Invasion History

First Non-native North American Tidal Record: 2024
First Non-native West Coast Tidal Record: 2024
First Non-native East/Gulf Coast Tidal Record:

General Invasion History:

Limnoperna fortunei is native to freshwater bodies within and south of the Yangtze River and within the Pearl River Basin, China (Morton and Dineson 2010, Zhang et al. 2022).  Its distribution expanded to Hong Kong in 1965, and then to Japan and Taiwan in the early 1990s (Morton 1975, Ricciardi 1998). This mussel had been widely introduced through ballast water contamination of veliger (larval) mussels in South America from 1991 (Pastorino et al. 1993, Darrigran & Damborenea 2005) and was found in freshwater ecosystems in Argentina, Brazil, Paraguay, Uruguay and Bolivia between 1991 and 1999; from Bagliardi Beach, Río de la Plata estuary Argentina (in 1991), and then along the Paraguay–Parana waterway into Brazil (Pastorino et al. 1993,  Darrigran 2002, Oliviera et al. 2006). In 2024 it was found in San Francisco Bay-Delta Estuary in California, the first known occurrence in North America (CDFW 2024).

North American Invasion History:

Invasion History on the West Coast:

Limnoperna fortunei was first reported from North American waters near Stockton, CA in October 2024, and is the first known occurrence in North America (CDFW 2024). It was discovered during a routine maintenance inspection of water quality monitoring equipment by the California Department of Water Resources. Since its discovery, it has been detected from surveys by the California Department of Fish and Wildlife (CDFW) invasive mussel early detection monitoring program in multiple locations in the Sacramento-San Joaquin River Delta, part of the larger San Francisco Bay Delta Estuary (USFWS 2025). As of March 2025, it has been verified as far west as the Contra Costa Canal near the Contra Loma Reservoir, and as far east as the San Joaquin river within the Stockton port complex. To the north, it has been verified in Three Mile Slough, and to the south it has been verified in at least two locations: the California Aquaduct just outside the San Luis Reservoir State Recreation Area; and the Coastal Aquaduct, an offshoot of the California Aquaduct, near Boulder Hill, about 165 miles south of its initial discovery in Stockton (USFWS 2025). The mussels that have been surveyed and collected are a mixture of juveniles and adults, and settle in clumps on a variety of hard and artificial substrates and samplers (USFWS 2025). Most recently in May and June 2025, Golden mussel veliger larvae were detected by light microscopy and PCR analysis in Quail Lake Los Angeles County, part of the State Water Project's West Branch canal network, the southernmost detection at the time and downstream from previous detections (CDFW, 2025). All stages of L. fortunei development have now been found in multiple locations within California's network of canals and aqueducts (CDFW 2024, 2025). Commercial shipping is a likely vector for its introduction to California, and as the port of Stockton is connected by natural rivers as well as a network of man-made canals and aqueducts, the golden mussel has multiple pathways to spread throughout the region. Human-mediated spread on recreational boat hulls and trailers overland and within water is also a critical means of distribution to new locations. The US Fish and Wildlife Service describe L. fortunei as a 'high risk' invasive species due to its climate affinity in the United States and economic and ecological impacts in areas it has currently invaded, including fouling and clogging aquatic infrastructure, and diet alteration and displacement of native species (USFWS 2024). 

 

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Invasion History Elsewhere in the World:

Limnoperna fortunei native range lies within and south of the Yangtze River and within the Pearl River Basin, China (Morton and Dineson 2010, Zhang et al. 2022) although it has spread into inland northern China through water transfer projects (Xu et al. 2015, Zhang et al. 2022). Its presence in Southeast Asia such as in Laos, Cambodia, Thailand and Vietnam is probably the result of historical human migrations (Morton & Dinesen 2010). Between 1965 and 1990, it spread into Hong Kong, Taiwan and Japan (Ricciardi 1998, Ito 2015). Around 1990 it appeared in South America, Argentina (Pastorino et al. 1993). By 2006, it had spread to Uruguay, Paraguay, Bolivia, and Brazil. In 2017, in South America it was present in two major basins (Río de La Plata, including the Paraguay-Paraná and Uruguay rivers, and the São Francisco basin), as well as several smaller watersheds (Mar Chiquita, Guaíba, Patos-Mirim, Tramandaí) (Oliveira et al. 2015, Barbosa et al. 2016). Introduction of the species into Japan and South America is thought to be from ballast water contamination of veliger (larval) mussels (Darrigran and Damborenea 2005). In the areas colonized, Limnoperna can reach very high abundances, often exceeding 200,000 ind. m-2 (Correa et al. 2015) and has become the sole species colonizing hard substrates in the Río de La Plata watershed (Boltovskoy et al. 2006). Spread of the species is likely through the free swimming or drifting of planktonic larva through water, hitchhiking, or movement of contaminated water in ships, or adults on the hulls of vessels (Boltovskoy et al. 2006). Like dreissenid mussels, this species has microscopic, planktonic larvae that are difficult to detect and can be transported in water accidentally. Adult mussels biofoul surfaces, attaching with byssal threads, which facilitates potential hitchhiking on boats and other materials that have been submerged in water long enough for veligers to settle and attach.


Description

Limnoperna fortunei commonly known as the golden mussel, is a small, sessile freshwater bivalve mollusk of the Mytilidae family. The golden mussel's common name is derived from the golden or yellowish-brown color of its shell. It has equivalve and heteromyarian shells. The shell is dark-brown above the umbonal keel and paler-yellow brown below. The shell’s inner surface has a purple mother-of-pearl layer above the keel and white below. This nacreous layer distinguishes it from members of the superfamily Dreissenacea, which includes zebra and quagga mussels. The shell’s outer periostracal layer is smooth and shiny, and thick where it curls inwards at the shell margin. The ventral margin of the shell can vary between straight or curved among individual specimens. It has subterminal umbones, smooth and glossy periostracum and a simple hinge ligament. It is a relatively unspecialized mytilid in terms of its shell architecture, lacking external ornamentation, internal teeth and crenulations, but it has internal specializations for greater efficiency in food collection and utilization (description from Darrigran 2022). 

After fertilization, L. fortunei larvae undergo a free swimming planktonic stage, aiding their dispersal, followed by a pedi-veliger and plantigrade stage, where the larvae settle out of the water column onto a substrate and lose their velum (swimming and feeding structure), to finally metamorphose into a juvenile dissoconch stage, when the final adult shell forms from the juvenile one, and secretes byssal threads that anchor them to the substrate (Zhang et al. 2022). Adult average shell length is around 20–30 mm though maximum shell length reaches up to 45 mm (Zhang et. al 2022). Adult L. fortunei can form very dense clusters on rigid substrates and even colonize semi-rigid substrates such as macrophytes at lower densities (Frau et al. 2012). 

 

 

 

 


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Mollusca
Class:   Bivalvia
Subclass:   Pteriomorphia
Order:   Mytiloida
Family:   Mytilidae
Genus:   Limnoperna
Species:   fortunei

Synonyms

Dreissena siamensis (Morelet, 1866)
Limnoperna coreana (Park & Choi, 2008)
Limnoperna depressa (Brandt & Temcharoen, 1971)
Limnoperna lacustris (E.von Martens, 1875)
Limnoperna lemeslei (Rochebrune, 1882)
Limnoperna supoti ( Brandt, 1974)
Modiola cambodgensis ( Clessin, 1888)
Modiola cambodjensis ( Clessin, 1889)
Modiola lacustris (Martens, 1875)
Mytilus martensi (Neumayer, 1898)
Volsella fortunei ( Dunker, 1857)

Potentially Misidentified Species

Corbicula

The morphological similarity between the larvae of the golden mussel and other bivalve molluscs, particularly those of the Corbicula genus, can pose challenges for accurate identification, even for experts (Darrigran and Damborenea 2009, de Paula et al. 2020).



Limnoperna fortunei kikuchii

Limnoperna fortunei kikuchii turns out to not be an Limnoperna fortunei at all. The wide morphological range of Limnopernae contributed to this confusion (Ricciardi 1998).



Mytella charruana

Mytella charruana is a brackish mytilid, native to the eastern coast of South America from Venezuela to Argentina, and in the Pacific from Mexico to El Salvador, and invasive in southeastern USA, Philippines, Singapore, Thailand, and India (Calazans et al., 2017, Boltovskoy et al. 2022). It only occasionally co-occurs with the golden mussel in estuarine conditions with frequent and strong salinity changes (Giberto & Sardiña, 2009).



Xenostrobus securis (Lamarck, 1819)

The Australian mussel Xenostrobus securis was initially misidentified and given this name in Japan in the 1970s. The wide morphological range of Limnopernae contributed to this confusion (Ricciardi 1998). 



Ecology

General:

Limnoperna fortunei is an effective ecosystem engineer, altering both structure and function of the ecosystems invaded, and a very successful opportunistic species. It has a wide environmental tolerance. In its native range, the climate is humid subtropical, with warm summers and no dry season (Darrigran & Damborenea 2005). It is found in many freshwater aquatic bodies such as lakes, rivers, streams) but like all mussels it requires a rigid substrate for byssal attachment. Its beds can have an extremely patchy distribution. It can survive up to 10 days with a salinity shock of 2 ppt (Angonesi et al. 2008) and has been found in tidal and slightly estuarine waters in China and Cambodia, with salinities < 1–2% ppt (Morton & Dinesen, 2010). 5°C has shown to be a threshold for the golden mussel for prolonged exposure in the lab experiments (de Oliveira et al. 2010) and it has not been reported from waterbodies where year-round temperatures are below 15–18 °C (Karatayev et al. 2015) but habitats suitability studies have predicted a higher latitudes distribution than previously considered (Xia et al. 2021). Compared to another successful invader, the zebra mussel Dreissena polymorpha, L. fortunei has higher resistance to anoxia, pollution (including eutrophication), pH, and high temperatures, longer reproduction periods and lower calcium requirements (3–4mg/L) (Karatayev et al. 2007). This broader tolerance indicates this species could have an even broader distribution in the Great Lakes if it were to be introduced there than D. polymorpha, except for depth as zebra mussels can inhabit depths up to 50 meters while L. fortunei has been noted to inhabit depths of 0.5 to 40 meters with an optimum depth of 10 meters (Darrigran 2022). It can alter the associated invertebrate community but also the properties of the water column by increasing the water transparency, decreasing the phytoplankton and zooplankton densities and enhancing macrophytes (Boltovskoy et al. 2009). Limnoperna fortunei is a suspension feeder and can continuously reproduce for 8–9 months over the year especially where temperature do not drop under 15°C during the winter (Boltovskoy & Cataldo 1999, Cataldo & Boltovskoy 2000). Even in the introduced area, the golden mussel, at every stage, has become a common prey in the diet of the local fish (Boltovskoy et al. 2006, Cataldo 2015b).

Food:

organic seston including: detritus, bacterioplankton, phytoplankton, and zooplankton

Consumers:

Fishes (the boga, Megaleporidens obtusidens)

Competitors:

Trophic Status:

Suspension feeder

Habitats

General HabitatNontidal FreshwaterNone
Salinity RangeOligohaline0.5-5 PSU
Salinity RangeLimnetic0-0.5 PSU
Salinity RangeMesohaline5-18 PSU

Life History

Limnoperna fortunei is a dioecious spawner with external fertilization (Darrigran 2022). Its life span is around 2–3 years (Morton 1977, Boltovskoy & Cataldo 1999). Studies from Asia and South America show that reproduction starts when water temperatures reach around 15–18 °C (Morton 1977, Choi & Shin 1985, Cataldo & Boltovskoy 2000, Nakano et al. 2010a, Brugnoli et al. 2011). The oocyte develops into the first trochophore stage after six hours. These larvae are capable of coordinated swimming and disperse in the water column. Larvae then develop into the veliger stages where the shell begins to form, and they begin to consume plankton. Free-swimming planktonic larvae, typical for most marine benthic animals (Thorson 1961, Pechenik 1999), are rare among freshwater benthic invertebrates (Moss,1988) and it has probably evolved from a recent marine ancestor. External fertilization and the development of planktonic larvae are the key for their rapid spread (Boltovskoy et al. 2015). The veliger larvae gradually transition into a plantigrade stage, when movement becomes restricted and an adhesive foot is formed and ends with the attachment of byssal threads. The larvae take 10–20 days to reach the settling stage (Cataldo et al. 2005). Once byssal attachment is complete, the larvae develop into juveniles and remain in place through adulthood (Boltovskoy 2015). Mean growth rates were in the range 0.4–3.5 mm/month. Sexual maturation occurs when the shell length reach 5–8 mm.

Limnoperna fortunei can reach densities of 5000–250,000 individuals/m2 on hard substrate, and 90–2000 individuals/m2 on softer substrate (Frau et al. 2012). Analysis of a L. fortunei population in Brazil found a high annual growth rate (K = 1.22) and estimated that 62.920 juveniles/m2 will be recruited annually (Ayroza et al. 2021). Limnoperna fortunei spawns continually in suitable warmer conditions (Boltovskoy et al. 2006). In China L. fortunei can produce three generations per year from March to November, each of which was associated with two reproductive peaks, resulting in six cohorts per year (Xu et al. 2015). Cyanobacterial blooms can suppress the reproduction as their toxins (microcystin) engender massive larval mortalities (Boltovskoy et al. 2013).


Tolerances and Life History Parameters

Minimum Temperature (ºC)5

Oliveira et al. (2010) found that the golden mussel reached 100% mortality after 38 days at 5–7 °C, and suggesting that 5 °C was a critical lower threshold for extended survival in winter.
Other authors found that it can survive at least 108 days in water < 5  °C (Ricciardi 1998, Oliveira et al. 2011, Xia et al. 2021)

Maximum Temperature (ºC)35

(Ricciardi 1998, Oliveira et al. 2011). At 34–36 °C, the total time of mortality was 25.0–644.3 h, from 38 to 43 °C, all mussels die after 0.7–17.5 h (Perepelizin & Bltovskoy 2011).
 

Minimum Salinity (‰)0

Mainly freshwater but can survive in brackish (Boltovskoy et al. 2015)

Maximum Salinity (‰)13.7

Golden mussels can tolerate short periods (hours) of salinities up to 23 ‰ without significant mortality (Deaton et al. 1989, Ricciardi 1998, Karatayev et al. 2007, Sylvester et al. 2013)

Minimum Dissolved Oxygen (mg/l)0.2

Below 0.16 mg/L DO, at 20oC 100% sample mortality (SM 100) of small mussels was attained in 20.7 days (LT50, lethal time to kill 50% of the mussels = 9.5 days) and large mussels in 29.3 days (LT50 = 18.0 days). At 27oC, small and large mussels achieved SM 100 much faster (11.6 and 10.2 days) and had correspondingly lower LT50 values (4.7 days for both sizes) (Perepelizin & Boltovskoy, 2011).

Maximum Duration10

Longevity can be Variable depending on the location: 2–5 years, even over 10 years in China (Zhang et al. 2022)
 

Growth RateNone

0.40-3.50 mm/month (Boltovskoy et al. 2015). 20 mm in shell size in a year (Goto 2002).
 

Minimum Settling Velocity10

Settlement velocity in days (Cataldo et al. 2005)

Maximum Settling Velocity20

Settlement velocity in days (Cataldo et al. 2005)

Maximum Length (mm)45

shell length of adult. The more usual size ranges from 20 to 30 mm (Morton 2015)
 

General Impacts

The general impact of the golden mussel can have both negative and positive impacts depending on the environmental factors of the new introduced area, so the risk analysis or the possible damage of its introduction must be carefully evaluated on a case-by-case basis (Burlakova et al. 2023). In 1998, Ricciardi identified Limnoperna fortunei among several biofouling pests that should be high quarantine priorities around the world (Ricciardi 1998). The Brazilian Institute of the Environment and Renewable Natural Resources (IBAMA) has classified L. fortunei as one of the top three priority invasive species for control, as its overpopulation can cause severe issues for native fauna and national ecosystems (IBAMA 2020, Ostrensky et al. 2025).

This species negatively affects burrowing invertebrates and unionids (Karatayev et al. 2010) in South America. Limnoperna fortunei modifies nutrient concentrations and proportions and promotes aggregation of solitary Microcystis spp. cells into colonies; both these effects can favor blooms of noxious cyanobacteria (Boltovskoy et al. 2009, Cataldo et al. 2012, Silva & Giani 2018).

Limnoperna fortunei can increase water transparency and decrease suspended matter, chlorophyll a, and primary production (Boltovskoy et al. 2009).Turbidity can decrease and  dissolved nitrogen can increase in mussel presence (Rojas Molina 2012). Increased habitat complexity led to significant (e.g., threefold) increase in community taxonomic richness. Shells increase surface area for settling organisms, and also provide refuges from predation and physical stressors (Darrigran et al. 1998, Darrigran & Damborenea 2011, Burlakova et al. 2012, Spaccesi & Capitulo 2012). This species transforms sand or mostly bare sediment into reef-like druses (Burlakova et al. 2012) or they can settle on and cause damage to manmade infrastructures, such as concrete and metal (Magara et al. 2001, Yao et al. 2017, Xiao et al. 2025).

As a biofouler, it causes major problems for industrial components, water treatments and power plants due to clogging and efficiency loss (Boltovskoy & Cataldo 1999, Zhang 2022). Mussels have been observed colonizing water transfer structures at over 10 cm in thickness (Xu et al, 2015) and have been the cause of massive fouling problems across China, Japan, Argentina, Paraguay, Uruguay, and Brazil (Zhang et al. 2022).  Mass attachment by L. fortunei resulting in clogged pipelines and corrosion at one main hydro-electric power plant on the São Francisco River is estimated to cost 700,000.00 USD a year for maintenance and cleaning (Uliano-Silva et al. 2017). Mussel die-offs inside and around structures can result in further water quality degradation (Xu et al. 2015).

L. fortunei can be temporarily dislodged through oxygen deprivation (Perepelizin & Boltovskoy 2015) and thermal treatment (Perepelizin & Boltovskoy 2011). Sodium hypochlorite can dissolve the byssus and clay or silt precipitation is recommended as a potential method for preventing the attachment of golden mussel to materials (Xu et al. 2015, Zhang et al. 2022). Heat treatment and dessication are other practical alternatives for efficient control in fouled systems (Montalto & Ezcurra de Drago 2003, Perepelizin & Boltovskoy 2011).


Regional Distribution Map


  Non-native  
  Native  
  Cryptogenic  
  Failed  
Leaflet | Tiles © Esri — Sources: GEBCO, NOAA, CHS, OSU, UNH, CSUMB, National Geographic, DeLorme, NAVTEQ, and Esri

Occurrence Map

Leaflet | Tiles © Esri — Source: Esri, i-cubed, USDA, USGS, AEX, GeoEye, Getmapping, Aerogrid, IGN, IGP, UPR-EGP, and the GIS User Community

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