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General and hinge views of the studied species: native Modiolus adriaticus from Dzharylhach Bay (A, C), Arcuatula senhousia from Sukhoy Liman (B, D, E) and Mytilopsis leucophaeata from the Southern Bug River (F, G); arrows indicate the diagnostic features of the species: hinge teeth (D) and crenulate anterior margin near the umbo (E) in A. senhousia (vs. smooth anterior margin in M. adriaticus); the apophysis of M. leucophaeata (G). Photos by M. Son.

General and hinge views of the studied species: native Modiolus adriaticus from Dzharylhach Bay (A, C), Arcuatula senhousia from Sukhoy Liman (B, D, E) and Mytilopsis leucophaeata from the Southern Bug River (F, G); arrows indicate the diagnostic features of the species: hinge teeth (D) and crenulate anterior margin near the umbo (E) in A. senhousia (vs. smooth anterior margin in M. adriaticus); the apophysis of M. leucophaeata (G). Photos by M. Son.

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Conrad's false mussel Mytilopsis leucophaeata (Conrad, 1831) and the Asian date mussel Arcuatula senhousia (Benson, 1842) are highly invasive bivalves actively spreading in the Ponto-Caspian region, which includes the basins of the Black, Azov, and Caspian seas. This study provides new records of both species from this region and gives a synopsis o...

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Context 1
... leucophaeata (Figure 2) can be recognized by the presence of an apophysis, a tubercular structure under the anterior interior of the shell, where the foot muscles are attached ( Figure 2G). This structure distinguishes this species from all local species of Dreissenidae and Mytilidae. ...
Context 2
... leucophaeata (Figure 2) can be recognized by the presence of an apophysis, a tubercular structure under the anterior interior of the shell, where the foot muscles are attached ( Figure 2G). This structure distinguishes this species from all local species of Dreissenidae and Mytilidae. ...
Context 3
... more reliable diagnostic features may be relatively small apophysis of M. leucophaeata, which is rounded or pointed posteriorly, in contrast to a large, anteriorly pointed apophysis in M. sallei (Marelli and Gray 1983). In most of our M. leucophaeata specimens the apophysis is not visible at all in standard projection, but it is clearly discernible when the shell is held at an angle ( Figure 2G). Arcuatula senhousia (Figure 2) differs from the native Modiolus adriaticus Lamarck, 1819 by its shell and hinge sculpture ( Figure 2D, E). ...
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... most of our M. leucophaeata specimens the apophysis is not visible at all in standard projection, but it is clearly discernible when the shell is held at an angle ( Figure 2G). Arcuatula senhousia (Figure 2) differs from the native Modiolus adriaticus Lamarck, 1819 by its shell and hinge sculpture ( Figure 2D, E). There are anteriorly several mild ridges causing a crenulate anterior margin of the shell of A. senhousia ( Zenetos et al. 2004), which is more clearly visible on the inside of the valves. ...
Context 5
... most of our M. leucophaeata specimens the apophysis is not visible at all in standard projection, but it is clearly discernible when the shell is held at an angle ( Figure 2G). Arcuatula senhousia (Figure 2) differs from the native Modiolus adriaticus Lamarck, 1819 by its shell and hinge sculpture ( Figure 2D, E). There are anteriorly several mild ridges causing a crenulate anterior margin of the shell of A. senhousia ( Zenetos et al. 2004), which is more clearly visible on the inside of the valves. ...

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... This species may form dense, continuous mats of ovoid cocoons anchored on the sediment with a matrix of byssal threads (Morton 1974;Sousa et al. 2009). Native to East Asia between the South China Sea and Siberia (Cheung et al. 1962in Morton 1974Kulikova 1978), the species has invaded several coastal ecosystems around the globe, including the Indian Ocean (India, Madagascar, Mauritius, Zanzibar, and the adjacent Persian Gulf) (Barash and Danin 1972;Behera et al. 2019;Yasser et al. 2023), coastal ecosystems scattered along the Eastern Atlantic Ocean (e.g., southern United Kingdom, Bay of Biscay, central Portugal, Guinea-Bissau) (Lourenço et al. 2018;Cabral et al. 2020;Watson et al. 2021;Massé et al. 2022), coastlines along the Mediterranean Sea (e.g., Spain, France, Italy, Croatia, Albania, Turkey, Israel, Tunisia, Algeria) (Hamza et al. 2022) and in the adjacent Black Sea (Zhulidov et al. 2021), off Venezuela on the Western Atlantic Ocean (Martínez-Escarbassiere et al. 2003), Eastern coastal ecosystems of the Pacific Ocean (e.g., Mexico, Colombia) (Crooks 1998;Arellano and Salgado-Barragán 2012;Osorio 2013), and in the southwestern Pacific Ocean (e.g., Australia, New Zealand) (Willan 1987;Creese et al. 1997). The Asian date mussel is an epifaunal bivalve observed in sites with salinities between 5 and 18 (Yamamuro et al. 2010;Hosozawa et al. 2020), but is able to colonize fully marine areas, as observed in the Solent (southern England) (Watson et al. 2021) and French localities along the Bay of Biscay (Massé et al. 2022). ...
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Ecosystem engineering bivalves can shape aquatic ecosystems because their high filtration capacity changes water quality and their shells increase the fractal dimension of benthic ecosystems with consequent abiotic and biotic effects. The Asian date mussel Arcuatula senhousia (W. H. Benson, 1842), native to East Asia between the South China Sea and Siberia, is one such bivalve that, despite its small size, can reshape a benthic ecosystem when forming dense, continuous mats. We describe here the first detected population of this non-indigenous species in southern Portugal. The Asian date mussel was found in the middle portion of the Guadiana estuary in 2022. There, river flow has been highly regulated since the construction of the biggest European reservoir in 2002, which may have been the precursor for the establishment of numerous non-indigenous species. We also discuss if this new non-indigenous species indicates an ongoing invasion meltdown process or if it can be framed under the empty niche or niche replacement hypothesis. So far, there is only circumstantial evidence supporting the niche theory hypotheses, but the interaction of several hypotheses promoting the spread and establishment of this species is also likely. Moving forward, better-informed management and conservation efforts should rely on new empirical and experimental evidence to understand the establishment mechanisms of nonindigenous species in the Guadiana estuary.
... Mytilopsis leucophaeata es originaria de la región atlántica norteamericana, desde el golfo de México hasta la bahía de Chesapeake en EEUU (Marelli & Gray, 1983;Kennedy, 2011;Eeuwes, 2018). Se trata de una especie propia de ecosistemas estuarinos y lagunas litorales, que ha invadido numerosos lugares en todo el mundo: norte de Brasil, costas europeas del Atlántico, Báltico, Mediterráneo, mar Negro y mar Caspio (Therriault et al., 2004;Eeuwes, 2018;Zhulidov et al., 2015Zhulidov et al., , 2021Zulian & Quaggiotto, 2020). Se considera que su invasión ha sido mediada principalmente por las embarcaciones, bien a través de larvas en aguas de lastre, o por incrustaciones de ejemplares en el casco de buques (Eeuwes, 2018). ...
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The second Iberian citation of Mytilopsis leucophaeata is given, after the finding of about two hundred specimens in the Guadiana River. This is a species with high invasive capability which may potentially colonize most of the surrounding estuarine and brackish environments. It is also the first record of the species from Portugal.
... Severe fragmentation of the estuary by dams and embayments, as well as its natural morphology, has resulted in the formation of many semi-isolated areas with different hydrological conditions. As a result, many alien species with diverse ecological preferences, including fully marine species, estuarine brackish water species, freshwater species and oligohaline Ponto-Caspian relicts, have become established and/or invasive (Zaitsev et al. 2006;Son 2007Son , 2008Krasnovyd et al. 2012;Vinogradov and Khutornoy 2013;Son et al. 2013Son et al. , 2020Zhulidov et al. 2021). This diversity in habitat conditions also provides a diversity of invasive species pathways, with marine routes, whether by shipping or naturally through the channel connecting the estuary to the sea, being the most important. ...
... This diversity in habitat conditions also provides a diversity of invasive species pathways, with marine routes, whether by shipping or naturally through the channel connecting the estuary to the sea, being the most important. Passage through the channel is typical for the many mollusk, worm, and crustacean species now established (Zaitsev et al. 2006;Son et al. 2013;Zhulidov et al. 2021); however, fish stocking in the estuary's desalinated areas and incidental introduction of alien 'hitchhikers' alongside these stocked fish has also had an important impact (Kvach and Kutsokon 2017;Son et al. 2020). Finally, exotic and Ponto-Caspian mollusks have been introduced into small tributaries and the upper parts of the Sukhyi Lyman through natural dispersal, possibly with the additional help of migratory waterfowl (Son 2007(Son , 2008. ...
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Two new fish species were recorded while monitoring the fish fauna of Sukhyi Lyman (or Sukhyi Estuary), an estuary in the northwestern Black Sea (Ukraine) close to the marine port of Chornomorsk. A single pumpkinseed, Lepomis gibbosus (Linnaeus, 1758), was observed in May 1999, followed by numerous observations in May 2021, while a single peacock blenny, Salaria pavo (Risso, 1810), was photographed during snorkeling observations in September 2021. The estuary is a known hub for invasive alien species and other neobionts due to the presence of the marine port. The new population of pumpkinseed, a North American invasive species, in the estuary represents the next stage in the species’ ongoing expansion along the Ukrainian Black Sea coast. Likewise, the appearance of the peacock blenny represents the latest stage in the “Mediterranization” of the Black Sea, a process that has been ongoing over the last several thousand years.
... As a result of these studies, important new findings of six species of aquatic macroinvertebrates were obtained (Table 1 and Table 2, Figure 2). The trumpet ram's-horn Menetus dilatatus (Gould, 1841) is first reported for the Danube Basin, and three marine species (Asian date mussel Arcuatula senhousia (Benson, 1842); spionid polychaetes Polydora cornuta Bosc, 1802 and Streblospio gynobranchiata Rice & Levin, 1998) for the first time for Data on the morphology and identification features of most of these species in the Danube Delta or nearby regions have been already published in several papers (Son 2007;Radashevsky and Selifonova 2013;Aleksandrov et al. 2014;Teacă et al. 2021;Zhulidov et al. 2021), so we will not elaborate on them. ...
... Some of the findings (A. senhousia, P. cornuta, and S. gynobranchiata) represent species already known from the marine ecosystems of the Northwest Black Sea (Bondarenko 2011;Teacă et al. 2021;Zhulidov et al. 2021), but reported for the first time in the estuarine habitats of the Danube Delta. ...
... This was, however, just a single finding of a live animal and there has been no further confirmation of the occurrence of this species in Romania. However, re-entry into the Azov-Black Sea basin began in 2015 and over several years has covered many locations in Russia, Ukraine, and Bulgaria (Kovalev et al. 2017;Chartosia et al. 2018;Zhulidov et al. 2021). ...
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Studies of alien aquatic invertebrates in 2021–2022 covered fresh marine and transitional waters mostly within the boundaries of the Danube Biosphere Reserve. As a result of these studies, important new findings of six species of aquatic macroinvertebrates were obtained: Menetus dilatatus (Gould, 1841) is first indicated for the Danube Basin, and three marine species (Arcuatula senhousia (Benson, 1842), Polydora cornuta Bosc, 1802 and Streblospio gynobranchiata Rice & Levin, 1998) for the first time for Ukrainian Danube Delta. For two species (Pectinatella magnifca (Leidy, 1851) and Ferrissia californica (Rowell, 1863)), new localities were found within the delta. In the case of M. dilatatus, the pathway of entry into the Danube Delta is an absolute mystery. The most likely pathway of dispersal of this species is natural dispersion, both along the river network and associated with waterfowl, however, the large distance between the Danube Delta and the nearest locations in Western Ukraine makes direct transportation unlikely. In all probability, the similarity of this species with local species leads to the formation of cryptic populations, not identified by researchers, which may be intermediate stages of species expansion towards the Northern Black Sea area.
... The required salinity for the survivorship and reproduction of M. leucophaeata are below 18-21 ppt (Kennedy 2011, Van der Gaag et al. 2016, Maia-Neto et al. 2020). This species is native from the Gulf of Mexico and Southeast USA, where it has low densities, and it was introduced in Europe, Asia, Caribbean, South America and Northeast USA (Fernandes et al. 2018, 2021, Lodeiros et al. 2019, Zhulidov et al. 2021, Rodrigues et al. 2022. In Brazil, the confirmed records of M. leucophaeata are in the city of Rio de Janeiro, i.e., in the Rodrigo de Freitas Lagoon and in the Marapendi Lagoon (Rizzo et al. 2014, Fernandes et al. 2020). ...
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Mytilopsis leucophaeata is an estuarine bivalve native from the Gulf of Mexico and Southeast USA, and it was introduced in Europe, Asia, Caribbean, South America and Northeast USA, showing massive colonization skills. In Brazil, the single invasion records of M. leucophaeata occur in the city of Rio de Janeiro, i.e., in the Rodrigo de Freitas Lagoon and in the Marapendi Lagoon. We conducted a new series of fieldworks in estuaries from the Rio de Janeiro state in order to evaluate the propagation of this invasive bivalve, aiming sites with proper salinities for the establishment of M. leucophaeata. A new record is given for the Maricá-Guarapina lagoon system, where M. leucophaeata mainly colonizes hard substrata (such as piers and rocks), reaching a mean density up to 43,375 specimens/m2; however, aggregates of M. leucophaeata were also observed in the soft substratum. Based on mitochondrial sequences, the taxonomic identification of the invasive bivalve was confirmed. The associated fauna to the agglomerates of M. leucophaeata in the lagoon system comprises amphipods, barnacles, tanaidaceans, isopods, crabs, polychaetes and snails. The expansion of M. leucophaeata requires a continuous investigation due to the great circulation of boats in the littoral of the Rio de Janeiro state and the increased chance of new introductions.
... Moreover, free-living planktonic larvae and high fecundity of dreissenids facilitate species transfer to new habitats and rapid colonization (Bij de Vaate et al., 2002). In recent decades, new introductions of the dark false mussel M. leucophaeata has grown substantially in association with increased long-range ship displacements and economic globalization (Heiler, Nahavandi & Albrecht, 2010;Zhulidov et al., 2021;Rodrigues et al., 2022). The major vectors of dark false mussel transportation to non-native brackish systems are possibly hull fouling and inadequate management of ballast water (Fernandes et al., 2018). ...
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The present study aimed to evaluate and compare feeding responses of the non-native and native bivalves, the dark false mussel Mytilopsis leucophaeata and the scorched mussel Brachidontes darwinianus, respectively, by offering different concentrations of seston from the coastal lagoon where these species coexist after dark false mussel introduction (Rodrigo de Freitas Lagoon, Rio de Janeiro-Brazil). For this purpose, independent laboratory experiments were carried out under five concentrations of seston to test the differences in clearance and ingestion rates of bivalves as a function of increasing concentrations of suspended particulate matter (SPM) on seston. In addition, from the integrated analysis of data obtained in experiments, it can be inferred about the efficiency levels of these species to remove SPM from seston and their effects on water turbidity and nutrient concentrations (total carbon, nitrogen, and phosphorus). Our hypothesis was that the non-native bivalve is more efficient to clear and ingest SPM from seston compared to the native one, which may lead to competitive advantages to the successful invasion of M. leucophaeata in coastal lagoons. Native species did not show a significant difference in clearance and ingestion rates with increasing concentrations of seston. Whereas the non-native bivalve showed a slight tendency to increase its clearance and ingestion rates with the increase in seston concentrations, evidencing its plasticity to adjust its feeding responses. The native bivalve was significantly more efficient to clear and ingest SPM at the lower seston concentration (i.e., close to natural concentrations found in the lagoon) compared to the non-native bivalve, which, on the other hand, showed a significant increase in its ingestion rates at the How to cite this article Rodrigues N, Ribeiro D, C Miyahira I, G M Portugal S, N Santos L, A F Neves R. 2023. Do feeding responses of a non-native bivalve outperform the native one in a coastal lagoon? A possible explanation for the invasion success of the dark false mussel Mytilopsis leucophaeata. PeerJ 11:e15848 Distributed under Creative Commons CC-BY 4.0 higher concentration tested (140 mg SPM L −1). Thus, the present results did not suggest food competition between the non-native M. leucophaeata and the native B. darwinianus in the introduced system. However, M. leucophaeata increased its feeding response with experimental increment in seston concentration, which suggests species ability to benefit from conditions of increased inputs of organic matter and higher primary production that could mediate its establishment in introduced systems.
... The native range of A. senhousia extends from southern Siberia (Kuril Islands) to Singapore (Chuang 1961). It has been introduced into numerous locations around the world over many decades, including New Zealand and Australia (Willan 1985(Willan , 1987, the United States (Crooks 1996;Wasson et al. 2001), India (Kutty et al. 2016), West Africa (Lourenço et al. 2018) and Europe including the Mediterranean Sea (Mistri et al. 2004;Bachelet et al. 2009; Barfield et al. 2018;Zhulidov et al. 2021;Massé et al. 2022). Arcuatula senhousia is an opportunistic suspension feeder capable of establishing enormous population densities, which can alter sediment structures, habitat, communities, and transform other aspects of ecosystem functioning (Crooks 1996). ...
... Arcuatula senhousia can attach to all manner of substrates via fine byssus threads, although it has most frequently been recorded from soft, mixed sediments (Crooks 1996;Mistri et al. 2004;Magni et al. 2006;Despalatović et al. 2013;Watson et al. 2021;Zhulidov et al. 2021). Because A. senhousia usually colonizes soft substrates, it is unlikely that ship or boat hull fouling would be the vector in this case (Crooks 1996;Hosozawa et al. 2020). ...
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The invasive Asian date mussel Arcuatula senhousia has now been recorded from the Shatt Al-Basrah Canal in the south of Iraq. High densities of up to 102 ind.m-2 were found on rocky substrates in the intertidal zone during March 2020. The species was confirmed based on morphological characters of the shell. We hypothesize that the initial introduction was associated with shipping, with secondary dispersion into the Shatt Al-Basrah canal. It is possible that the species could extend its distribution into the Shatt Al-Arab and northern Arabian Gulf naturally or by additional anthropogenic means.
... Arcuatula senhousia has successfully colonised various areas of the world, including the Pacific coast of North America, Australia, New Zealand, the Mediterranean Sea, the French Atlantic coast, the English Channel, West Africa, the Sea of Azov and the Black Sea (Cohen 2005;Bachelet et al. 2009;Chartosia et al. 2018;Watson et al. 2021;Zhulidov al. 2021). In the Mediterranean Sea, A. senhousia was first recorded in 1964 as Modiolus arcuatulus (from the coastal Bardawil Lagoon, southern Levantine Basin, Egypt) and Arcuatula arcuatula (Hanley, 1843) (from "Tel Baruch", eastern Levantine Basin, Tel Aviv) Danin 1971, 1973) (Figure 1). ...
... Mytilopsis leucophaeata (Conrad 1831) is an estuarine bivalve native from the Gulf of Mexico and possibly part of the eastern USA, where it is usually found in low densities (Kennedy 2011a). The invasion of M. leucophaeata is widely documented in many regions around the globe, such as northeastern USA (e.g., Kennedy 2011a; Richardson and Hammond 2016), Caribbean (e.g., Mohammed et al. 2018;Lodeiros et al. 2019;Minchin and Cottier-Cook 2020), Brazil (e.g., Rizzo et al. 2014;Fernandes et al. 2020), Europe (e.g., Verween et al. 2010;Heiler et al. 2010;Zhulidov et al. 2015;Forsström et al. 2016), and the Ponto-Caspian region (e.g., Heiler et al. 2010;Mumladze et al. 2019;Zhulidov et al. 2021). Such invasions have substantially increased in the last decades due to faster ship dislodgements and increased global commerce, although old establishments were also detected (Oliver 2015). ...
... Explosive densities of M. leucophaeata in invaded sites may promote severe ecological and economic impacts (Verween et al. 2010;Kennedy 2011a;Florin et al. 2013), regarding their role as ecosystem-engineers (Neves et al. 2020;Rodrigues et al. 2021). Despite the main colonization of this species on hard substrata, the formation of druses or even biogenic reefs on soft substrata was also recorded (Fernandes et al. 2020;Zhulidov et al. 2021). ...
... But the apparent gap in the current range of M. leucophaeata suggests the North lineage is better adapted to cold environments: after the intense ship traffic and oyster trade between the southern USA and NE USA during centuries (Kennedy 2011a), it would be expected that other COI haplotypes were present in NE USA if southern populations of M. leucophaeata were equally cold-tolerant. This tolerance of the North lineage to cold environments is also observed in the invasive populations in Europe; for example, Zhulidov et al. (2021) suggested that these bivalves face water temperatures down to 0 °C in the Ponto-Caspian region, whereas Van der Gaag et al. (2020) indicated a tolerance of settled specimens in Western Europe to at least 2.9 °C. Even though, severe winters may promote mass mortality on the invasive population in Europe (Schütz 1969). ...
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Mytilopsis leucophaeata is an estuarine bivalve native from the Gulf of Mexico and possibly part of the eastern USA, which invaded many regions around the globe. Despite the previous discovery that two genetic lineages of this species exist, one comprising the northeastern USA and Eurasia and another having invaded South America, there were no genetic data about specimens from native areas. We reveal for the first time the COI signatures of two native populations of M. leucophaeata, one in the eastern coast of Florida, another in its western coast, besides analyzing the nuclear gene 28S. The COI sequences from South America (Venezuela and Brazil) are similar to those from Florida, mainly to the population from the western coast, whereas the clade composed of sequences from NE USA and Eurasia diverges by 2.4–3.2% from remaining sequences. The 28S tree also confirmed the distinction of this lineage. We suspect that the cold-tolerant lineage may represent a distinct species originated in the mid-eastern or NE USA, possibly derived from glaciations or the formation of the Labrador Current, and having colonized Eurasia after human introductions. Based on data about ship trades, we also raise the hypothesis that the invasion in South America was composed of two stages, with an initial invasion in Venezuela or nearby Caribbean countries, followed by a secondary invasion in SE Brazil, although requiring further studies. These results may alter deeply our understanding about the invasion of M. leucophaeata around the globe.
... The native area of A. senhousia ranges from Siberia to Singapore (Chuang 1961), but it has been introduced in numerous localities worldwide over many decades such as in New-Zealand and Australia (Willan 1985(Willan , 1987, the USA (Crooks 1996(Crooks , 1998Wasson et al. 2001), India (Kutty et al. 2016) and Africa (Lourenço et al. 2018). Since 1989, it has been regularly reported from European coasts ( Figure 2): in the Mediterranean Sea (Hoenselaar and Hoenselaar 1989;Lazzari and Rinaldi 1994;Mistri et al. 2000;Mistri 2002;Mastrototaro et al. 2003;Zenetos et al. 2003;Doğan et al. 2014;Como et al. 2016;Scirocco and Urbano 2018); in the Black Sea (Kovalev et al. 2017;Chartosia et al. 2018;Zhulidov et al. 2021); in the Atlantic Ocean in the South of the Bay of Biscay (Bachelet et al. 2009); in UK in the Solent and particularly from Southampton Water estuary since 2007 (Barfield et al. 2018;Holman et al. 2019;Worsfold et al. 2020;Watson et al. 2021); and in the Netherlands (Faasse 2018). ...