1 | PE | PPO | |
| Lepidoptera | Noctuidae | Pyrrhia umbra (Hufnagel, 1766) | 1 | NPE | PPO | |
| Lepidoptera | Noctuidae | Rhyacia simulans (Hufnagel, 1766) | 1 | NPE | PPO | |
| Lepidoptera | Noctuidae | Shargacucullia verbasci (Linnaeus, 1758) | 1 | NPE | PPO | |
| Lepidoptera | Noctuidae | Thalpophila matura (Hufnagel, 1766) | 1 | NPE | PPO | |
| Lepidoptera | Noctuidae | Trachea atriplicis (Linnaeus, 1758) | 1 | NPE | PPO | |
| Lepidoptera | Noctuidae | Tyta luctuosa (Denis & Schiffermüller, 1775) | 1 | NPE | PPO | |
| Lepidoptera | Noctuidae | Xestia c-nigrum (Linnaeus, 1758) | 1 | PE | PPO | |
| Lepidoptera | Noctuidae | Xestia xanthographa (Denis & Schiffermüller, 1775) | 1 | NPE | PPO | |
| Lepidoptera | Nymphalidae | Araschnia levana (Linnaeus, 1758) | 1 | NPE | PO | |
| Lepidoptera | Nymphalidae | Coenonympha pamphilus (Linnaeus, 1758) | 1 | NPE | PO | |
| Lepidoptera | Nymphalidae | Nymphalis c-album (Linneus, 1758)* | 1 | NPE | PO | |
| Lepidoptera | Pieridae | Colias croceus (Geoffroy in Fourcroy, 1785) | 1 | NPE | PO | |
| Lepidoptera | Pyralidae | Etiella zinckenella (Treitschke, 1832) | 1 | PE | PPO | |
| Lepidoptera | Pyralidae | Hypsopygia costalis (Fabricius, 1775) | 1 | PE | PPO | |
| Lepidoptera | Pyralidae | Ostrinia nubilalis (Hübner, 1796) | 1 | PE | PPO | |
| Lepidoptera | Pyralidae | Salebria semirubella (Scopoli, 1763) | 1 | PE | PPO | |
| Lepidoptera | Sesiidae | Synanthedon vespiformis (Linnaeus, 1761) | 1 | NPE | PO | |
| Mecoptera | Panorpidae | Panorpa communis (Linnaeus, 1758) | 1 | NPE | PPO | |
| Orthoptera | Acrididae | Calliptamus italicus (Linnaeus, 1758) | 1 | PE | FLO | |
| Orthoptera | Gryllidae | Oecanthus pellucens (Scopoli, 1763) | 1 | NPE | PPO | |
| Orthoptera | Tettigoniidae | Tettigonia viridissima (Linnaeus, 1758) | 1 | NPE | PPO |
Most identified species were common and widespread in Hungary. The major part of the assemblage consisted of agricultural pest species (e.g., Hemiptera spp., Ostrinia nubilalis, Autographa gamma, Agrotis spp., Helicoverpa armigera), primarily associated with alfalfa fields. In contrast, the majority of individuals and species caught in apple orchards were considered beneficial. Additionally, two protected species (Bombus argillaceus, B. ruderatus) were recorded in the apple orchards, and one (Nymphalis c-album) in the alfalfa fields.
Although Lepidoptera had the highest species-richness, they accounted for 47.25% of all specimens, with 21.75% belonging to the Noctuidae family. Within Noctuidae, the Plusiinae (RF%=53.79), and Heliothinae (RF%=37.50) subfamilies were dominant. Lepidoptera was followed by Hymenoptera (RF%=15.76), Hemiptera (RF%=6.95), and Diptera (RF%=3.64).
Notable differences were found between the flower visiting assemblages of the two habitat types. Arable lands supported a more diverse assemblage, with 24,834 specimens representing 108 species, compared to only 2,705 individuals and 22 species in the orchards.
In arable lands, Lepidoptera dominated species-richness (S=95), with other groups contributing only a few species each. However, Lepidoptera accounted for only 67.83% of individuals. Hymenoptera (RF%=15.97), Hemiptera (RF%=10.10) and Neuroptera (RF%=4.93) were also relatively abundant given their species numbers.
In orchards, only 22 species were recorded. Almost half belonged to Hymenoptera (S=9), while Lepidoptera, Diptera, and Neuroptera were represented by only a few species. Relative frequencies followed a similar pattern: Hymenoptera were dominant (RF%=53.13), followed by Diptera (RF%=34.44), Neuroptera (RF%=6.12), and Lepidoptera (RF%=6.29).
Across all habitats, the mean ratio (MR%) of potential pollinators (MR%=90.462±15.049) was significantly higher than that of floriphagous insects (MR%=9.537±15.049) (Fig. 3). Similarly, the mean ratio of pest species (MR%=29.361±28.860) was significantly lower than that of non-pest species (MR%=70.638±28.860).
Figure 3.

Mean ratio (ratio/location [%]) of potential pollinator, floriphagous (A), pest, and non-pest species (B). Means with the same letter within a diagram are not significantly different at P=0.05 by Mann-Whitney U non-parametric test.
Comparing habitat types, both the mean ratio of potential pollinators (MR%=98.239±1.049) and non-pest species (MR%=95.420±0.691) were significantly higher in orchards than in arable lands (potential pollinators: MR%=84.629±18.610; non-pests: MR%=52.052±24.305) (Fig. 4). Accordingly, arable lands had a much higher ratio of pests (MR%=47.947±24.305) compared to orchards (MR%=4.579±0.691).
Figure 4.

Mean ratio of potential pollinator, floriphagous, pest, and non-pest species in the different habitat types (orchard and arable land). Means with the same letter within a diagram are not significantly different at P=0.05 by Mann-Whitney U non-parametric test.
The dominant pollinator groups in orchards were Hymenoptera (MR%=51.537±4.275), Diptera (MR%=34.453±2.913), and Lepidoptera (MR%=6.313±0.564) and Neuroptera (MR%=5.934±2.913). In arable lands, the most abundant pollinators were Lepidoptera (MR%=63.445±28.774), Hymenoptera (MR%=15.876±27.294), and Neuroptera (MR%=4.209±6.031).
Floriphagous and pest species had significantly higher mean ratios in arable lands (floriphagous: MR%=15.371±18.610; pests: MR%=47.947±24.305) than in orchards (floriphagous: MR%=1.760±1.049; pests: MR%=4.579±0.691). The most abundant pest groups in arable lands were Lepidoptera (MR%=32.576±16.283) and Hemiptera (MR%=15.222±18.697), with the latter also comprising the only notable floriphagous group. In orchards, the most abundant pest groups were Lepidoptera (MR%=2.819±0.424) and Hymenoptera (MR%=1.760±1.049), the latter also being the sole floriphagous group.
Discussion
A total of 123 flower-visiting insect species were recorded in arable lands and apple orchards of the eastern Pannonian Lowland. The insect assemblages included a wide range of taxa from multiple orders, reflecting the diverse communities occurring in these agroecosystems. Notably, three protected species – Bombus argillaceus, B. ruderatus, and Nymphalis c-album – were recorded, underscoring the ecological importance of these habitats.
The observed species richness, especially within Lepidoptera, highlights the attractiveness of phenylacetaldehyde as a floral scent. The dominance of the Noctuidae family within Lepidoptera is consistent with their known responsiveness to floral volatiles (Szanyi et al. 2019, Szanyi et al. 2024). Nonetheless, Hymenoptera and Diptera also contributed substantially, especially in orchards, suggesting that these groups play there the major role in pollination.
Our findings demonstrate that phenylacetaldehyde-based volatile traps, originally designed for pest monitoring, are highly effective at capturing a broad spectrum of flower-visiting insects belonging several orders. This supports earlier findings (Landolt et al. 2013, Hesler 2016, Nagy et al. 2023a, Nagy et al. 2023b, Szanyi et al. 2024) and highlights the potential of such traps for broader ecological and faunistic applications beyond pest surveillance.
The study revealed notable differences in the composition of flower-visiting insect communities in arable lands and orchards. Arable lands supported a more diverse assemblage, which may be partly due to the higher sampling effort and longer sampling duration in these habitats, as the flowering stage of the investigated arable crop – alfalfa – is much longer than that of the apple orchard. However, the relative frequency and ratio patterns suggest that these differences are not solely methodological. From a functional perspective, potential pollinator species made up the majority of the assemblages in both habitat types, although their proportion was significantly higher in orchards. Accordingly, pest and floriphagous species were more abundant in arable lands.
These observed differences underline the importance of multi-taxa monitoring in agroecosystems. Focusing solely on bees or model pollinators would overlook important contributors to pollination and ecosystem functioning. Moreover, the detection of protected species like Bombus argillaceus, B. ruderatus and Nymphalis c-album reinforces the conservation value of these habitats and the relevance of including non-target data collected with volatile traps in faunistic studies.
In conclusion, standardized volatile-baited traps can be a valuable tool for studying flower-visiting insects and their interactions with plants in various ecosystems. Their ability to provide consistent and replicable data makes them especially suitable for long-term monitoring and comparative studies in both natural and agricultural habitats.
Supplementary Material
Relative frequences
Ősz A., Szanyi K., Arnóczkyné Jakab D., Szanyi Sz., Koczor S., Tóth M., Szarukán I., Nagy A.
Data type
Occurences
Brief description
Relative frequences (RF%) of insect orders, families (only in the case of Hymenoptera and Lepidoptera orders) and subfamilies (only in the case of Noctuidae family) calculated based on all caught individuals and individual numbers of insects identified to species level.
File: oo_1494994.xlsx
Acknowledgements
Supported by the University of Debrecen Program for Scientific Publication.
Hosting institution
Faculty of Agriculture and Food Sciences and Environmental Management, Institute of Plant Protection, University of Debrecen
Author contributions
AŐ and AN conducted the experiments. SZSZ, DAJ, SK, AŐ identified the collected insect material. KSZ analysed data and conducted statistical analyses. AŐ, KSZ, MT, ISZ and AN wrote the manuscript. All authors read and approved the manuscript.
Conflicts of interest
No conflict of interest to declare
Disclaimer: This article is (co-)authored by any of the Editors-in-Chief, Managing Editors or their deputies in this journal.
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Associated Data
This section collects any data citations, data availability statements, or supplementary materials included in this article.
Supplementary Materials
Relative frequences
Ősz A., Szanyi K., Arnóczkyné Jakab D., Szanyi Sz., Koczor S., Tóth M., Szarukán I., Nagy A.
Data type
Occurences
Brief description
Relative frequences (RF%) of insect orders, families (only in the case of Hymenoptera and Lepidoptera orders) and subfamilies (only in the case of Noctuidae family) calculated based on all caught individuals and individual numbers of insects identified to species level.
File: oo_1494994.xlsx

