New Guests in the Romanian Fauna and Pathways for the Introduction of Alien Bush-Crickets and Grasshoppers (Insecta: Orthoptera)
Abstract
1. Introduction
2. Materials and Methods
3. Results
3.1. Taxonomy


3.2. Genetic Data on Meconema meridionale
4. Discussion
4.1. Anthropogenic Pathways and Urban Habitats
4.2. Genetic Insights into the Eastern Colonization Margin
4.3. The Romanian Context and Regional Biogeography
4.4. Management and Biosecurity Recommendations
Author Contributions
Funding
Institutional Review Board Statement
Data Availability Statement
Acknowledgments
Conflicts of Interest
References
- Richardson, D.M.; Pyšek, P.; Rejmanek, M.; Barbour, M.G.; Panetta, F.D.; West, C.J. Naturalization and invasion of alien plants: Concepts and definitions. Divers. Distrib. 2000, 6, 93–107. [Google Scholar] [CrossRef]
- Pyšek, P.; Hulme, P.E.; Simberloff, D.; Bacher, S.; Blackburn, T.M.; Carlton, J.T.; Dawson, W.; Essl, F.; Foxcroft, L.C.; Genovesi, P.; et al. Scientists’ warning on invasive alien species. Biol. Rev. 2020, 95, 1511–1534. [Google Scholar] [CrossRef]
- Turbelin, A.; Diagne, C.; Hudgins, E.; Moodley, D.; Kourantidou, M.; Novoa, A.; Haubrock, P.; Bernery, C.; Gozlan, R.; Francis, R.; et al. Introduction pathways of economically costly invasive alien species. Biol. Invasions 2022, 24, 2061–2079. [Google Scholar] [CrossRef]
- Haubrock, P.; Turbelin, A.; Cuthbert, R.; Nóvoa, A.; Taylor, N.; Angulo, E.; Ballesteros-Mejia, L.; Bodey, T.; Capinha, C.; Diagne, C.; et al. Economic costs of invasive alien species across Europe. NeoBiota 2021, 67, 153–190. [Google Scholar] [CrossRef]
- Hulme, P.E. Trade, transport and trouble: Managing invasive species pathways in an era of globalization. J. Appl. Ecol. 2009, 46, 10–18. [Google Scholar] [CrossRef]
- Seebens, H.; Blackburn, T.M.; Dyer, E.E.; Genovesi, P.; Hulme, P.E.; Jeschke, J.M.; Pagad, S.; Pyšek, P.; Winter, M.; Arianoutsou, M.; et al. No saturation in the accumulation of alien species worldwide. Nat. Commun. 2017, 8, 14435. [Google Scholar] [CrossRef]
- Wang, S.; Li, W.; Zhang, J.; Luo, Z.; Li, Y. Alien range size, habitat breadth, origin location, and domestication of alien species matter to their impact risks. Integr. Zool. 2024, 20, 520–534. [Google Scholar] [CrossRef] [PubMed]
- Hernández-Triana, L.M.; Brugman, V.A.; Nikolova, N.I.; Ruiz-Arrondo, I.; Barrero, E.; Thorne, L.; de Marco, M.F.; Krüger, A.; Lumley, S.; Johnson, N.; et al. DNA Barcoding of British Mosquitoes (Diptera, Culicidae) to Support Species Identification, Discovery of Cryptic Genetic Diversity and Monitoring Invasive Species. ZooKeys 2019, 832, 57–76. [Google Scholar] [CrossRef] [PubMed]
- Milián-García, Y.; Young, R.; Madden, M.; Bullas-Appleton, E.; Hanner, R.H. Optimization and Validation of a Cost-Effective Protocol for Biosurveillance of Invasive Alien Species. Ecol. Evol. 2021, 11, 1999–2014. [Google Scholar] [CrossRef] [PubMed]
- Comtet, T.; Sandionigi, A.; Viard, F.; Casiraghi, M. DNA (Meta)Barcoding of Biological Invasions: A Powerful Tool to Elucidate Invasion Processes and Help Managing Aliens. Biol. Invasions 2015, 17, 905–922. [Google Scholar] [CrossRef]
- Garcia-Vazquez, E.; Ardura, A.; Planes, S. DNA mtCOI Barcodes for Maritime Biosecurity: A Proof of Concept in French Polynesia Ports. Front. Ecol. Evol. 2020, 8, 179. [Google Scholar] [CrossRef]
- Mahon, A.R.; Grey, E.K.; Jerde, C.L. Integrating Invasive Species Risk Assessment into Environmental DNA Metabarcoding Reference Libraries. Ecol. Appl. 2023, 33, e2730. [Google Scholar] [CrossRef]
- Barney, J.N.; Tekiela, D.R.; Dollete, E.S.; Tomasek, B.J. What is the “real” impact of invasive plant species? Front. Ecol. Environ. 2013, 11, 322–329. [Google Scholar] [CrossRef]
- Bradshaw, C.J.; Leroy, B.; Bellard, C.; Roiz, D.; Albert, C.; Fournier, A.; Barbet-Massin, M.; Salles, J.M.; Simard, F.; Courchamp, F. Massive yet grossly underestimated global costs of invasive insects. Nat. Commun. 2016, 7, 12986. [Google Scholar] [CrossRef] [PubMed]
- Diagne, C.; Leroy, B.; Gozlan, R.E.; Vaissière, A.-C.; Assailly, C.; Nuninger, L.; Roiz, D.; Jourdain, F.; Jarić, I.; Courchamp, F. InvaCost, a public database of the economic costs of biological invasions worldwide. Sci. Data 2020, 7, 277. [Google Scholar] [CrossRef] [PubMed]
- Essl, F.; Zuna-Kratky, T. The checklist of alien orthopterans (Orthoptera) and mantises (Mantodea) in Austria (2nd edition). BioInv. Rec. 2021, 10, 991–996. [Google Scholar] [CrossRef]
- Demetriou, J.; Radea, C.; Peyton, J.; Groom, Q.; Roques, A.; Rabitsch, W.; Seraphides, N.; Arianoutsou, M.; Roy, H.; Martinou, A. The Alien to Cyprus Entomofauna (ACE) database: A review of the current status of alien insects (Arthropoda, Insecta) including an updated species checklist, discussion on impacts and recommendations for informing management. NeoBiota 2023, 83, 11–42. [Google Scholar] [CrossRef]
- Musolin, D.; Kirichenko, N.; Karpun, N.; Aksenenko, E.; Golub, V.; Kerchev, I.; Mandelshtam, M.; Vasaitis, R.; Volkovitsh, M.; Zhuravleva, E.; et al. Invasive Insect Pests of Forests and Urban Trees in Russia: Origin, Pathways, Damage, and Management. Forests 2022, 13, 521. [Google Scholar] [CrossRef]
- Drăghici, A.-C.; Pintilioaie, A.-M.; Murariu, D.; Manci, C.-O.; Ruzzier, E. New additions and further records of non-native Coleoptera in Romania. Zootaxa 2025, 5575, 409–428. [Google Scholar] [CrossRef]
- Renault, D.; Laparie, M.; McCauley, S.; Bonte, D. Environmental Adaptations, Ecological Filtering, and Dispersal Central to Insect Invasions. Annu. Rev. Entomol. 2018, 63, 345–368. [Google Scholar] [CrossRef]
- Krištín, A.; Capka, J.; Zlámal, N.; Demeš, P. First records of the alien bush-crickets Rhacocleis annulata Fieber, 1853 and Yersinella raymondii (Yersin, 1860) (Orthoptera) in Slovakia. BioInv. Rec. 2022, 11, 383–389. [Google Scholar] [CrossRef]
- Kulessa, A.K.; Kouba, A.; Renault, D.; Soto, I.; Haubrock, P.J. Assessing non-native invasive Orthoptera in Europe. Res. Sq. 2023. [Google Scholar] [CrossRef]
- Jaskuła, R.; Sulikowska-Drozd, A.; Jabłońska, A.; Banaś, K.; Rewicz, T. Undesirable immigrants: Hobbyist vivaria as a potential source of alien invertebrate species. PeerJ 2019, 7, e7617. [Google Scholar] [CrossRef]
- Cigliano, M.M.; Braun, H.; Eades, D.C.; Otte, D. Orthoptera Species File. Available online: http://orthoptera.speciesfile.org/ (accessed on 30 August 2025).
- Capinera, J.L.; Scott, R.D.; Walker, T.J. Field Guide to Grasshoppers, Katydids and Crickets of the United States; Cornell University Press: Ithaca, NY, USA, 2004; pp. 1–280. [Google Scholar]
- Hochkirch, A.; Massa, B.; Zuna-Kratky, T.; Krištín, A.; Puskás, G.; Ivković, S.; Presa, J.J.; Skejo, J.; Monnerat, C.; Szövényi, G.; et al. Meconema meridionale. The IUCN Red List of Threatened Species 2016: e.T68427413A74540416. Available online: https://www.iucnredlist.org/species/68427413/74540416 (accessed on 12 October 2025).
- Tilmans, J.; Odé, B.; Willemse, L. Rhacocleis andikithirensis a new bush-cricket from Greece (Orthoptera: Tettigoniidae: Tettigoniinae). J. Orthoptera Res. 2016, 25, 25–38. [Google Scholar] [CrossRef]
- Willemse, F.; Willemse, L. Two new cryptic aberrant species of Rhacocleis Fieber 1853, living under stones in the mountains of Greece (Orthoptera; Tettigoniidae). J. Orthoptera Res. 2005, 14, 161–172. [Google Scholar] [CrossRef]
- Hochkirch, A.; Massa, B.; Monnerat, C.; Skejo, J.; Gomboc, S.; Willemse, L.P.M.; Rutschmann, F.; Chobanov, D.P.; Kleukers, R.; Krištín, A.; et al. Yersinella raymondii. The IUCN Red List of Threatened Species: e.T68486449A74625126. Available online: https://www.iucnredlist.org/species/68486449/74625126 (accessed on 12 October 2025).
- Zuna-Kratky, T. Eingeschleppte, nicht dauerhaft etablierte Arten. In Die Heuschrecken Österreichs; Zuna-Kratky, T., Landmann, A., Illich, I., Zechner, L., Essl, F., Lechner, K., Ortner, A., Weißmair, W., Wöss, G., Eds.; Denisia: Linz, Austria, 2017; Volume 39, pp. 816–818. [Google Scholar]
- Heller, K.G.; Mosny, K. Fund einer Kleinen Strauchschrecke Yersinella raymondii (Yersin, 1860) (Orthoptera, Tettigoniidae) im Rhein-Neckar-Raum. Articulata 2019, 33, 69–72. [Google Scholar]
- Ivković, S.; Đurđević, A.; Popović, M. New data on the occurrence of Anacridium aegyptium (Orthoptera: Acrididae) in Serbia. Acta Ent. Ser. 2015, 20, 163–166. [Google Scholar]
- Sardet, E.; Roesti, C.; Braud, Y. Grasshoppers of Britain and Western Europe. A Photographic Guide; Bloomsbury Publishing Plc: London, UK, 2021; pp. 1–304. [Google Scholar]
- Colgan, D.J.; Hutchings, P.A.; Brown, S. Phylogenetic relationships within the Terebellomorpha. J. Mar. Biol. Assoc. UK 2001, 81, 765–773. [Google Scholar] [CrossRef]
- Larsson, A. AliView: A fast and lightweight alignment viewer and editor for large data sets. Bioinformatics 2014, 30, 3276–3278. [Google Scholar] [CrossRef] [PubMed]
- Ratnasingham, S.; Wei, C.; Chan, D.; Agda, J.; Agda, J.; Ballesteros-Mejia, L.; Ait Boutou, H.; El Bastami, Z.M.; Ma, E.; Manjunath, R.; et al. BOLD v4: A Centralized, Bioinformatics Platform for DNA-Based Biodiversity Data. In DNA Barcoding. Methods in Molecular Biology; Humana: New York, NY, USA, 2024; pp. 403–441. [Google Scholar]
- Altschul, S.F.; Gish, W.; Miller, W.; Myers, E.W.; Lipman, D.J. Basic local alignment search tool. J. Mol. Biol. 1990, 215, 403–410. [Google Scholar] [CrossRef]
- Clark, K.; Karsch-Mizrachi, I.; Lipman, D.J.; Ostell, J.; Sayers, E.W. GenBank. Nucleic Acids Res. 2016, 44, D67–D72. [Google Scholar] [CrossRef] [PubMed]
- Nei, M. Molecular Evolutionary Genetics; Columbia University Press: Manhattan, NY, USA, 1987; pp. 1–505. [Google Scholar] [CrossRef]
- Tajima, F. Unbiased estimation of evolutionary distance between nucleotide sequences. Mol. Biol. Evol. 1993, 10, 677–688. [Google Scholar] [CrossRef] [PubMed]
- Rozas, J.; Ferrer-Mata, A.; Sánchez-DelBarrio, J.C.; Guirao-Rico, S.; Librado, P.; Ramos-Onsins, S.E.; Sánchez-Gracia, A. DnaSP6: DNA sequence polymorphism analysis of large data sets. Mol. Biol. Evol. 2017, 34, 3299–3302. [Google Scholar] [CrossRef]
- Bandelt, H.J.; Forster, P.; Röhl, A. Median-joining networks for inferring intraspecific phylogenies. Mol. Biol. Evol. 1999, 16, 37–48. [Google Scholar] [CrossRef]
- Leigh, J.W.; Bryant, D. PopART: Full-feature software for haplotype network construction. Methods Ecol. Evol. 2015, 6, 1110–1116. [Google Scholar] [CrossRef]
- Tamura, K.; Stecher, G.; Kumar, S. MEGA11: Molecular evolutionary genetics analysis version 11. Mol. Biol. Evol. 2021, 38, 3022–3027. [Google Scholar] [CrossRef]
- Ćato, S.; Zagorac, D. Unexpected faunistic records of Rhacocleis annulata; Eyprepocnemis plorans; and Xya pfaendleri (Orthoptera) from Croatia and Slovenia. Nat. Croat. 2021, 30, 501–511. [Google Scholar] [CrossRef]
- Ferenca, R.; Budrys, E. First records of southern oak bush-cricket Meconema meridionale Costa, 1860: (Orthoptera, Tettigoniidae) in Lithuania. Lietuvos Entomologų Draugijos Darbai 2025, 9, 5–9. [Google Scholar]
- Marchioro, C.A.; Krechemer, F.S. Prevention is better than cure: Integrating habitat suitability and invasion threat to assess global biological invasion risk by insect pests under climate change. Pest Manag. Sci. 2021, 77, 4510–4520. [Google Scholar] [CrossRef]
- Tăuşan, I.; Markó, B. New records of the pharaoh ant Monomorium pharaonis (Linnaeus, 1758) (Hymenoptera: Formicidae) in Romania. Entomol. Romanica 2011, 16, 23–26. [Google Scholar]
- Kulessa, A.K.; Balzani, P.; Soto, I.; Kouba, A.; Renault, D.; Tarkan, A.S.; Haubrock, P.J. The neglect of nonnative orthopterans as potential invaders: A call for awareness. Insect Sci. 2023, 31, 994–1000. [Google Scholar] [CrossRef] [PubMed]
- Hemala, V.; Svetlík, J.; Nuhlíčková, S. First record of the Common Crevice-cricket Gryllomorpha dalmatina (Ocskay, 1832) (Orthoptera: Gryllidae) in Slovakia: Evidence of a viable population. BioInv. Rec. 2023, 12, 555–562. [Google Scholar] [CrossRef]
- Reinhardt, K.; Köhler, G. A locust Acanthacris ruficornis (Fabricius, 1787) in a flower shop of Tübingen, Germany (Acrididae: Cyrtacanthacridinae). Articulata 2014, 29, 93–97. [Google Scholar]
- Gojznikar, J.; Poljanec, N.; Mlakar Medved, M. An interesting new record of Egyptian locust Anacridium aegyptium (Linnaeus, 1764) (Orthoptera: Acrididae) for Slovenian inland. Nat. Slov. 2018, 20, 33–34. [Google Scholar] [CrossRef]
- Pavlović, M. Egyptian locust (Anacridium aegyptium) (Acrididae: Cyrtacanthacridinae) in the Pannonian part of Croatia. Articulata 2019, 34, 95–100. [Google Scholar]
- Vlk, R.; Balvín, O.; Krištín, A.; Marhoul, P.; Hrúz, V. Distribution of the Southern Oak Bush-cricket Meconema meridionale (Orthoptera; Tettigoniidae) in the Czech Republic and Slovakia. Folia Oecol. 2012, 39, 155–165. [Google Scholar]
- Olenici, N.; Bălăcenoiu, F.; Tomescu, R.; Netoiu, C.; Buzatu, A.; Alexandru, A. Invasive alien forest insect species in south-eastern Romania. Not. Bot. Horti. Agrobo. 2022, 50, 12618. [Google Scholar] [CrossRef]
- Sîrbu, C.; Miu, I.; Gavrilidis, A.; Grădinaru, S.; Niculae, I.; Preda, C.; Oprea, A.; Urziceanu, M.; Camen-Comănescu, P.; Nagodă, E.; et al. Distribution and pathways of introduction of invasive alien plant species in Romania. NeoBiota 2022, 75, 1–21. [Google Scholar] [CrossRef]
- Hochkirch, A.; Nieto, A.; García Criado, M.; Cálix, M.; Braud, Y.; Buzzetti, F.M.; Chobanov, D.; Odé, B.; Presa Asensio, J.J.; Willemse, L.; et al. European Red List of Grasshoppers, Crickets and Bush-Crickets; Publications Office of the European Union: Luxembourg, 2016; pp. 1–86. [Google Scholar]
- Hochkirch, A.; Willemse, L.P.M.; Szövényi, G.; Rutschmann, F.; Presa, J.J.; Krištín, A.; Kleukers, R.; Chobanov, D.P. Meconema thalassinum. The IUCN Red List of Threatened Species: e.T68427417A74540428. Available online: https://www.iucnredlist.org/species/68427417/74540428 (accessed on 12 October 2025).
- Rohde, K.; Hau, Y.; Weyer, J.; Hochkirch, A. Wide prevalence of hybridization in two sympatric grasshopper species may be shaped by their relative abundances. BMC Evol. Biol. 2015, 15, 191. [Google Scholar] [CrossRef][Green Version]

| Crt. No. | Country | Collecting Site | Coordinates | Bold ID | Haplotype |
|---|---|---|---|---|---|
| 1. | Poland | Pulawy | 51.415° N, 21.956° E | BGEPL2159-25 | Hap_6 |
| 2. | Poland | Mroczk | 52.028° N, 22.225° E | BGEPL2272-25 | Hap_7 |
| 3. | Poland | Pulawy | 51.415° N, 21.956° E | BGEPL2281-25 | Hap_9 |
| 4. | Poland | Pustynia, Kozlowska | 51.404° N, 15.647° E | BGEPL2297-25 | Hap_9 |
| 5. | Poland | Pustynia, Kozlowska | 51.404° N, 15.647° E | BGEPL2322-25 | Hap_10 |
| 6. | Slovakia | Rybnicna, Bratislava | 48.215° N, 17.18° E | UZINS242-23 | Hap_1 |
| 7. | Slovakia | Rybnicna, Bratislava | 48.215° N, 17.18° E | UZINS243-23 | Hap_1 |
| 8. | United Kingdom | - | 52.145° N, −0.025° E | NTWE5731-25 | Hap_9 |
| 9. | United Kingdom | - | 51.291° N, −0.656° E | PBRI772-24 | Hap_7 |
| 10. | United Kingdom | Canvey Wick | - | DTNHM8266-23 | Hap_9 |
| 11. | Germany | Muenchen, Bavaria | 48.167° N, 11.504° E | FBORT177-09 | Hap_9 |
| 12. | Germany | Karlsruhe, Baden-Wuerttemberg | 49.006° N, 8.400° E | FBORT430-10 | Hap_6 |
| 13. | Germany | Karlsruhe, Baden-Wuerttemberg | 49.006° N, 8.400° E | FBORT431-10 | Hap_6 |
| 14. | Germany | Karlsruhe, Baden-Wuerttemberg | 49.006° N, 8.400° E | FBORT432-10 | Hap_6 |
| 15. | Germany | Kulmain, Bavaria | 49.896° N, 11.897° E | FBORT514-13 | Hap_4 |
| 16. | Germany | Bobenheim-Roxheim, Rhineland-Palatinate | 50° N, 8° E | GBMIX3300-16 | Hap_9 |
| 17. | Austria | Vienna | 48.203° N, 16.359° E | TDAOE2907-23 | Hap_5 |
| 18. | France | Ile-de-France, Paris | 48.845° N, 2.3625° E | GMFPT579-18 | Hap_9 |
| 19. | France | Corsica | - | LPRCH586-23 | Hap_2 |
| 20. | France | Campu di Bonza, Corsica | 41.771° N, 9.124° E | LPRCT662-21 | Hap_3 |
| 21. | Bulgaria | Botanical Garden, Varna | - | PORTH057-10 | Hap_9 |
| 22. | Romania | Făgăraș, BV | 45.841° N, 24.987° E | Mec | Hap_9 |
| 23. | Romania | Fălticeni, SV | 47.451° N, 26.295° E | Mh1 | Hap_9 |
| 24. | Romania | Suceava, SV | 47.641° N, 26.244° E | Mh2 | Hap_9 |
| 25. | Romania | Suceava, SV | 47.642° N, 26.246° E | Mh3 | Hap_9 |
| 26. | Romania | Fălticeni, SV | 47.451° N, 26.295° E | Mh4 | Hap_9 |
| 27. | Romania | Suceava, SV | 47.642° N, 26.247° E | Mh8 | Hap_8 |
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Iorgu, I.-Ș.; Lobiuc, A.; Ungurean, E.-A.; Tăușan, I.; Cazacu, A.; Iorgu, E.I. New Guests in the Romanian Fauna and Pathways for the Introduction of Alien Bush-Crickets and Grasshoppers (Insecta: Orthoptera). Diversity 2026, 18, 167. https://doi.org/10.3390/d18030167
Iorgu I-Ș, Lobiuc A, Ungurean E-A, Tăușan I, Cazacu A, Iorgu EI. New Guests in the Romanian Fauna and Pathways for the Introduction of Alien Bush-Crickets and Grasshoppers (Insecta: Orthoptera). Diversity. 2026; 18(3):167. https://doi.org/10.3390/d18030167
Chicago/Turabian StyleIorgu, Ionuț-Ștefan, Andrei Lobiuc, Erica-Alexandra Ungurean, Ioan Tăușan, Alexandru Cazacu, and Elena Iulia Iorgu. 2026. "New Guests in the Romanian Fauna and Pathways for the Introduction of Alien Bush-Crickets and Grasshoppers (Insecta: Orthoptera)" Diversity 18, no. 3: 167. https://doi.org/10.3390/d18030167
APA StyleIorgu, I.-Ș., Lobiuc, A., Ungurean, E.-A., Tăușan, I., Cazacu, A., & Iorgu, E. I. (2026). New Guests in the Romanian Fauna and Pathways for the Introduction of Alien Bush-Crickets and Grasshoppers (Insecta: Orthoptera). Diversity, 18(3), 167. https://doi.org/10.3390/d18030167

