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The Nonindigenous Occurrences section of the NAS species profiles has a new structure. The section is now dynamically updated from the NAS database to ensure that it contains the most current and accurate information. Occurrences are summarized in Table 1, alphabetically by state, with years of earliest and most recent observations, and the tally and names of drainages where the species was observed. The table contains hyperlinks to collections tables of specimens based on the states, years, and drainages selected. References to specimens that were not obtained through sighting reports and personal communications are found through the hyperlink in the Table 1 caption or through the individual specimens linked in the collections tables.

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Common name: Golden Mussel

Synonyms and Other Names: Limnoperna siamensis, Limnoperna lacustris

Identification: Limnoperna fortunei (common name: golden mussel) is a sessile, bivalve mollusk in the Mytilidae family with equivalve and heteromyarian shells. It is dark-brown above the umbonal keel and paler-yellow brown below. The golden mussel's common name is derived from the golden or yellowish-brown color of its shell. The shell’s inner surface has a purple mother-of-pearl layer above the keel and white below. 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 (Darrigran 2022).

 

 

Size: 20-30 mm, max 42-46 mm

Native Range: Limnoperna fortunei is a freshwater mytilid of South East Asia; native to the lakes and rivers of China, and also occurs naturally in Laos, Cambodia, Vietnam, Korea, Indonesia and Thailand (Ricciardi 1998).

Alaska	Hawaii	Puerto Rico & Virgin Islands	Guam Saipan

Hydrologic Unit Codes (HUCs) Explained
Interactive maps: Point Distribution Maps

Ecology:  

Habitat

The climate in Limnoperna fortunei’s native range is humid subtropical, with warm summers and no dry season (Darrigran and Damborenea 2005). L. fortunei is able to inhabit a variety of aquatic habitats (i.e. lakes, rivers, streams) but requires a hard substrate for byssal attachment, though it can attach to aquatic plants. However, it can also attach to soft substrates if it is sufficiently compacted (Boltovskoy et al. 2006). L. fortunei has a wide environmental tolerance: see the table below for a description of physiological tolerances.

Parameter Description	Numeric Value	Reference
Depth	< 10m	Karatayev et al. 2010
Temperature (Adult Survival)	5 - 35 °C	Ricciardi 1998 Oliveira et al. 2010
Temperature (Larval Development)	16 - 28 °C	Ricciardi 1998
Salinity	800 mOsm 0 - 12 ‰ 13.7 %	Deaton et al. 1989 Ricciardi 1998 Karatayev et al. 2007
pH	≥ 6.4	Ricciardi 1998
Calcium	≥ 3 mg/l	Ricciardi 1998 Karatayev et al. 2007

 

The mussel can survive (90%) up to a salinity shock of 2 ppt for periods of at least 10 days (Angonesi et al. 2008). This species overwinters in South Korea, with water temperatures as low as 0°C (Oliveira et al. 2010). In Japan, the minimum temperature in a reservoir with mussels was 4.2°C (Nakano et al. 2011). Experimental research supports the 5°C threshold for prolonged exposure (Oliveira et al. 2010). Experiments to examine L. fortunei overwintering survival found that populations at the northern invasion front can survive 6 days, 41 days, and 108 days at <1°C, <2°C, and <5°C respectively. Overall survival was 27% at these temperatures. An accompanying species distribution model implies suitable habitats at higher latitudes than previously considered (Xia et al. 2021). Compared to the zebra mussel Dreissena polymorpha, Limnoperna 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 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).

Food Web

Limnoperna fortunei consumes a variety of phytoplankton and zooplankton (Rojas Molina et al. 2010, Rojas Molina et al. 2012, Frau et al. 2012). Adults do well despite low food availability (Oliveira et al. 2011). This species negatively affects burrowing invertebrates and unionids (Karatayev et al. 2010) in South America, and may do the same in the Great Lakes given its high densities, such competitive exclusion is also seen in a similar species, the zebra mussel Dreissena polymorpha. L. fortunei modifies nutrient concentrations and proportions, and promotes aggregation of solitary Microcystis spp. cells into colonies; both these effects can favor blooms of this often noxious cyanobacteria (Cataldo et al. 2012).

Limnoperna fortunei negatively impacts zooplankton, and part of zooplankton decline may be due to starvation (i.e., mussel outcompetes zooplankton for food resources) (Rojas Molina et al. 2012). Limnoperna fortunei has increased food availability in the benthic zone, which in part has increased invertebrate (excluding mussels) density 1.9 to 22 times and biomass 1.7 to 19 times (Burlakova et al. 2012). Limnoperna fortunei has homogenized benthic communities (Darrigran and Damborenea 2011, Sardiña et al. 2011); e.g., in one study 99.9% of community biomass consisted of the filtering collector trophic group (Burlakova et al. 2012). Limnoperna fortunei has shunted the dominant nutrient cycling from the pelagic to the benthic zone (Darrigran and Damborenea 2011, Cataldo et al. 2012, Rojas Molina 2012). Limnoperna fortunei has significantly reduced phytoplankton densities (>60%) and changed composition of the algae assemblages, most notably an increase in the flagellate group, relative to the diatom, single-celled and colonial groups (Frau et al. 2012).

Limnoperna fortunei filters water substantially faster than D. polymorpha (Karatayev et al. 2010). Limnoperna fortunei brings increases in water transparency, and a decrease in suspended matter, chlorophyll a, and primary production (Boltovskoy et al. 2009). It also brings a decrease in turbidity and an increase in dissolved nitrogen 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 and Damborenea 2011, Burlakova et al. 2012, Spaccesi and Capitulo 2012). This species transforms sand or mostly bare sediment into reef-like druses (Burlakova et al. 2012).

Life History

Limnoperna fortunei requires external fertilization to reproduce and is considered a dioecious spawner with an equal ratio of males to females (Darrigran 2022). After fertilization, the oocyte develops into the first trochophore stage after six hours. This is the first active larval stage and at this point 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. External fertilization and the development of planktonic larvae support the rapid spread of golden mussel (Boltovskoy et al. 2015), similar to other successful invaders in the Great Lakes (i.e. Dreissena polymorpha and D. bugensis). The veliger larvae gradually transition into a plantigrade stage, at this stage movement becomes restricted and an adhesive foot is formed. The plantigrade stage involves exploration for suitable substrate and ends with the attachment of byssal threads. Once byssal attachment is complete, the larvae develop into juveniles and remain in place through adulthood (Boltovskoy 2015).

No data was found on natural Limnoperna fortunei fecundity, though very high rates of colonization suggest it is high (Karatayev et al. 2007). L. fortunei can reach densities of 5000-250,000 individuals/m² on hard substrate, and 90-2000 individuals/m² on soft 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). L. fortunei spawns continually in suitable conditions, as opposed to batch spawning that is observed in similar species (i.e. zebra mussel) (Boltovskoy et al. 2006).

Means of Introduction: 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). Spread of the species is likely through the freeswimming or drifting of plantonic larva through water, hitchhiking, or movement of contaminated water. 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 hitchking on boats and other materials that have been submerged in water long enough for veligers to settle and attach.

Impact of Introduction:

Summary of species impacts derived from literature review. Click on an icon to find out more...

Ecological	Economic	Human Health	Other

References: (click for full references)

2008. Invasive Species Compendium. CAB International. www.cabi.org. Accessed on 01/20/2017.

Angonesi, L.G., N.G. da Rosa, and C.E. Bemvenuti. 2008. Tolerance to salinities shocks of the invasive mussel Limnoperma fortunei under experimental conditions. Iheringia Serie Zoologia 98(1):66-69. http://www.scielo.br/scielo.php?script=sci_arttext&pid=S0073-47212008000100009&lng=en&tlng=en.

Boelman, S.F., F.M. Neilson, E.A. Dardeau, Jr., T. Cross. 1997. Zebra mussel (Dreissena polymorpha) control handbook for facility operators, first edition. Volume EL-97-1. US Army Corps of Engineers, Waterways Experiment Station, Vicksburg, MS .

Boltovskoy, D., N. Correa, D. Cataldo, and F. Sylvester. 2006. Dispersion and ecological impact of the invasive freshwater bivalve Limnoperna fortunei in the Río de la Plata watershed and beyond. Biological Invasions 8(4):947-963. http://link.springer.com/article/10.1007%2Fs10530-005-5107-z?LI=true.

Boltovskoy, D., F. Sylvester, A. Otaegui, V. Leites, and D.H. Cataldo. 2009. Environmental modulation of reproductive activity of the invasive mussel Limnoperna fortunei: implications for antifouling strategies. Austral Ecology 34(7):719-730. http://doi.wiley.com/10.1111/j.1442-9993.2009.01974.x.

Boltovskoy, D., A. Karatayev, L. Burlakova, D. Cataldo, V. Karatayev, F. Sylvester, and A. Mariñelarena. 2009. Significant ecosystem-wide effects of the swiftly spreading invasive freshwater bivalve Limnoperna fortunei. Hydrobiologia 636(1):271-284. dx.doi.org/10.1007/s10750-009-9956-9.

Bowers, R., and F.A. de Szalay. 2007. Fish predation of zebra mussels attached to Quadrula quadrula (Bivalvia: Unionidae) and benthic molluscs in a Great Lakes coastal wetland. Wetlands 27(1):203-208. http://www.bioone.org/doi/full/10.1672/0277-5212%282007%2927%5B203%3AFPOZMA %5D2.0.CO%3B2 BioOne.

Burlakova, L.E., A.Y. Karatayev, and V.A. Karatayev. 2012. Invasive mussels induce community changes by increasing habitat complexity. Hydrobiologia 685(1):121-134. dx.doi.org/10.1007/s10750-011-0791-4.

Carlsson, N., O. Sarnelle, and D.L. Strayer. 2009. Native predators and exotic prey: an acquired taste? Frontiers in Ecology and the Environment 7(10):525-532. http://www.jstor.org/stable/25595245.

Cataldo, D., D. Boltovskoy, J.L. Hermosa, and C. Canzi. 2005. Temperature-dependent rates of larval development in Limnoperna fortunei (Bivalvia : Mytilidae). Journal Molluscan Studies 71(1):41-46. dx.doi.org/10.1093/mollus/eyi005.

Cataldo, D., D. Boltovskoy, and M. Pose. 2003. Toxicity of chlorine and three nonoxidizing molluscicides to the pest mussel Limnoperna fortunei. Journal (American Water Works Association) 95(1):66-78. http://www.jstor.org/stable/41298216?seq=1#page_scan_tab_contents.

Cataldo, D., A. Vinocur, I. O'Farrell, E. Paolucci, V. Leites, and D. Boltovskoy. 2012. The introduced bivalve Limnoperna fortunei boosts Microcystis growth in Salto Grande reservoir (Argentina): evidence from mesocosm experiments . Hydrobiologia 680(1):25-38. http://link.springer.com/article/10.1007/s10750-011-0897-8.

Cataldo, D., I. O'Farrell, E. Paolucci, F. Sylvester, and D. Boltovskoy. 2012. Impact of the invasive golden mussel (Limnoperna fortunei) on phytoplankton and nutrient cycling. Aquatic Invasions 7(1):91-100. dx.doi.org/10.3391/ai.2012.7.1.010.

Darrigran, G.A., M.E. Maronas, and D.C. Colautti. 2004. Air exposure as a control mechanism for the Golden Mussel, Limnoperna fortunei, (Bivalvia: Mytilidae). Journal of Freshwater Ecology 19(3):461-464. http://www.tandfonline.com/doi/abs/10.1080/02705060.2004.9664920.

Darrigran, G., and C. Damborenea. 2011. Ecosystem engineering impact of Limnoperna fortunei in South America. Zoological Science 28(1):1-7. dx.doi.org/10.2108/zsj.28.1.

Darrigran, G.A., and M.C. Damborenea. 2005. A South American bioinvasion case history: Limnoperna fortunei (Dunker, 1857), the golden mussel. Amer. Malac. Bull. 20: 105-112.

Darrigran, G., S.M. Martin, B. Gullo, and L. Armendariz. 1998. Macroinvertebrates associated with Limnoperna fortunei (Dunker, 1857) (Bivalvia, Mytilidae) in Río de Plata, Argentina. Hydrobiologia 367(1):223-230. dx.doi.org/10.1023/A:1003244603854.

Darrigran, G.A., D.C. Colautti, and M.E. Maronas. 2007. A potential biocide for control of the Golden Mussel, Limnoperna fortunei. Journal of Freshwater Ecology 22(2):359-360. http://www.tandfonline.com/doi/abs/10.1080/02705060.2007.9665060.

Deaton, L.E., J.G.S. Derby, N. Subhedar, and M.J. Greenberg. 1989. Osmoregulation and salinity tolerance in two species of bivalve mollusc: Limnoperna fortunei and Mytilopsis leucophaeta. Journal of Experimental Marine Biology and Ecology 133(1-2):67-79. http://dx.doi.org/10.1016/0022-0981(89)90158-5.

Di Fiori, E., H. Pizarro, M. dos Santos Afonso, and D. Cataldo. 2012. Impact of the invasive mussel Limnoperna fortunei on glyphosate concentration in water . Ecotoxicology and Environmental Safety 81:106-113. http://dx.doi.org.proxy.lib.umich.edu/10.1016/j.ecoenv.2012.04.024.

Frau, D., F.R. Molina, M. Devercelli, and S. José de Paggi. 2012. The effect of an invading filter-feeding bivalve on a phytoplankton assemblage from the Paraná system: a mesocosm experiment. Marine and Freshwater Behaviour and Physiology 45(5):303-316. http://dx.doi.org.proxy.lib.umich.edu/10.1080/10236244.2012.735419.

Gazulha, V., M.C.D. Mansur, L.F. Cybis, and S.M.F.O. Azevedo. 2012. Grazing impacts of the invasive bivalve Limnoperna fortunei (Dunker, 1857) on single-celled, colonial and filamentous cyanobacteria. Brazilian Journal of Biology 72(1):33-39. http://dx.doi.org/10.1590/S1519-69842012000100004.

GLMRIS. 2012. Appendix C: Inventory of available controls for aquatic nuisance species of concern, Chicago area waterway system. U.S. Army Corps of Engineers.

Goto, Y. 2002. Behavior of nuisance mussel, Limnoperna fortunei, in water supply facilities. Water Science and Technology 46(11-12):45-50. http://wst.iwaponline.com.proxy.lib.umich.edu/content/46/11-12/45.

Isaac, A., A. Fernandes, M.J.M. Ganassin, and N.S. Hahn. 2014. Three invasive species occurring in the diets of fishes in a neotropical floodplain. Brazilian Journal of Biology 74(3):16-22. http://dx.doi.org/10.1590/1519-6984.18312.

Karatayev, A.Y., D. Boltovskoy, D.K. Padilla, and L.E. Burlakova. 2007. The invasive bivalves Dreissena polymorpha and Limnoperna fortunei: Parallels, contrasts, potential spread and invasion impacts. Journal of Shellfish Research 26(1):205-213. http://dx.doi.org/10.2983/0730-8000(2007)26[205:TIBDPA]2.0.CO;2.

Karatayev, A.Y., L.E. Burlakova, V.A. Karatayev, and D. Boltovskoy. 2010. Limnoperna fortunei versus Dreissena polymorpha: population densities and benthic community impacts of two invasive freshwater bivalves. Journal of Shellfish Research 29(4):975-984. http://dx.doi.org/10.2983/035.029.0432.

Nakano, D., T. Kobayashi, N. Endo, and I. Sakaguchi. 2011. Growth rate and settlement of Limnoperna fortunei in a temperature reservoir. Journal of Molluscan Studies 77(2):142-148. https://doi-org.proxy.lib.umich.edu/10.1093/mollus/eyq048.

Oliveira, C.R.C., R. Fugi, K.P. Brancalhao, and A.A. Agostinho. 2010. Fish as Potential Controllers of Invasive Mollusks in a Neotropical Reservoir. Brazilian Journal of Nature Conservation 8(2):140-144. dx.doi.org/10.4322/natcon.00802006.

Oliveira, M.D., S.K. Hamilton, and C.M. Jacobi. 2009. Forecasting the expansion of the invasive golden mussel Limnoperna fortunei in Brazilian and North American rivers based on its occurrence in the Paraguay River and Pantanal wetland of Brazil. Aquatic Invasions 5(1):59-73. dx.doi.org/10.3391/ai.2010.5.1.8.

Oliveira, M.D., D.F. Calheiros, C.M. Jacobi, and S.K. Hamilton. 2011. Abiotic factors controlling the establishment and abundance of the invasive golden mussel Limnoperna fortunei. Biological Invasions 13(3):717-729. dx.doi.org/10.1007/s10530-010-9862-0.

Paoulucci, E.M., E.V. Thuesen, D.H. Cataldo, and D. Boltovskoy. 2010. Veligers of an introduced bivalve, Limnoperna fortunei, are a new food resource that enhances growth of larval fish in the Paraná River (South America). Freshwater Biology 55(9):1831-1844. http://onlinelibrary.wiley.com/doi/10.1111/j.1365-2427.2010.02418.x/full.

Molina, F.R., S.J. de Paggi, and D. Frau. 2012. Impacts of the invading golden mussel Limnoperna fortunei on zooplankton: a mesocosm experiment. Zoological Studies 51(6):733-744.

Molina, F.R., J.C. Paggi, and M. Devercelli. 2010. Zooplanktophagy in the natural diet and selectivity of the invasive mollusk Limnoperna fortunei. Biological Invasions 12(6):1647-1659. dx.doi.org/10.1007/s10530-009-9578-1.

Rackl S., M. Koivunen, P. Marrone, H. Huang, and R. Asolkar. (2012). Agents for the control of Limnoperna sp. United States Patent and Trademark Office (Pub. No. 20120121745A1)

Ricciardi, A. 1998. Global range expansion of the Asian mussel Limnoperna fortunei (Mytilidae): another fouling threat to freshwater systems. Biofouling 13(2):97-106.

Sardiña, P., E. Chaves, and M. Marchese. 2011. Benthic community responses to invasion by the golden mussel, Limnoperna fortunei Dunker: biotic homogenization vs environmental driving forces. Journal of the North American Benthological Society 30(4):1009-1023. http://www.journals.uchicago.edu.proxy.lib.umich.edu/doi/10.1899/10-170.1.

Spaccesi, F.G., and A.R. Capitulo. 2012. Benthic communities on hard substrates covered by Limnoperna fortunei Dunker (Bivalvia, Mytilidae) at an estuarine beach (Río de la Plata, Argentina). Journal of Limnology 71(1):144-153. http://www.jlimnol.it/index.php/jlimnol/article/view/jlimnol.2012.e15.

Sylvester, F., D.H. Cataldo, C. Notaro, and D. Boltovskoy. 2013. Fluctuating salinity improves survival of the invasive freshwater golden mussel at high salinity: implications for the introduction of aquatic species through estuarine ports. Biological Invasions 15(6):1355-1366. http://link.springer.com/article/10.1007/s10530-012-0373-z.

Xu, M., G. Darrigran, Z. Wang, N. Zhao, C.C. Lin, and B. Pan. 2015. Experimental study on control of Limnoperna fortunei biofouling in water transfer tunnels. Journal of Hydro-environment Research 9(2):248-258. dx.doi.org/10.1016/j.jher.2014.06.006.

Other Resources:
CABI Compendium Datasheet for Limnoperna fortunei

US Fish and Wildlife Service Ecological Risk Screening Summary for Limnoperna fortunei

Author: Fusaro, A., A. Davidson, K. Alame, M. Gappy, E. Baker, G. Nunez, J. Larson, W. Conard, P. Alsip, C. Shelly, and Cayla Morningstar

This information is preliminary or provisional and is subject to revision. It is being provided to meet the need for timely best science. The information has not received final approval by the U.S. Geological Survey (USGS) and is provided on the condition that neither the USGS nor the U.S. Government shall be held liable for any damages resulting from the authorized or unauthorized use of the information.