The spread of Aoroides longimerus Ren & Zheng, 1996 across the Mediterranean and the Atlantic: genetic diversity, anthropogenic transport, and ecological implications
Published:
Oct 17, 2025
Keywords:
early detection non-indigenous Amphipoda taxonomic expertise scientific collaboration aquaculture recreational boating
Abstract
Accurate knowledge of the global distribution of non-indigenous species (NIS) is essential for understanding their invasion dynamics and for implementing timely management measures. This study reports the first records of the marine amphipod Aoroides longimerus, putatively native to the East Asian coast, in Italy (2018), Spain (2019) and Tunisia (2022) and provides the earliest documented record of the species in its introduced European range (Portugal, 2011). Furthermore, it expands the species’ known distribution in mainland Portugal (including the northernmost record), Macaronesia (with first records in the Canary Islands and Madeira), and confirms its presence along the Atlantic coasts of France and the Netherlands. Aquaculture facilities, particularly those associated with oyster farming, are probably the primary vector of introduction, while recreational boating may have contributed significantly to secondary dispersal. In comparison to other exotic amphipods, A. longimerus remains absent from many marinas, ports and/or aquaculture facilities in the Mediterranean Sea and adjacent regions, such as the Red Sea. This suggests that the species may still be in the early stages of expansion, or that its dispersal ability across marinas and ports is more limited than that of other amphipods. The present study, however, highlights several key ecological traits of A. longimerus: (i) it can survive year-round despite seasonal fluctuations, and reach high local densities, (ii) it exhibits strong colonisation capacity, as shown by its rapid establishment on settlement plates, and (iii) it displays opportunistic feeding behaviour, primarily consuming detritus. These characteristics underscore the importance of continuous surveillance and effective communication with stakeholders to prevent further expansion of this species.
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GUERRA-GARCÍA, J. M., DESIDERATO, A., MUCCIOLO, S., NAVARRO-BARRANCO , C., ROS, M., FERRARIO, J., MORETTI, P., MARCHINI, A., MOSBAHI, N., RUIZ-VELASCO, S., GESTOSO, I., RAMALHOSA, P., CANNING-CLODE, J., OLABARRIA, C., CACABELOS, E., TRONCOSO, J. S., PEZY, J.-P., RAOUX, A., DAUVIN, J.-C., CHARBONNELLE, M., SCHLICKLIN, F., ARIYAMA, H., MANCINI, E., BONIFAZI , A., LE GARREC, V., BUREL, T., GOUILLIEUX, B., SANTO, C. E., CHAINHO, P., RIBEIRO, R. S., AFONSO , I., WANG, Y., LECLERC, J.-C., MOAL, M., HOUBIN, C., VIARD, F., THIÉBAUT, E., FERNANDEZ-GONZALEZ, V., NAVARRO-MAYORAL, S., DÍAZ-VERGARA, S., GITTENBERGER, A., HUGHES , L. E., LAMPA, M., CHATZIGEORGIOU , G., PLAITIS, W., TIRALONGO, F., MARTÍNEZ-LAIZ, G., CABEZAS, M. P., DUARTE , S., PINHEIRO, J. P., REIS, G. O., REVANALES, T., JUAN , S.-V., AYLAGAS, E., POLISENO , A., CHEBAANE, S., CARVALHO, S., GEROVASILEIOU, V., FRUTOS, I., & SAENZ-ARIAS, P. (2025). The spread of Aoroides longimerus Ren & Zheng, 1996 across the Mediterranean and the Atlantic: genetic diversity, anthropogenic transport, and ecological implications. Mediterranean Marine Science, 26(4), 842–859. https://doi.org/10.12681/mms.41784
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References
Almón, B., Pérez, J., Bañón, R., Trigo, J., 2014. First record of Caprella mutica from the Iberian Peninsula: expansion southwards in European waters. Marine Biodiversity Records, 7, e30.
Arfianti, T., Costello, M.J., 2020. Global biogeography of marine amphipod crustaceans: latitude, regionalization, and beta diversity. Marine Ecology Progress Series, 638, 83-94.
Ariyama, H., 2004. Nine species of the genus Aoroides (Crustacea: Amphipoda: Aoridae) from Osaka Bay, Central Japan. Publications of the Seto Marine Biological Laboratory, 40 (1-2), 1-66.
Ariyama, H., 2022. Guidebook of Gammaridean Amphipoda. Kaibundo, Tokyo, 160 pp. (in Japanese).
Ariyama, H., Kawabe, K., 2022. Two new species of Aoridae from Chichijima Island, the Ogasawara Islands in Japan (Crustacea: Amphipoda). Species Diversity, 27 (1), 113-128. A
riyama, H., Kohtsuka, H., 2022. Three new species of the family Aoridae collected from Sagami Bay, central Japan (Crustacea: Amphipoda). Zootaxa, 5159 (3), 393-413.
Bailey, S.A., Brown, L., Campbell, M.L., Canning‐Clode, J., Carlton, J.T. et al., 2020. Trends in the detection of aquatic non‐indigenous species across global marine, estuarine and freshwater ecosystems: A 50‐year perspective. Diversity and Distributions, 26 (12), 1780-1797.
Barnard, J.L., 1970. Sublittoral Gammaridea (Amphipoda) of the Hawaiian Islands. Smithsonian Contributions to Zoology, 34, 1-286.
Barnard, J.L., Karaman, G.S., 1991. The families and genera of marine gammaridean Amphipoda (except marine gammaroids). Records of the Australian Museum, (Suppl. 13) Part 1, 1-417.
Bell, S.S., 1991. Amphipods as insect equivalents? An alternative view. Ecology, 72 (1), 350-354.
Bello, C.L., Cabrera, M.I., 1999. Uso de la técnica microhistológica de Cavender y Hansen en la identificación de insectos acuáticos. Boletín Entomológico de Venezuela, 14, 77-79.
Beermann, J., Hall-Mullen, A.K., Havermans, C., Coolen, J.W.P., Crooijmans, R.P.M.A. et al., 2020. Ancient globetrotters connectivity and putative native ranges of two cosmopolitan biofouling amphipods. PeerJ, 8, e9613.
Carvalho, S., Shchepanik, H., Aylagas, E., Berumen, M.L., Costa, F.O. et al., 2023. Hurdles and opportunities in implementing marine biosecurity systems in data-poor regions. BioScience, 73 (7), 494-512.
Casquet, J., Thebaud, C., Gillespie, R.G., 2012. Chelex without boiling, a rapid and easy technique to obtain stable amplifiable DNA from small amounts of ethanol-stored spiders. Molecular Ecology Resources, 12, 136-141.
Castro, N., Carlton, J.T., Costa, A.C., Marques, C.S., Hewitt, C. et al., 2022. Diversity and patterns of marine non‐native species in the archipelagos of Macaronesia. Diversity and Distributions, 28 (4), 667-684.
Chevreux, E., Fage, L., 1925. Amphipodes. Volume 9. Paris: Office Central de Faunistique, Librairie de la Faculté des Sciences.
Clark, K., Karsch-Mizrachi, I., Lipman, D.J., Ostell, J., Sayers, E.W., 2016. GenBank. Nucleic Acids Research, 44 (D1), D67-72. CIEMAR, 2021. Substrato duro subtidal. Monitorização de Ambientes Marinhos do Porto de Sines - MAPSi 2018/2020. Relatório final. Laboratório de Ciências do Mar da Universidade de Évora. 191 pp. (report for the Port of Sines Administration).
CIEMAR, 2025. Substrato duro subtidal. Monitorização de Ambientes Marinhos do Porto de Sines - MAPSi 2021/2023. Relatório final. Laboratório de Ciências do Mar da Universidade de Évora. 275 pp. (report for the Port of Sines Administration).
Cuthbert, R.N., Pattison, Z., Taylor, N.G., Verbrugge, L., Diagne, C. et al., 2021. Global economic costs of aquatic invasive alien species. Science of the Total Environment, 775, 145238.
Conlan, K.E., Bousfield, E.L., 1982. The superfamily Corophioidea in the North Pacific region. Family Aoridae: systematics and distributional ecology. National Museum of Natural Sciences (Ottawa), Publications in Biological Oceanography, 10, 77-101.
Dauvin, J.-C., Pezy, J.-P., Raoux, A., 2020. First records of Aoroides longimerus Ren and Zheng, 1996, and A. semicurvatus Ariyama, 2004 (Crustacea, Amphipoda), in the English Channel, France. BioInvasions Records, 4, 753-762.
DQEM, 2020. Estratégia Marinha. Relatório do 2º ciclo, Parte D - Reavaliação do Estado Ambiental e Definição de Metas, subdivisão dos Açores. Governo dos Açores, Diretiva Quadro Estratégia Marinha, março 2020.
Faasse, M., Gittenberger, A., 2021. De exotische vlokreeften Aoroides longimerus en Erichthonius didymus bereiken Nederland (Crustacea: Amphipoda). Nederlandse Faunistische Mededelingen, 57, 81-88.
Galanidi, M., Gratsia, E., Zenetos, A., 2025. Backdating first records of non-indigenous species in the Mediterranean: are initial findings underestimated? Mediterranean Marine Science, 26 (3), 642-656.
Gouillieux, B., Lavesque, N., Leclerc, J.-C., Le Garrec, V., Viard, F. et al., 2015. Three non-indigenous species of Aoroides (Crustacea: Amphipoda: Aoridae) from the French Atlantic Coast. Journal of the Marine Biological Association of the United Kingdom, 96 (8), 1651-1659.
Guerra-García, J.M., Tierno de Figueroa, J.T., Navarro-Barranco, C., Ros, M., Sánchez-Moyano, J.E. et al., 2014. Dietary analysis of the marine Amphipoda (Crustacea: Peracarida) from the Iberian Peninsula. Journal of Sea Research, 85, 508-517.
Guerra-García, J.M., Revanales, T., Saenz-Arias, P., Navarro- Barranco, C., Ruiz-Velasco, S. et al., 2023. Quick spreading of the exotic amphipod Laticorophium baconi (Shoemaker, 1934): another small stowaway overlooked? Mediterranean Marine Science, 24 (3), 644-665.
Guindon, S., Dufayard, J., Lefort, V., Anisimova, M., Hordijk, W.O.G. et al., 2010. New algorithms and methods to estimate maximum-likelihood phylogenies: Assessing the performance of PhyML 3 .0. Systematic Biology, 59 (3), 307-321.
Gurjanova, E., 1938. Amphipoda, Gammaroidea zalikov Siaukhu i Sudzukhe (Yaponskoe More). [Amphipoda, Gammaroidea of Siaukhu Bay and Sudzukhe Bay (Japan Sea)]. Reports of the Japan Sea Hydrobiological Expedition of the Zoological Institute of the Academy of Sciences USSR in 1934. 1, 241-404.
Hou, Z., Fu, J., Li, S.A., 2007. Molecular phylogeny of the genus Gammarus (Crustacea: Amphipoda) based on mitochondrial and nuclear gene sequences. Molecular Phylogenetics and Evolution, 45 (2), 596-611.
ICES, 2018. Interim Report of the Working Group on Introductions and Transfers of Marine Organisms (WGITMO). 7-9 March 2018, Madeira, Portugal. ICES CM 2018/HAPISG: 11. 179 pp.
Iitembu, J.A., Fitzgerald, D., Altintzoglou, T., Boudry, P., Britz, P. et al., 2023. Comparative description and analysis of oyster aquaculture in selected Atlantic regions: production, market dynamics, and consumption patterns. Fishes, 8 (12), 584.
Kim, J., Choi, J. H., Kim, Y. J., Im, H.W., Kim, Y. H., 2024. Two new species of the family Aoridae (Crustacea, Malacostraca, Amphipoda) from Korean waters. ZooKeys, 1210, 117-132.
Larkin, M.A., Blackshields, G., Brown, N.P., Chenna, R., McGettigan, P.A. et al., 2007. Clustal W and Clustal X version 2.0. Bioinformatics, 23, 2947-2948.
Lefort, V., Longueville, J.-E., Gascuel, O., 2017. SMS: Smart model selection in PhyML. Molecular Biology and Evolution, 34 (9), 2422-2424.
Lincoln, R.J., 1979. British marine Amphipoda: Gammaridea. London: British Museum (Natural History).
Lobo, J., Costa, P.M., Teixeira, M.A., Ferreira, M.S.G., Costa, M.H. et al., 2013. Enhanced primers for amplification of DNA barcodes from a broad range of marine metazoans. BMC Ecology, 13, 34.
Macdonald III, K.S., Yampolsky, L., Duffy, J.E., 2005. Molecular and morphological evolution of the amphipod radiation of Lake Baikal. Molecular Phylogenetics and Evolution, 35, 323-343.
Marchini, A., Cardeccia, A., 2017. Alien amphipods in a sea of troubles: cryptogenic species, unresolved taxonomy and overlooked introductions. Marine Biology, 164 (4), 69.
Martínez-Laiz, G., Ros, M., Guerra-García, J.M., Marchini, A., Fernandez-Gonzalez, V. et al., 2020. Scientific collaboration for early detection of invaders results in a significant update on estimated range: Lessons from Stenothoe georgiana Bynum & Fox 1977. Mediterranean Marine Science, 21 (2), 464-481.
Martínez-Laiz, G., MacLeod, C.D., Hesketh, A.V., Konecny, C.A., Ros, M. et al., 2022. The journey of hull-fouling mobile invaders: basibionts and boldness mediate dislodgement risk during transit. Biofouling, 38 (8), 837-851.
Mosbahi, N., Boudaya, L., Neifar, L., Dauvin, J.C., 2020. Do intertidal Zostera noltei meadows represent a favourable habitat for amphipods? The case of the Kneiss Islands (Gulf of Gabès: Central Mediterranean Sea). Marine Ecology, 41 (3), 1-16.
Myers, A.A., 1995. The Amphipoda (Crustacea) of Madang Lagoon: Aoridae, Isaeidae, Ischyroceridae and Neomegamphopidae. In Lowry, J.K. (Ed.). The Amphipoda (Crustacea) of Madang Lagoon, Papua New Guinea, Part 1. Records of the Australian Museum, Supplement, 22, 25-95.
Myers, A.A., 2009. Aoridae. In: Lowry, J.K., Myers, A.A. (Eds.) Benthic Amphipoda (Crustacea: Peracarida) of the Great Barrier Reef, Australia. Zootaxa, 2260, 220-278.
Navarro-Barranco, C., Guerra-García, J.M., Sánchez-Tocino, L., Ros, M., Florido, M. et al., 2015. Colonization and successional patterns of the mobile epifaunal community along an environmental gradient in a marine cave. Marine Ecology Progress Series, 521, 105-115.
Navarro-Barranco, C., Florido, M., Ros, M., González-Romero, P., Guerra-García, J.M., 2018. Impoverished mobile epifaunal assemblages associated with the invasive macroalga Asparagopsis taxiformis in the Mediterranean Sea. Marine Environmental Research, 141, 44-52.
Navarro-Barranco, C., Muñoz-Gómez, B., Saiz, D., Ros, M., Guerra-García, J.M. et al., 2019. Can invasive habitat-forming species play the same role as native ones? The case of the exotic marine macroalga Rugulopteryx okamurae in the Strait of Gibraltar. Biological Invasions, 21, 3319-3334.
Navarro‐Barranco, C., Moreira, J., Espinosa, F., Ros, M., Rallis, I. et al., 2021. Evaluating the vulnerability of coralligenous epifauna to macroalgal invasions. Aquatic Conservation: Marine and Freshwater Ecosystems, 31 (9), 2305-2319.
Navarro-Barranco, C., Lanza-Arroyo, P., Gutiérrez-Serrano, J., Moreira, J., 2023. Amphipod assemblages associated with native habitat-forming seaweeds of the Alboran Sea: influence by environmental protection and biogeographical patterns. Marine and Freshwater Research, 74, 50-64.
Ojaveer, H., Galil, B.S., Carlton, J.T., Alleway, H., Goulletquer, P. et al., 2018. Historical baselines in marine bioinvasions: Implications for policy and management. PloS ONE, 13 (8), e0202383.
Palumbi, S.R., 1996. What can molecular genetics contribute to marine biogeography? An urchin’s tale. Journal of Experimental Marine Biology and Ecology, 203 (1), 75-92.
Pascual, S., Villalba, A., Abollo, E., Garci, M., González, A.F. et al., 2010. The mussel Xenostrobus securis: a well-established alien invader in the Ria de Vigo (Spain, NE Atlantic). Biological Invasions, 12, 2091-2103.
Ratnasingham, S., Wei, C., Chan, D., Agda, J., Agda, J., et al. 2024. BOLD v4: A centralized bioinformatics platform for DNA-based biodiversity data. Methods in Molecular Biology, 2744, 403-441.
Ren, X., Zheng, C., 1996. Fouling Amphipoda (Crustacea) from Dayawan, Guangdong Province China (South China Sea). Annual Research Reports, Marine Biology Research Station at Dayawan, South China Sea Institute of Oceanology, the Chinese Academy of Sciences, 1, 58-78 [in Chinese with English abstract and descriptions of new species].
Rius, M., Turon, X., Bernard, G., Volckaert, F., Viard, F., 2015. Marine invasion genetics: from spatial and temporal patterns to evolutionary outcomes. Biological Invasions, 17 (3), 869-885.
Ros, M., Guerra-García, J.M., 2012. On the occurrence of the tropical caprellid Paracaprella pusilla Mayer, 1890 (Crustacea: Amphipoda) in Europe. Mediterranean Marine Science, 13 (1), 134-139.
Ros, M., Guerra-García, J.M., Navarro-Barranco, C., Cabezas, M.P., Vázquez-Luis, M., 2014. The spreading of the non-native caprellid (Crustacea: Amphipoda) Caprella scaura Templeton, 1836 into southern Europe and northern Africa: a complicated taxonomic history. Mediterranean Marine Science, 15 (1), 145-155.
Ruffo, S. (Ed.), 1982. The Amphipoda of the Mediterranean. Part 1. Gammaridea (Acanthonotozomatidae to Gammaridae). Mémoires de l’Institut Océanographique de Monaco, 13, 1-364.
Ruiz, C., Abad, M., Sedano, F., Garcia-Martin, L.O., Sánchez- López, J.L., 1992. Influence of seasonal environmental changes on the gamete production and biochemical composition of Crassostrea gigas (Thunberg) in suspended culture in El Grove, Galicia, Spain. Journal of Experimental Marine Biology and Ecology, 155 (2), 249-262.
Saenz-Arias, P., Navarro-Barranco, C., Ros, M., Moreira, J., Guerra-García, J.M., 2022a. Exploring biocontamination in associated macrofaunal assemblages in marinas: Soft bottoms vs artificial hard substrate. Where and what to look for? Marine Pollution Bulletin, 185, 114346.
Saenz-Arias, P., Navarro-Barranco, C., Guerra-García, J.M., 2022b. Influence of environmental factors and sessile biota on vagile epibionts: The case of amphipods in marinas across a regional scale. Mediterranean Marine Science, 23 (1), 1-13.
Spalding, M.D., Fox, H.E., Allen, G.R., Davidson, N., Ferdaña, Z.A. et al., 2007. Marine ecoregions of the world: a bioregionalization of coastal and shelf areas. BioScience, 57 (7), 573-583.
Sun, Y., Wong, E., Keppel, E., Williamson, J.E., Kupriyanova, E.K., 2017. A global invader or a complex of regionally distributed species? Clarifying the status of an invasive calcareous tubeworm Hydroides dianthus (Verrill, 1873) (Polychaeta: Serpulidae) using DNA barcoding. Marine Biology, 164, 1-12.
Tamura, K., Stecher, G., Kumar, S., 2021. MEGA11: Molecular Evolutionary Genetics Analysis version 11. Molecular Biology and Evolution, 38 (7), 3022-3027.
Ulman, A., Ferrario, J., Occhpinti-Ambrogi, A., Arvanitidis, C., Bandi, A. et al., 2017. A massive update of non-indigenous species records in Mediterranean marinas. PeerJ, 5, e3954.
Walker, A.O., 1898. Crustacea collected by W.A. Herdmann, F.R.S., in Puget Sound, Pacific coast of North America, September 1897. Transactions of the Liverpool Biological Society, 12, 268-287.
WoRMS Editorial Board, 2025. World register of marine species. https://www. marinespecies.org. at VLIZ.
Zenetos, A., Gratsia, E., Cardoso, A.C., Tsiamis, K., 2019. Time lags in reporting of biological invasions: the case of Mediterranean Sea. Mediterranean Marine Science, 20 (2), 469-475.
