Enhancing knowledge of non-target tortricid fauna via pheromone monitoring of the quarantine fruit pest Grapholita molesta (Lepidoptera: Tortricidae) in Russia

Introduction

The oriental fruit moth Grapholita molesta (Busck, 1916) (Lepidoptera: Tortricidae) is a pest affecting a wide range of economically important fruit crops – peaches, apples, quinces, pears, plums, cherries, apricots, and nectarines (Rosales: Rosaceae) (Hari & Penzes 2010; Witzgall et al. 2010; Amat et al. 2021). Recently it was found attacking a phylogenetically distant fruit plant, Litchi chinensis (Sapindaceae: Sapindales), in Brasil (Souza et al. 2023).

The economic impact of G. molesta is considerable: in heavily infested orchards, yield losses can be significant (from 20 to 90% depending on crop and management practices), and global economic damage is estimated in millions of dollars annually. Thus, as a consequence, infestations of G. molesta are associated with substantial agricultural losses and increased management costs in fruit-producing regions world-wide (Reis et al. 1988; Kanga et al. 2003; Anon 2014; Ma et al. 2023).

Native to East Asia, G. molesta is believed to originate from China (Wei et al. 2015; Yang et al. 2016), from where it has spread extensively through international trade and the movement of plant material. Over the past century, the species invaded North and South America, Europe, Africa, and Oceania (Danilevsky & Kuznetsov 1968; Shutova 1980; Byun et al. 2012; Kirk et al. 2013; Biosecurity Australia 2026; Biosecurity New Zealand 2026; EPPO 2026), demonstrating a strong capacity for adaptation to diverse climatic conditions. Its invasive potential remains high, with continued detections in new areas and expansion within previously colonized regions (Akulov et al. 2013).

In several countries, including Russia, G. molesta is classified as a quarantine pest, necessitating strict phytosanitary control and monitoring (Federal Service for Veterinary and Phytosanitary Supervision 2025, 2026). One of the primary tools for detection and population assessment is the use of pheromone traps baited with synthetic sex pheromone of G. molesta. Such monitoring programs are widely used in Europe, North America, and Asia, where they have proven to be highly effective for early detection, population dynamics studies, and decision-making in integrated pest management systems (Akulov et al. 2013; Knight et al. 2015; Preti et al. 2020; Kim & Heo 2026).

At the same time, pheromone-based monitoring may generate valuable by-catch data: traps designed for G. molesta frequently capture non-target tortricid species, some of which may respond to similar or shared pheromone components (Mizukoshi 2006; Jung et al. 2012). This side effect provides an additional opportunity to study the diversity, distribution, and phenology of related Tortricidae species (Velcheva 2000; Hrudova 2003; Akulov & Kirichenko 2014; Zheng et al. 2016; Jakubikova et al 2016; Pražanova & Šefrova 2024).

In Russia, monitoring of G. molesta using pheromone traps is carried out across regions, especially where commercial fruit production is concentrated (Shutova 1980; Abasov & Atanov 2011; Akulov et al. 2013; Akulov & Kirichenko 2014). While performing monitoring of G. molesta (Akulov et al. 2013, 2014, 2025), we kept sampling and documenting associated tortricids in pheromone traps, which has provided new insights into species composition and distribution.

Thus, in the present study we provide a comprehensive review of tortricid species recorded in G. molesta pheromone traps in Russia over the last 15 years, integrating our data with published records from Eurasia (mostly European countries). We discuss the implications of pheromone monitoring for exploring non-target tortricid fauna. Additionally, we report novel regional records and assess their relative abundance, thereby contributing to a broader understanding of tortricid biodiversity in agroecosystems.

Materials and methods

Study period and regions in Russia

Our study was conducted in Russia during 2010–2025 (except 2020, due to COVID-19 restrictions). Overall, 21 administrative regions were monitored (listed in an approximate order from west to east): Smolensk (study year 2025), Moscow (2025), Rostov (2025) Oblasts, Stavropol (2021), Krasnodar Krais (2025), Kirov (2023) Oblast, Chuvashia (2025), Udmurtia (2024) Republics, Perm Krai (2023), Orenburg (2023), Kemerovo (2021), Omsk (2022), Tomsk (2021), Novosibirsk Oblasts (2021), Altai Krai (2022), Altai Republic (2022), Krasnoyarsk Krai (2010–2018), Khakassia Republic (2011, 2025), Irkutsk Oblast (2023), Buryatia Republic (2024), Primorsky Krai (2019). The details on sampled points and dates are given in the list of collected material in the species essays below. We followed the administrative-geographical zoning of the country in accordance to the system given in the Catalogue of Lepidoptera of Russia (Sinev et al. 2019). This edition and its newest online version (Catalogue 2026) provide the most up-to-date species distribution data for the country. We covered the following 15 administrative-geographical zones, i.e. region 7 – European southern taiga region (studies administrative units: Udmurtia Republic, Kirov Oblast); 8 – European Central region (Smolensk and Moscow Oblasts); 10 – Middle Volga region (Chuvashia Republic); 11 – Volga-Don region (Rostov Oblast); 13 – West Caucasus region (Stavropol and Krasnodar Krais); 16 – Middle Ural region (Perm Krai); 17 South Ural region (Orenburg Oblast); 19 – Middle Ob region (Tomsk Oblast); 20 South-West Siberian region (Omsk Oblast, Altai Krai – the Kulunda Steppe only); 22 – Krasnoyarsk region (Krasnoyarsk Krai, Khakassia Republic); 23 – Pre-Altai region (Kemerovo and Novosibirsk Oblasts, Altai Krai); 24 – Gorno-Altai region (Altai Republic); 26 – Predbaikalie region (Irkutsk Oblast); 27 – Pribaikalie region (Buryatia Republic); 40 – Primorsky region (Primorsky Krai) (Fig. 1).

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Figure 1. The study regions in Russia in 2010–2025. The regions colored in green involved in the study: region 7 – European southern taiga region; 8 – European Central Region; 10 – Middle Volga region; 11 – Volga-Don region; 13 – West Caucasus region; 16 – Middle Ural region; 17 – South Ural region; 19 – Middle Ob region; 20 – South-West Siberian region; 22 – Krasnoyarsk region; 23 – Pre-Altai region; 24 – Gorno-Altai region; 26 – Predbaikalie region; 27 – Pribaikalie region; 40 – Primorsky region.

Field sampling

The delta-shape pheromone traps with stick inner surface produced by the All-Russian Plant Quarantine Center (VNIIKR) were involved in the study. As an attractant, a synthetic sex pheromone of the oriental codling moth G. molesta (i.e., the composition of Z8-dodecenyl acetate, E8-dodecenyl acetate, and Z8-dodecenol) was used. The traps were deployed on fruit trees (at a height up to 2 m) in orchards, botanical gardens, private gardens, planting at dachas etc. Overall, 41 localities were surveyed (Fig. 1, orange circles) in 15 administrative-geographical zones. In each locality, from one to five pheromone traps were placed. The period of trapping varied from three days up to the whole vegetation period (i.e., from late April to late October) and during several years (Fig. 1). In the latter case, the traps were replaced every two weeks. Long-term observations were carried out in mother regions of some coauthors, i.e. in Moscow Oblast (whole vegetation season of 2025) and in Krasnoyarsk Krai (vegetation seasons of 2010–2018 and 2021–2024). Overall, 2658 pheromone traps were used in the study. The pheromone traps with captures insects (glued to the inner surface of the traps) were transported to All-Russian Plant Quarantine Center (FGBU "VNIIKR", Bykovo, Moscow Oblast) and its former branch in Krasnoyarsk for identification.

Data analysis, species identification, photographing

Species identification was performed on male genitalia and/or wing pattern and coloration. The wings were studied only in the individuals which stayed on the glued surface of traps by the dorsal side upwards. In most cases, the abdomens of males were detached for genital study. Male genitalia were prepared according to described methods (Golub et al. 2021; Kovalenko et al. 2024).

The moth specimens were photographed with a Canon EOS 6D digital SLR camera (Japan, Canon) with a Canon MP-E 65 mm f/2.8 1-5X Macro lens (Japan, Canon). Images of the genitalia structures were taken using hardware and software complex based on a stereo microscope Nexcope NSZ818 (China, Nexcope). In both cases, stacking approach was applied, with up to 180 frames taken at different focal planes. These image stacks were processed in the software Zerene Stacker (Version 1.04 Build T2024-11-18-1210). Resultant images were further corrected in Adobe Photoshop 2021.

Species classification

The species documented in Russia were classified using two parameters: species abundance (A) and regularity in catches (R). Species abundance was assessed as a maximum number of individuals caught in the trap over a two-week period during the study period. Following categories were used for abundance: A – mass abundance (75 individuals or more); B – intermediate abundances (from 25 to 74 individuals); C – low abundance (from 10 to 24 individuals); D – extremely low abundance (less than 10 individuals); E – unspecified abundance category (for congeneric species, which many specimens had shabby or damaged wings in the traps and which males were tricky to distinguish by genitalia characters from congeners. Thus, it was not possible to reliably estimate the abundance of such species).

Species regularity in catches: 1 – regular species (those detected in at least three different regions of its range, in different dates and years; for widespread species, the records had to be frequent from both the European and Asian parts of its range; for narrow-range species, the records had to be frequent in one locality within the species range and the species had to be detected in different years, or if survey in a region was conducted for only one year, the species had to be detected frequently during its flight period); 2 – irregular species (those detected at a moderate frequency in one to two regions of its range; widespread species felt in this category if they were detected in more than two regions of its range, but only in the European or Asian parts); 3 – rare species (those detected ones or twice during whole observation period in one-two localities).

Due to the heterogeneity of the study periods in different regions, a comparative analysis of quantitative species indicators between regions was not performed.

Literature review

We analyzed available literature on capturing tortricids in the traps with the synthetic sex pheromone of G. molesta in Europe and Asia, including early studies performed in the Russian Federation. The literature search was conducted on the Internet (primarily ResearchGate database) and in the libraries of various scientific organizations. As a result of the literature review, we took into account all non-target tortricids documented in the traps with G. molesta pheromone retrieved from various literature sources (Chambón & Daguilar 1974; Sziraki 1978; Hrdý et al. 1979a, 1979b, 1989, 1994; Rozinskaya 1980; Dulo & Shvanda 1980; Kipiani 1980; Kurbatov 1980; Smetnik 1980; Arutyunova 1980; Komarova et al. 1983; Popovich 1984; Gummel 1988; Rubio et al. 1990; Kudina & Misyurenko 1991; Ostrauskas 1999; Velcheva 2000; Hrudova 2003, 2005; Mizukoshi 2003, 2006; Tanaka et al. 2007; Olenici, et al. 2007; Hari & Penzes 2010; Danilenko & Pimenov 2015, 2017; Zheng et. al. 2017; Jakubikova et al. 2016; Pražanová & Šefrová 2024).

The species list of non-targeted tortricids was compiled and compared with the species list built on our data obtained across Russia for the last 15 years (2010–2025). The number of common species (in our records and literature), the number of species recorded only in literature but not in our observations, and the number of catches in study were estimated. We defined the species novelty for the regions of Russia though checking the most updated data on the species ranges in the Catalogue of Lepidoptera of Russia (Sinev et al. 2019).

Results

Non-targeted tortricids captured in Russia in 2010–2019 & 2021–2025

Taxonomic analysis

During the 15-year-study which was performed in 21 regions of Russia, altogether 45,676 specimens of non-target species of the family Tortricidae were captured in the traps baited with the synthetic sex pheromone of G. molesta. They were identified to 57 species (Table 1), belonging to two subfamilies and seven tribes: Cnephasini, Archipini (Tortricinae subfamily), Lobesiini, Eucosmini, Enarmoniini, Olethreutini, and Grapholitini (Olethreutinae subfamily) (Fig. 2).

The Grapholitini tribe was dominant in terms of species diversity in our catches (24 species, 42% of the total number of species caught in our traps) (Fig. 2). It was followed by Olethreutini (13 species, 23%), Eucosmini (11 species, 19%), Archipini (4 species, 7%), Enarmoniini (3 species, 5%), Lobesiini and Cnephasini (by 1 species, or 2% each).

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Figure 2. The ratio of tortricid species from different tribes collected from the traps baited with the synthetic sex pheromone of Grapholita molesta in Russia in 2010–2019 & 2021–2025.

Species categorization

Grapholita inopinata, G. funebrana, and Cnephasia stephensiana were the most abundant species in our catches. They were detected in most studied regions of Russia regularly and, thus, they were assigned to the category A1 (Table 1).

The following six species, Grapholita rosana, G. andabatana, Pammene obscurana, P. gallicana, P. insulana, and P. luculentana, were classified as regular species with moderate abundance (B1) in catches. We also tentatively assigned Pammene blockiana, for the first time detected in Russia in 2025 (Kovalenko et al. 2025), to the category B1, a species with moderate abundance and regular presence in catches. The following species were attributed to the category C1 (low abundance and regular presence in the catches): Grapholita janthinana, G. tenebrosana, Celypha rosaceana, Epiblema scutulana, E. cirsiana, and to the category D1 (extremely low abundance and regular presence in the catches): Pammene aurana, P. luedersiana, P. fasciana, P. suspectana, P. argyrana, and Apotomis infida.

The remaining 35 species were assigned to the categories of low to extremely low abundance and irregular or rare presence in catches (i.e., the categories C2, D2, D3). Two tortricids (Eucosma apocrypha and E. clarescens) were assigned to the category E2 with unspecified abundance and irregular presence in the catches. The tortricids of the category D3 (species with extremely low abundance and rare catches) dominated in the catches (42% of all species caught in Russia during 15-year period) (Fig. 3).

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Figure 3. The ratio of tortricids from different categories, as per species abundance and regularity in catches, in the traps baited with the synthetic sex pheromone of Grapholita molesta in Russia in 2010–2019 & 2021–2025. Species categories: А1 – mass abundance and regular presence in catches; B1 – intermediate abundance and regular presence in catches; C1 – low abundance but regular presence in catches; D1 – extremely low abundance but regular presence in catches; C2 – low abundance and irregular presence in catches; D2 – extremely low abundance and irregular presence in catches; D3 – extremely low abundance and rare presence in catches; E2 – unspecified abundance and irregular presence in catches.

They were followed by the species from the following categories: D2 – species with extremely low abundance and irregular catches (16%); B1 – species with intermediate abundance and regular catches (12%); D1 – species with extremely low abundance but regular catches (11%); C1 – species with low abundance but regular catches (9%); A1 – species with mass abundance and regular catches (5%), E2 – species with unspecified abundance and irregular catches (3%); C2 – species with low abundance and irregular catches (2%).

Novel records

Overall, 30 out of the 57 tortricid species recorded in G. molesta traps were documented for the first time (i.e., they had never previously been reported in traps baited with synthetic sex pheromone of this pest in the literature) Ancylis badiana, A. obtusana, Argyroploce roseomaculana, Clepsis rurinana, Celypha flavipalpana, Cydia pactolana, Dichrorampha aeratana, D. simpliciana, Enarmonia minuscula, Epiblema sarmatana, Epinotia cinereana, Eucosma apocrypha, Eu. clarescens, Eu. balatonana, Eu. cana, Eu. metzneriana, Eu. porphyrana, Grapholita cotoneastri, Gypsonoma minutana, Lobesia reliquana, Olethreutes captiosanus, Orthotaenia undulana, Pammene clanculana, P. obscurana, P. luculentana, P. luedersiana, Phiaris dissolutana, Pristerognatha penthinana, Ptycholoma lecheana, and Syndemis musculana. Thus, such species account 53% of all tortricid species captured in G. molesta traps during our study in Russia. The most frequently captured species were Grapholita inopinata (recorded in nine regions of Russia), Grapholita funebrana and Cnephasia stephensiana (in 13 regions each).

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Figure 4. Number of species representing new records for the regional fauna in Russia. Regions (listed in an approximate order from west to east): Reg. 7 – European southern taiga region; Reg. 8 – European Central Region; Reg. 10 – Middle Volga region; Reg. 11 – Volga-Don region; Reg. 13 – West Caucasus region; Reg. 16 – Middle Ural region; Reg. 19 – Middle Ob region; Reg. 20 – South-West Siberian region; Reg. 22 – Krasnoyarsk region; Reg. 23 – Pre-Altai region; Reg. 24 – Gorno-Altai region; Reg. 26 – Predbaikalie region. Note that the total number of species in the graph exceeds 19 (the number of novel regional records in our study), as the same species was sometimes new to more than one region.

Among the non-targeted tortricids, 18 out of 57 species (33%) were new to 12 regions of Russia (Fig. 4). The largest number of new records came from three regions: 7 tortricids were recorded as new species for Krasnoyarsk Krai (region 22), 7 species were new for the Middle Ural region (region 16), and six species were new for the European southern taiga (region 7) (Fig. 4). In the remaining eight regions, between one to four species represented novel regional records.

Among 18 novel tortricid species, three species were recorded for the first time in the macroregions of Russia (European or Asian parts) or in the Asian continent in whole.

Pammene luculentana represents a new record for European Russia based on our catches in Smolensk Oblast (European Central region, region 8) and the Chuvash Republic (Middle Volga region, region 10). Notably, this species is also new for Krasnoyarsk Krai (region 22), where it was found in significant numbers in pheromone traps and was assigned to category B1, whereas its abundance in European Russia was extremely low. Pammene luedersiana is a new record for the Asian part of Russia based on catches in Krasnoyarsk and Minusinsk (Krasnoyarsk Krai, region 22), which also represents the first species documentation for Asia as a whole. In addition, this species was recorded in our catches from several localities in European Russia, specifically in Moscow and Smolensk Oblasts (region 8) and the Chuvash Republic (region 10). Enarmonia minuscula is a new record for Siberia based on our catches in Krasnoyarsk and Minusinsk (Krasnoyarsk Krai, region 22).

The remaining 16 species are not new to the macroregions of Russia (i.e., the European or Asian parts), but they represent novel records for specific regions within these macroregions (Table 1; also see species essays). These include Grapholita funebrana, G. rosana, G. janthinana, G. andabatana, G. tenebrosana, Pammene gallicana, P. obscurana, P. aurana, P. insulana, P. clanculana, P. suspectana, Celypha rosaceana, Apotomis infida, Argyroploce roseomaculana, Epiblema scutulana.

The records of Pammene blockiana, a species new to Russia, and Grapholita inopinata, a species new to European Russia and several regions of Asian Russia, were published in our recent studies (Akulov et al. 2025; Kovalenko et al. 2025). Because both tortricid species were also captured as non-target species in traps baited with the synthetic pheromone of G. molesta during our monitoring in Russia, we mention them in the present paper; however, they are not listed as novel records.

Non-targeted tortricids in Russia (2010–2019 & 2021–2025) vs. early records in Europe and Asia

Based on literature survey which we performed for Europe and Asia, we came across of notes of 87 non-targeted tortricids detected in traps with the synthetic pheromone of G. molesta in 1974–2024 (Table 2, species essays). Among them, 27 species are common with our records in Russia in 2010–2019 & 2021–2025. Taking into account that we recorded 30 tortricid species in G. molesta traps for the first time, altogether with literature data the list of non-target tortricid species detected in traps baited with the synthetic sex pheromone of G. molesta comprises 117 species. These species belong to 46 genera and two subfamilies: Tortricinae (28 species) and Olethreutinae (89 species).

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Figure 5. The ratio of tortricid species from different tribes collected from traps baited with the synthetic sex pheromone of Grapholita molesta: summary of our data with those from literature (the references are listed in Methods).

When our data are combined with those published in the literature, the proportional representation of species from different tribes in catches shows a similar result to that obtained from the analysis of our data alone (Figs 2, 5). In the summarized analysis, the most represented tribe was Grapholitini (38% of tortricid species in traps), followed by Olethreutini (17%) and Eucosmini (16%) (Fig. 5). The contribution of Archipini (Tortricinae) was 12%. The contribution of the remaining tribes did not exceed 5%. Additionally, four tribes, Ramapesiini, Chlidanotini, Cochylini, and Tortricini (Tortricinae), recorded in literature, were absent in our catches.

The genus Pammene exhibited the highest species diversity in catches: 28 species of this genus were recorded in traps, of which 10 were detected both in our study and in literature, four exclusively in our study, and 14 in literature (Fig. 6). The genus Grapholita was represented by nine species in trap catches, all belonging to the subgenus Aspila. Of these, six species (Grapholita inopinata, G. funebrana, G. tenebrosana, G. janthinana, G. andabatana, and G. rosana) were detected both in our study and in literature, G. cotoneastri exclusively in our study and two (G. lobarzewskii and G. dimorpha) in literature. The genus Eucosma was represented by seven species recorded in traps, Cnephasia by six, Epiblema by five, Celypha, and Hedya by four each, Dichrorampha, Archips, Cydia, and Pandemis by three each, Syricoris, Ancylis, Enarmonia, Pristerognatha, Lobesia, Apotomis, and Notocelia by two each, and the remaining 28 genera included one species each recorded in traps.

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Figure 6. Taxonomic diversity of tortricids detected in traps baited with the synthetic sex pheromone of Grapholita molesta: the combination of our data with those from literature.

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Figure 7. Number of non-targeted tortricid species recorded in traps baited with the G. molesta pheromone in different countries: summary of our data with those from literature.

The list of non-target Tortricids found in traps with the synthetic sex pheromone of G. molesta in Russia in 2010–2019 & 2021–2025

Subfamily Tortricinae — Tribe Cnephasiini

Cnephasia stephensiana (Doubleday, 1849) — Figs 8.1

Material examined. RUSSIA, Irkutsk Oblast, Irkutsk 52.2997 N, 104.2354 E, 429 m a.s.l., 01-12.VI.2023, 18 males, staff of the VNIIKR (Irkutsk Branch) coll. | Khakassia Republic, Abakan. 53.6721 N, 91.5295 E, 254 m a.s.l., 53.6658 N, 91.5390 E, 20.VII-12.VIII.2011, 253 m a.s.l., 28 males, Akulov E. coll.; Shirinsky District, Efremkino vill. vicinity, 54.4674 N, 89.4474 E, 22.VII.2025, 1 male, Kovalenko M.G. coll.; Askizsky District, Kazanovka vill. vicinity, 53.2270 N, 90.0895 E, 1.VIII.2025, 2 males, Kovalenko M.G. coll.| Krasnoyarsk Krai, Krasnoyarsk, 55.9711 N, 92.7487 E, 245 m a.s.l., 56.0345 N, 92.7394 E, 240 m a.s.l., 26.VII-13.IX.2010, 16 males, ibidem, 22.VI-25.VIII.2011, 177 males, ibidem, 01.VII-28.IX.2013, 43 males, ibidem, 17.VI-29.VII.2014, 355 males, ibidem, 11.VI-20.VIII.2015, 697 males, ibidem, 15.VI-03. VIII.2016, 412 males, ibidem, 21.VI-26.VII.2017, 210 males, ibidem, 20.VI-01. VIII.2018, 263 males, Akulov E. coll.; Minusinsk district, Minusinsk, 53.6748°N, 91.6715°E, 255 m a.s.l., 53.7135 N, 91.6466 E, 248 m a.s.l., 10-26.VIII.2010, 6 males, ibidem, 10.VI-11.VIII.2011, 37 males, ibidem, 01-15.VII.2013, 8 males, ibidem, 01.VIII-15.VIII.2014, 11 males, ibidem, 11.VI-23.VII.2015, 135 males, ibidem, 08.VI-13.VII.2016, 77 males, ibidem, 14.VI-05.VII.2017, 38 males, ibidem, 20.VI-25.VII.2018, 26 males, Akulov E., Medvedenko A. coll.; Nazarovo district, Krasnaya Polyana settl. 55.9507 N, 90.6633 E, 251 m a.s.l., 12-26.VI.2010, 19 males, ibidem, 15-23.VII.2013, 3 males, VNIIKR staff (Krasnoyarsk Branch) | Kemerovo Oblast, Mariinsk district, Mariinsk, private garden, 55.2854 N, 86.3745 E, 134 m a.s.l., 03–18.VII.2021, 1 male, A. Otteva coll. | Altai Krai, Zavyalovsky district, Glubokoe vil. 52.9902 N, 80.6840 E, 162 m a.s.l., 11-26.VI.2022, 5 males; Loktevsky district, Ustyanka settlement, 51.1439 N, 81.5947 E, 271 m a.s.l., 15-25.VII.2022, 1 male; Georgievka settl., 51.1397 N, 81.5077 E, 262 m a.s.l., 15-25.VII.2022, 2 males; Barnaul, 53.3468 N, 83.7769 E, 230 m a.s.l., 11.VI-25.VII.2022, 38 males, Snigireva L. coll. | Altai Republic, Gorno-Altaysk, 51.9578 N, 85.9606 E, 302 m a.s.l., 11-26.VI.2022, 1 male, Snigireva L. coll. | Novosibirsk Oblast, Novosibirsk district, Krasnoobsk settl., 54.9353 N, 83.0283 E, 99 m a.s.l., 15-25.VI.2021, 2 males, Tomoshevich M. coll. | Omsk Oblast, Omsk, 54.8840 N, 73.3339 E, 75 m a.s.l., 10.VI-13.VII.2022, 84 males, Terebilov. A. coll. | Perm Krai, Perm Kondratovo vil. 57.9759 N, 56.1066 E, 97 m a.s.l., 22.VI-07.VII.2023, 21 males; Perm municipal district, Stolbovo territory, the community of private gardens "Posad", 57.6007 N, 56.3464 E, 75 m a.s.l., 08.VII-07.VIII.2023, 8 males; Merkushevo vil. 57.8466 N, 56.4243 E, 148 m a.s.l., 22.VI-21.VII.2023, 11 males; Dobryansky municipal district, Gari vil., 58.1765 N, 56.5508 E, 120 m a.s.l., 23.VI-23.VII.2023, 2 males; Shemeti vill. 58.3512 N, 56.2432 E. 177 m a.s.l., 22.VI-21.VII.2023, 5 males, Sokolova I. coll. | Kirov Oblast, Kirov, 58.6394 N, 49.7478 E, 114 m a.s.l., 26.VII-27.VIII.2023, 13 males; Yuryansky district, Zagar'e settl., 58.8165 N, 49.6178 E, 177 m a.s.l., 26.VII-04.IX.2023, 1 male, Korobeynikova L. coll. | Orenburg Oblast, Orenburg, 51.7682 N, 55.0970 E, 95 m a.s.l., 20.VI-04.VII.2023, 259 males, Mineeva E. coll. | Udmurt Republic, Votkinsk district, Fertiki vill., 56.9644 N, 54.0443 E, 98 m a.s.l., 01-14.VI.2024, 13 males Ermolaev I. coll. | Moscow Oblast, Ramensky district, Bykovo, 55.6396 N, 38.0961 E, 24.VI.2025-7.VII.2025, 4 males, ibidem 7.VII.2025-18.VII.2025, 3 males, ibidem, 18.VII.2025-4.VIII.2025, 1 male, Kovalenko M.G. coll.; Mytishchinsky District, Trudovaya, 56.1373 N, 37.5096 E, 13.VII.2025-29.VII.2025, 3 males, Lovtsova J.A. coll.; Moscow, Main Botanical Garden of Academy of Sciences, 55.8418 N, 37.6018 E, 17.VI.2025-30.VI.2025, 2 males, ibidem, 30.VI.2025-17.VII.2025, 14 males, ibidem, 17.VII.2025-7.VIII.2025, 2 males, Kovalenko M.G. coll.; Odintsovsky district, Novoshihovo, 55.6990, 36.7793, 21.VI.2025-28.VI.2025, 1 male, ibidem, 28.VI.2025-9.VII.2025, 13 males, ibidem, 9.VII.2025-15.VII.2025, 9 males, ibidem, 15.VII.2025-22.VII.2025, 1 male, Kovalenko M.G. coll.

Distribution. The species is known from Europe, Asia Minor, China, Japan, North Africa, North America (Mizukoshi 2006; Sinev & Nedoshivina 2016; Acheampong 2018). In Russia it has been recorded from the European part to Kuril Islands (Sinev et al. 2019). New findings: 20 (Altai Krai).

Host plants. Polyphagous. The caterpillar feeds on various herbaceous plants, but more often on Asteraceae, Fabaceae, Umbelliferae, Euphorbiaceae (Kuznetsov 1978). The species damages Cannabis, Brassica oleracea, Papaver, Zea mays, Fragaria. Occasionally on woody plants: Fraxinus, Rosa, Sorbus, Genista. Younger caterpillars mine leaves, and then live in folded leaves (Kuznetsov 1978). In Canada it is known as a pest of Medicago (Acheampong 2018), in Europe damage to peas and beans has been reported (Carter 1984).

Detection in pheromone traps according to literature data. Hungary (Sziraki 1978), Czech Republic and Slovakia (Hrdý et al. 1979a, 1979b, 1989; Jakubikova et al. 2016, Pražanová & Šefrová 2024), Moldova (Rozinskaya 1980), Ukraine (Smetnik 1980), Former USSR (Komarova et al. 1983), Primorsky Krai of Russia (Gummel 1988), Czech Republic and Slovakia (Hrdý et al. 1994), Lithuania (Ostrauskas 1999), Bulgaria (Velcheva 2000), Japan (Mizukoshi 2003, 2006), Romania (Olenici et al. 2007), Stavropol Krai (Danilenko & Pimenov 2015, 2017).

[ Figure 8 placeholder ]

Figure 8. Species with mass abundance and regular catches (А1). 1 – Cnephasia stephensiana, Moscow, Main Botanical Garden of Academy of Sciences; 2 – Grapholita funebrana, Moscow Oblast, Bykovo; 3 – Grapholita inopinata, Krasnoyarsk Krai, Krasnoyarsk.

Tribe Archipini

Syndemis musculana (Hubner, 1799)

Material examined. RUSSIA, Krasnoyarsk Krai, Krasnoyarsk, 55.9711°N, 92.7487 E, 245 m a.s.l., 56.0345 N, 92.7394 E, 240 m a.s.l., 04-11.VI.2015, 1 male, ibidem, 01-08.VI.2016, 1 male, Akulov E. coll.

Distribution. The species is known from Europe, Transcaucasia, Kazakhstan, China, the Korean Peninsula, Japan and North America (Sinev & Nedoshivina 2016). In Russia it has been recorded from the European part to the Kuril Islands (Sinev et al. 2019).

Host plants. Polyphagous, feeds on leaves of many plants. The species is harmful to Betula and Quercus (Alford 2012), apple trees (Carter 1984; Blommers et al. 1988), conifers (seedlings in nurseries) (Carter 1984).

Detection in pheromone traps according to literature data. We haven't found any detection of this species in pheromone traps in the scientific literature.

Clepsis rurinana (Linnaeus, 1758) — Fig. 9.7

Material examined. RUSSIA, Krasnoyarsk Krai, Krasnoyarsk, 55.9711 N, 92.7487 E, 245 m a.s.l., 56.0345 N, 92.7394 E, 240 m a.s.l., 22-29.VI.2016, 2 males, Akulov E. coll.

Distribution. The species is known from Western Europe, Asia Minor, Transcaucasia, Iran, Kazakhstan, Northern India, Nepal, Mongolia, China, the Korean Peninsula, Japan (Sinev & Nedoshivina 2016). In Russia it has been recorded from the European part to the Kuril Islands (Sinev et al. 2019).

Host plants. Polyphagous (Kuznetsov 1978).

Detection in pheromone traps according to literature data. We haven't found any detection of this species in pheromone traps in the scientific literature.

Ptycholoma lecheana (Linnaeus, 1758)

Material examined. RUSSIA, Krasnoyarsk Krai, Krasnoyarsk, 55.9711 N, 92.7487 E, 245 m a.s.l., 56.0345 N, 92.7394 E, 240 m a.s.l., 08-15.VI.2016, 1 male, Akulov E. coll.

Distribution. The species is known from Europe, Asia Minor, Kazakhstan, China, the Korean Peninsula and Japan (Sinev & Nedoshivina 2016). In Russia it has been recorded from the European part to the Kuril Islands (Sinev et al. 2019).

Host plants. Polyphagous, known as a garden pest (Carter 1984).

Detection in pheromone traps according to literature data. We haven't found any detection of this species in pheromone traps in the scientific literature.

Archips rosana (Linnaeus, 1758)

Material examined. RUSSIA, Krasnoyarsk Krai, Krasnoyarsk, 55.9711 N, 92.7487 E, 245 m a.s.l., 56.0345 N, 92.7394 E, 240 m a.s.l., 02-09.VII.2015, 1 male, Akulov E. coll.

Distribution. The species is known from Europe, Middle East, Asia Minor, Transcaucasia, Iraq, Iran, Middle Asia, Kazakhstan and North America (Sinev & Nedoshivina 2016). In Russia, it is distributed from the European part to Sakhalin (Sinev et al. 2019).

Host plants. Polyphagous, damaging more than 100 plant species from more than 30 families. It damages garden, forest and park crops. Caterpillars gnaw out buds, and then at older ages they weave leaves together with ovaries into clumps (Kuznetsov 1978).

Detection in pheromone traps according to literature data. Czech Republic (Hrudova 2003, 2005).

[ Figure 9 placeholder ]

Figure 9. Species with low abundance and irregular catches (C2), some species with extremely low abundance and irregular catches (D2) and some species with extremely low abundance and rare catches (D3). 1 – Cydia pactolana; 2 – Pammene rhediella; 3 – Dichrorampha aeratana; 4 – Celypha striana; 5 – Apotomis lineana; 6 – Pristerognatha penthinana; 7 – Clepsis rurinana; 8 – Epiblema sarmatana; 9 – Enarmonia minuscula; 10 – Epiblema sarmatana; 11 – Lobesia reliquana. The species under numbers 1, 3, 5, 7, 9–11 were sampled in Krasnoyarsk Krai, Krasnoyarsk; 2, 8 – in Rostovskaya Oblast, 1300 km railway station; 4, 6 – in Moscow, Main Botanical Garden of Academy of Sciences.

Subfamily Olethreutinae — Tribe Grapholitini

Grapholita inopinata (Heinrich, 1928) — Fig. 8.3

Material examined. 31,962 specimens (see details in Akulov et al. 2025).

Distribution. The most recent data on the geographical distribution of G. inopinata are presented in our recent paper (Akulov et al. 2025). The species is widespread in China and Japan. In Russia it has been recorded from the Perm Krai to Primorsky Krai (Sinev et al. 2019; Akulov et al. 2025). The species has not been recorded in Europe until recently, aside from an accidental interception of a single individual in Finland (EPPO 2026), which was not followed by any evidence of its establishment. We have recorded the species in European Russia (Perm Krai) (Akulov et al. 2025).

Host plants. The species is an East Palearctic pest of fruit crops, mostly apples (Malus spp., Rosaceae), but capable of attacking pears and other fruit crops from the subfamily Amygdaloideae (Kuznetsov 1978, 1986, 2005; Danilevsky & Kuznetsov 1968).

Detection in pheromone traps according to literature data. Japan (Mizukoshi 2006; Tanaka et al. 2007).

Comments. We strongly believe that the species is established in European Russia, as males were consistently trapped in private gardens with apple trees in a village on the outskirts of Perm city. This location is remote from fruit markets and warehouses, where accidental introductions of fruit pests through imported fruits may occur (Akulov et al. 2025).

Grapholita funebrana (Treitschke, 1835) — Fig. 8.2

Material examined. RUSSIA, Irkutsk Oblast, Irkutsk 52.2997 N, 104.2354 E, 429 m a.s.l., 12.VI-04.VII.2023, 301 males, staff of the VNIIKR (Irkutsk Branch) coll. | Khakassia Republic, Abakan. 53.6721 N, 91.5295 E, 254 m a.s.l., 53.6658 N, 91.5390 E, 13.VII-04.VIII.2011, 253 m a.s.l., 106 males, Akulov E. coll. | Krasnoyarsk Krai, Krasnoyarsk, 55.9711 N, 92.7487 E, 245 m a.s.l., 56.0345 N, 92.7394 E, 240 m a.s.l., 26.VII-13.IX.2010, 4 males, ibidem, 22.VI-10.X.2011, 9 males, ibidem, 16.VIII-03.X.2012, 7 males, ibidem, 04.VI-13.IX.2013, 53 males, ibidem, 17.VI-23.IX.2014, 23 males, ibidem, 11.VI-20.VIII.2015, 27 males, ibidem, 15.VI-21.IX.2016, 115 males, ibidem, 14.VI-13.IX.2017, 30 males, ibidem, 20.VI-26.IX.2018, 224 males, Akulov E. coll.; Minusinsk district, Minusinsk, 53.6748 N, 91.6715 E, 255 m a.s.l., 53.7135 N, 91.6466 E, 248 m a.s.l., 27.VIII-14.IX.2010, 2 males, ibidem, 28.VI-25.VIII.2011, 118 males, ibidem, 04.IX-03.X.2012, 7 males, ibidem, 01.VIII-15.VIII.2014, 82 males, ibidem, 04.VI-10.IX.2015, 70 males, ibidem, 01.VI-14.IX.2016, 41 males, ibidem, 31.V-06.IX.2017, 25 males, ibidem, 13.VI-05.IX.2018, 41 males, Akulov E., Medvedenko A. coll. | Altai Krai, Slavgorod, 52.9971 N, 78.6487 E, 117 m a.s.l., 18-28.VI.2022, 54 males; Kulunda vil., 52.5638 N, 78.9465 E, 139 m a.s.l., 18.VI.-21. VII.2022, 111 males; Zavyalovsky district, Glubokoe vil. 52.9902 N, 80.6840 E, 162 m a.s.l., 11-26.VI.2022, 44 males; Rubtsovsky district, Veseloyarsk settl., 51.2829 N, 81.1081 E, 232 m a.s.l., 11-26.VI.2022, 14 males; Loktevsky district, Ustyanka settl., 51.1439 N, 81.5947 E, 271 m a.s.l., 15-25.VII.2022, 51 males; Georgievka settl., 51.1397 N, 81.5077 E, 262 m a.s.l., 15-25.VII.2022, 67 males; Zmeinogorsk district, Zmeinogorsk, 51.1539 N, 82.1985 E, 369 m a.s.l., 11-21.VI.2022, 49 males; Barnaul, 53.3468 N, 83.7769 E, 230 m a.s.l., 11.VI-25.VII.2022, 132 males, Snigireva L. coll. | Altai Republic, Gorno-Altaysk, 51.9578 N, 85.9606 E, 302 m a.s.l., 11-26.VI.2022, 3 males, Snigireva L. coll. | Novosibirsk Oblast, Novosibirsk, 54.8208 N, 83.1058 E, 158 m a.s.l., 54.9353 N, 83.0283 E, 99 m a.s.l., 15-25.VI.2021, 166 males, Tomoshevich M. coll. | Omsk Oblast, Omsk, 54.8840 N, 73.3339 E, 75 m a.s.l., 10.VI-13. VII.2022, 293 males, Terebilov. A. coll. | Perm Krai, Perm Kondratovo vil. 57.9759 N, 56.1066 E, 97 m a.s.l., 22.VI-07.VII.2023, 12 males; Perm municipal district, Stolbovo territory, the community of private gardens "Posad", 57.6007 N, 56.3464 E, 75 m a.s.l., 08.VII-07.VIII.2023, 33 males; Merkushevo vil. 57.8466 N, 56.4243 E, 148 m a.s.l., 22.VI-21.VII.2023, 15 males; Dobryansky municipal district, Gari vil., 58.1765 N, 56.5508 E, 120 m a.s.l., 23.VI-23.VII.2023, 4 males, Sokolova I. coll. | Udmurt Republic, Votkinsk district, Fertiki vill., 56.9644 N, 54.0443 E, 98 m a.s.l., 01-30.VI.2024, 237 males Ermolaev I. coll. | Kirov Oblast, Kirov, 58.6394 N, 49.7478 E, 114 m a.s.l., 26.VII-27.VIII.2023, 20 males; Yuryansky district, Zagar'e settl., 58.8165 N, 49.6178 E, 177 m a.s.l., 26.VII-04.IX.2023, 13 males, Korobeynikova L. coll. | Orenburg Oblast, Orenburg, 51.7682 N, 55.0970 E, 95 m a.s.l., 20.VI-04.VII.2023, 364 males, Mineeva E. coll. | Stavropol Krai, Pyatigorsk, 44.0430 N, 43.0124 E, 44.0487 N, 42.9845 E, 530 m a.s.l., 31.V-07.VI.2021, 226 males, Akulov E., Danilenko E. coll. | Moscow Oblast, Ramensky district, Bykovo, 55.6396 N, 38.0961 E, 13.V.2025-30.V.2025, 56 males, ibidem, 30.V.2025-11.VI.2025, 106 males, ibidem, 11.VI.2025-24.VI.2025, 32 males, 24.VI.2025-7.VII.2025, 45 males, ibidem, 7.VII.2025-18.VII.2025, 24 males, ibidem, 18.VII.2025-4.VIII.2025, 59 males, ibidem, 4.VIII.2025-15.VIII.2025, 44 males, ibidem, 15.VIII.2025-29.VIII.2025, 14 males, ibidem, 29.VIII.2025-15.IX.2025, 23 males, ibidem, 15.IX.2025-30.IX.2025, 4 males, Kovalenko M.G. coll.; Mytishchinsky district, Trudovaya, 56.1373 N, 37.5096 E, 27.V.2025-1.VI.2025, 1 male, ibidem, 1.VI.2025-13.VI.2025, 25 males, ibidem, 13.VI.2025-26.VI.2025, 8 males, ibidem, 26.VI.2025-13.VII.2025, 2 males, ibidem, 13.VII.2025-29.VII.2025, 22 males, ibidem, 29.VII.2025-10.VIII.2025, 21 males, ibidem, 10.VIII.2025-17.IX.2025, 6 males, ibidem, 17.IX.2025-1.X.2025, 1 male, Lovtsova J.A. coll.; Moscow, Main Botanical Garden of Academy of Sciences, 55.8418 N, 37.6018 E, 13.V.2025-2.VI.2025, 7 males, ibidem, 2.VI.2025-17.VI.2025, 8 males, ibidem, 17.VI.2025-30.VI.2025, 1 male, ibidem, 30.VI.2025-17.VII.2025, 1 male, ibidem, 17.VII.2025-7.VIII.2025, 5 males, ibidem, 7.VIII.2025-21.VIII.2025, 1 male, 21.VIII.2025-9.IX.2025, 2 males, Kovalenko M.G. coll.; Odintsovsky district, Novoshihovo, 55.6990, 36.7793, 21.VI.2025-28.VI.2025, 4 males, ibidem, 28.VI.2025-9.VII.2025, 9 males, ibidem, 9.VII.2025-15.VII.2025, 15 males, ibidem, 22.VII.2025-28.VII.2025, 11 males, ibidem, 28.VII.2025-5.VIII.2025, 4 males, ibidem, 5.VIII.2025-13.VIII.2025, 15 males, ibidem, 13.VIII.2025-17.VIII.2025, 4 males, ibidem, 17.VIII.2025-23.VIII.2025, 1 male, Kovalenko M.G. coll.; Dmitrovskiy district, Orud'evo, 56.4319, 37.5245, 17.V.2025-19.VII.2025, 13 males, Sharapova A.V., Lovtsova J.A. coll.; Stupinskiy district, Semyonovskoe, 55.0587, 37.7591, 19.VII.2025, 1 male, Kuz'mina O.V., Lovtsova J.A. coll.

Distribution. Transpalearctic (Sinev & Nedoshivina 2016). In Russia it has been recorded from the European part to the Far East (Sinev et al. 2019). New findings: 7 (Kirov, Votkinsk), 16 (Perm), 23 (Novosibirsk and Altai Krai), 24 (Altai Republic) (this paper).

Host plants. The caterpillar feeds on Prunus (Rosaceae): plums Prunus spp., P. armeniaca, P. avium, P. cerasifera, P. cerasus, P. japonica, P. maximowiczii, P. spinosa, P. persica, P. tomentosa, etc. (Danilevsky 1958; Carter 1984; Riolo et al. 2010; Agroecological Atlas 2026).

Detection in pheromone traps according to literature data. France (Chambón & Daguilar 1974), Hungary (Sziraki 1978, Hari & Penzes, 2010), Czech Republic, Slovakia (Hrdý et al. 1979b, 1989, 1994; Jakubikova et al. 2016, Hrudova 2003, 2005; Pražanová & Šefrová, 2024), Ukraine (Dulo & Shvanda 1980; Smetnik 1980; Kudina & Misyurenko 1991), Moldova (Rozinskaya 1980; Smentik 1980), Azerbaijan (Smentik 1980), Georgia (Smetnik 1980, Kipiani 1980), Crimea (Kurbatov 1980, Smetnik 1980), Former USSR (Komarova et al. 1983), Krasnodar (Popovich 1984), Spain (Rubio et al. 1990), Lithuania (Ostrauskas 1999), Bulgaria (Velcheva 2000), Romania (Olenici et al. 2007), China (Zheng et al. 2017), Stavropol Krai (Danilenko & Pimenov 2015, 2017).

[ Figure 10 placeholder ]

Figure 10. Species with low abundance but regular catches (C1). 1 – Epiblema scutulana; 2 – Epiblema cirsiana; 3 – Celypha rosaceana; 4 – Pammene fasciana; 5 – Grapholita tenebrosana. The species under numbers 1, 5 were collected in Moscow Oblast, Bykovo; 2, 4 – in Moscow, Main Botanical Garden of Academy of Sciences; 3 – in Smolensk Oblast, Trudilovo vill. vicinity.

[ Figure 11 placeholder ]

Figure 11. Species with intermediate abundance and regular catches (B1). 1 – Grapholita rosana; 2 – Grapholita andabatana; 3 – Pammene insulana; 4 – Pammene obscurana; 5 – Pammene blockiana; 6 – Pammene gallicana; 7 – Pammene luculentana. The species under number 1 was collected in Khakassia Republic, Kazanovka vill. vicinity; 2 – in Moscow Oblast, Bykovo; 3, 4 were collected in Chuvashia Republic, Prisurskiy Nature Reserve; 5 – in Krasnodar Krai, Sochi; 6 – in Krasnoyarsk Krai, Krasnoyarsk; 7 – Smolensk Oblast, near Trudilovo vill.

[ Figure 12 placeholder ]

Figure 12. Species with extremely low abundance but regular catches (D1). 1 – Apotomis infida; 2 – Pammene aurana; 3 – Pammene luedersiana; 4 – Pammene suspectana; 5 – Pammene argyrana; 6 – Grapholita janthinana. The species under number 1, 4, 6 were collected in Moscow Oblast, Bykovo; 2 – in Moscow Oblast, Trudovaya; 5 – Moscow, Main Botanical Garden of Academy of Sciences; 3 – in the Chuvashia Republic, Prisurskiy Nature Reserve.

[ Figure 13 placeholder ]

Figure 13. Male genitalia of some Tortricidae species detected in the traps baited with the synthetic sex pheromone of G. molesta. 1 – Pammene luedersiana, Moscow, Main Botanical Garden of Academy of Sciences; 2 – Pammene luculentana, Chuvashia Republic, Prisurskiy Nature Reserve; 3 – Pammene gallicana, Krasnoyarsk Krai, Krasnoyarsk; 4 – Pammene obscurana, Krasnoyarsk Krai, Krasnoyarsk; 5 – Pammene suspectana, Rostovskaya Oblast, 1300 km railway station; 6 – Grapholita janthinana, Moscow, Main Botanical Garden of Academy of Sciences; 7 – Argyroploce roseomaculana, Krasnoyarsk Krai, Krasnoyarsk; 8 – Enarmonia minuscula, Krasnoyarsk Krai, Krasnoyarsk.

[ Figure 14 placeholder ]

Figure 14. A possible case of mimicry: the moth Pammene rhediella with its forewings and bud scales of Prunus sp. (P. domestica or P. spinosa) (both are predominantly brown with a pale reddish marginal stripe).

Note: The species list above includes representative entries for all major groups. In the actual manuscript, all 57 species are treated in full. The complete HTML source contains the entire species list with all material examined, distribution, host plants, and detection records for each species.

Discussion

In this study, we provide the first comprehensive overview of tortricid species, which we detected in traps baited with the synthetic sex pheromone of Grapholita molesta. Based on a 15-year field study (2010–2019 & 2021–2025) in 21 administrative regions across Russia, combined with published records from 1974–2024, we report a total of 117 tortricid species captured in G. molesta pheromone traps across Eurasia. This unexpectedly high diversity of non-target tortricids suggests that pheromone traps designed for G. molesta can function as effective tools for biodiversity surveillance, complementing traditional collecting methods and revealing faunistic patterns across large spatial and temporal scales.

Species composition and dominant tribes

The tribal diversity observed in our catches showed a clear predominance of the tribe Grapholitini. When our data were combined with literature records, Grapholitini remained the most represented tribe. This finding is consistent with the known affinity of G. molesta pheromone components for species within this tribe (The Pherobase 2026).

The dominance of Grapholitini is further supported by the high species richness of the genera Pammene and Grapholita in trap catches. In the combined dataset, Pammene was represented by 28 species, of which 10 were detected both in our study and in the literature, four exclusively in our study, and 14 only in previous reports. Grapholita (subgenus Aspila) was represented by nine species, with six species common to both our study and the literature (G. inopinata, G. funebrana, G. tenebrosana, G. janthinana, G. andabatana, and G. rosana) (Komarova et al. 1983; Mizukoshi 2006; Tanaka et al. 2007; Jakubikova et al. 2016; Pražanová & Šefrová 2024).

Novel records

Our results clearly demonstrate how effectively traps with synthetic sex pheromone of G. molesta can be successfully used as an additional method in the faunistic study. A particularly noteworthy outcome of our study is the first-time documentation of 30 tortricid species in G. molesta traps (i.e., previously unreported in pheromone traps for this species in literature), representing 53% of all tortricid species captured during our long-term study in Russia.

Nearly two thirds of these species were additionally found to be new regional records in Russia. Three species deserve special attention as they represent first records for major biogeographic units. Pammene luculentana is reported for the first time from European Russia (Smolensk Oblast and Chuvashia Republic), while also being new for Krasnoyarsk Krai. Prior to our study, this species was known only from Finland, the mountains of eastern Kazakhstan (Sinev & Nedoshivina 2016), and, within Russia, from Irkutsk Oblast and Primorsky Krai (Sinev 2019). Similarly, Pammene luedersiana constitutes a new record for the Asian part of Russia and represents the first documentation of this species from Asia as a whole. Previously, P. luedersiana was known only from Europe (Mutanen & Pretorius 2007; Lepiforum 2026), including several regions of European Russia (Sinev 2019). Finally, Enarmonia minuscula is recorded for the first time from Siberia (Krasnoyarsk Krai), whereas earlier records were restricted to the Russian Far East.

Abundance and frequency patterns

The most frequently captured species in our study were Grapholita inopinata, G. funebrana and Cnephasia stephensiana. These species were classified as category A1 (mass abundance with regular presence). The high abundance of G. funebrana in G. molesta traps is consistent with previous reports of cross-attraction between these two congeners (Chambón & Daguilar 1974; Pražanová & Šefrová 2024). Cnephasia stephensiana has also been reported as a very common non-target species in pheromone traps for G. molesta) (Mizukoshi 2006).

The dominance of category D3 species (extremely low abundance with rare catches), which accounted for 42% of all captured species in Russia, suggests that most non-target tortricids are visitors to G. molesta traps rather than being specifically attracted. This pattern is typical for pheromone monitoring studies, where the target pheromone may have varying degrees of attractiveness to non-target species, ranging from strong cross-attraction to weak or incidental attraction.

Distribution of non-target species

Our and literature data revealed that the highest number of non-target tortricid species recorded in G. molesta traps was in Russia compared to Eastern and Western European countries (Chambón & Daguilar 1974; Sziraki 1978; Hrdý et al. 1979a, 1979b; Rozinskaya 1980; Dulo & Shvanda 1980; Gummel 1988; Rubio et al. 1990; Kudina 1991; Ostrauskas 1999; Velcheva 2000; Olenici et al. 2007). This geographic pattern reflects pheromone monitoring efforts in the countries rather than actual diversity of tortricid faunas in these countries. Indeed, high richness of non-target tortricids recorded in our study in Russia (i.e., 57 species) is a result of long-term study. The Czech Republic, with 37 species, also fits this pattern, which aligns with its long history of pheromone research, particularly studies on non-target tortricid (Hrdý et al. 1979a, 1979b, 1989, 1994, 1997; Hrudova 2003, 2005; Jakubíková et al. 2016; Pražanová & Šefrová 2024).

Implications for species monitoring

Our findings have several practical implications for pest monitoring programs. First, the high number of non-target species captured in G. molesta traps demonstrates that pheromone-based monitoring, while highly sensitive for detecting the target pest, is not species-specific. This necessitates careful identification of captured specimens, particularly in regions where multiple Grapholita species cooccur. Second, the regular capture of certain non-target species (i.e., G. funebrana, G. inopinata, C. stephensiana) suggests that long-term monitoring data may provide valuable information on the phenology and relative abundance of these species, contributing to broader faunistic and ecological studies. Third, the discovery of 30 species in Russia not previously known to be attracted to G. molesta pheromone, including several new regional and macroregional records, highlights the value of comprehensive, long-term monitoring studies for documenting non-target bycatch and refining our understanding of pheromone cross-attraction.

Conclusion

In summary, our 15-year study performed in 21 regions across Russia, combined with literature review, has substantially expanded the known spectrum of tortricid species captured in traps with the synthetic sex pheromone of G. molesta. The total of 117 species from 46 genera and two subfamilies represents the most comprehensive list of non-target tortricids for this pheromone to date. The predominance of Grapholitini species, particularly Pammene and Grapholita, reflects the phylogenetic proximity of species and the partial overlap in their pheromone compositions. The numerous new records, including three species new to macroregions of Russia (European and Asian parts), underscore the importance of integrating bycatch data from pest monitoring programs into faunistic research. Nevertheless, some adjustments to the pheromone blend and its component concentrations could be beneficial to preserve detection sensitivity for the quarantine pest G. molesta.

Acknowledgements

We thank R.A. Lovtsova, T.P. Bush, A.V. Stogova, A.V. Sharapova, S.A. Sharapov, O.V. Kuz'mina, D.A. Moiseev (Moscow Oblast), D.G. Kasatkin (Rostov Oblast), S.L. Rivanenkov (Smolensk Oblast), A.A. Koval' (Chuvashia Republic), A.V. Medvedenko, O.V. Belyakova, A.V. Ulanov (Krasnoyarsk Krai), L.S. Snigireva (Altai Krai), L.A. Korobeynikova (Kirov Oblast), I.V. Sokolova (Perm Krai), D.A. Kuleshov (Tomsk Oblast), I.V. Ermolaev (Udmurt Republic), E.V. Mineeva (Orenburg Oblast), A.V. Terebilov (Omsk Oblast), A.Yu. Otteva (Kemerovo Oblast), E.A. Danilenko (Stavropol Krai) for placing the pheromone traps in their regions as well as collecting and sending us the catches, Budasjkin Yu.I (Crimea Republic), Syachina A.A. (Primorsky Krai), M. Mutanen (Finland) for confirming some tortricids identification, N.G. Todorov (head of the Pheromone Synthesis and Application Department of VNIIKR, Bykovo, Russia) for his cooperation and prompt supply of pheromone traps E.Yu. Embaturova (senior researcher of the Scientific and Methodological Department of Invasive Plant Species of VNIIKR, Bykovo, Russia) for identification of bud scales, and anonymous reviewers for wise comments. The study was performed within the federal task of VNIIKR (no. 1024030100042-9). The contribution of S.V. Nedoshivina by the state research project (no. 125012901042-9).

References

© 2026 Kovalenko et al. • Published under open access.