Quagga Mussel
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The quagga mussel (''Dreissena rostriformis'', also known as ''Dreissena bugensis'' or ''Dreissena rostriformis bugensis'') is a species (or subspecies) of
freshwater mussel Freshwater bivalves are one kind of freshwater mollusc, along with freshwater snails. They are bivalves that live in fresh water as opposed to salt water, which is the main habitat type for bivalves. The majority of species of bivalve molluscs ...
, an aquatic bivalve
mollusk Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is e ...
in the family
Dreissenidae The Dreissenidae are a family of small freshwater mussels, aquatic bivalve molluscs. They attach themselves to stones or to any other hard surface using a byssus. The shells of these bivalves are shaped somewhat like those of true mussels, and ...
. It has an average lifespan of 3 to 5 years. The species is indigenous to the
Dnieper River } The Dnieper () or Dnipro (); , ; . is one of the major transboundary rivers of Europe, rising in the Valdai Hills near Smolensk, Russia, before flowing through Belarus and Ukraine to the Black Sea. It is the longest river of Ukraine an ...
drainage of
Ukraine Ukraine ( uk, Україна, Ukraïna, ) is a country in Eastern Europe. It is the second-largest European country after Russia, which it borders to the east and northeast. Ukraine covers approximately . Prior to the ongoing Russian inv ...
, and is named after the
quagga The quagga ( or ) (''Equus quagga quagga'') is a subspecies of the plains zebra that was endemic to South Africa until it was hunted to extinction in the late 19th century. It was long thought to be a distinct species, but early genetic ...
, an extinct subspecies of African
zebra Zebras (, ) (subgenus ''Hippotigris'') are African equines with distinctive black-and-white striped coats. There are three living species: the Grévy's zebra (''Equus grevyi''), plains zebra (''E. quagga''), and the mountain zebra (''E. zebr ...
, possibly because, like the quagga, its stripes fade out towards the ventral side. The invasive quagga mussel is currently of major concern as it spreads in the rivers and lakes of Europe and also in the
Great Lakes The Great Lakes, also called the Great Lakes of North America, are a series of large interconnected freshwater lakes in the mid-east region of North America that connect to the Atlantic Ocean via the Saint Lawrence River. There are five lak ...
of North America where it was brought to by overseas shippers that use the
Saint Lawrence Seaway The St. Lawrence Seaway (french: la Voie Maritime du Saint-Laurent) is a system of locks, canals, and channels in Canada and the United States that permits oceangoing vessels to travel from the Atlantic Ocean to the Great Lakes of North Ameri ...
.


Appearance

The quagga mussel shell is generally black, yellow, and/or zig-zagged. However, a large range of shell morphologies is seen, including a distinct morph in
Lake Erie Lake Erie ( "eerie") is the fourth largest lake by surface area of the five Great Lakes in North America and the eleventh-largest globally. It is the southernmost, shallowest, and smallest by volume of the Great Lakes and therefore also h ...
that is pale or completely white. The shell has a rounded
carina Carina may refer to: Places Australia * Carina, Queensland, a suburb in Brisbane * Carina Heights, Queensland, a suburb in Brisbane * Carina, Victoria, a locality in Mildura Serbia * Carina, Osečina, a village in the Kolubara District ...
and a convex ventral side. The quagga mussel resembles the
zebra mussel The zebra mussel (''Dreissena polymorpha'') is a small freshwater mussel. The species originates from the lakes of southern Russia and Ukraine, but has been accidentally introduced to numerous other areas and has become an invasive species in ma ...
, just as its namesake (quagga) resembles the zebra. The quagga mussel shell can be distinguished from the zebra mussel shell because it is paler toward the end of the hinge. It is also slightly larger than the zebra mussel, about wide, roughly about the size of an adult human's thumbnail.


Diet

The quagga mussel is a filter feeder; it uses its cilia to pull water into its shell cavity through an incurrent siphon, where the desirable particulate matter is removed. Each adult mussel is capable of filtering one liter or more of water each day, where they remove phytoplankton,
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
, algae, and even their own
veliger A veliger is the planktonic larva of many kinds of sea snails and freshwater snails, as well as most bivalve molluscs (clams) and tusk shells. Description The veliger is the characteristic larva of the gastropod, bivalve and scaphopod ...
s. Any undesirable particulate matter is bound with
mucus Mucus ( ) is a slippery aqueous secretion produced by, and covering, mucous membranes. It is typically produced from cells found in mucous glands, although it may also originate from mixed glands, which contain both serous and mucous cells. It ...
, known as
pseudofeces Pseudofeces or pseudofaeces are a specialized method of expulsion that filter-feeding bivalve mollusks (and filter-feeding gastropod mollusks) use in order to get rid of suspended particles such as particles of grit which cannot be used as food, ...
, and ejected out the incurrent siphon. The particle-free water is then discharged out the excurrent siphon.


Reproduction

The quagga mussel is a prolific breeder, possibly contributing to its spread and abundance. ''Dreissena'' spp. are dioecious (either male or female) with external fertilization. A fully mature female mussel is capable of producing up to one million eggs per year. After fertilization, pelagic microscopic larvae, or veligers, develop within a few days and these veligers soon acquire minute bivalve shells. Free-swimming veligers drift with the currents for 3 to 4 weeks feeding by their hair-like cilia while trying to locate suitable substrata to settle and secure
byssal thread A byssus () is a bundle of filaments secreted by many species of bivalve mollusc that function to attach the mollusc to a solid surface. Species from several families of clams have a byssus, including pen shells ( Pinnidae), true mussels (Mytili ...
s. Mortality in this transitional stage from planktonic veliger to settled juvenile may exceed 99%. In 2019, the genome of a quagga mussel from the
Danube River The Danube ( ; ) is a river that was once a long-standing frontier of the Roman Empire and today connects 10 European countries, running through their territories or being a border. Originating in Germany, the Danube flows southeast for , pa ...
in Austria was sequenced, revealing how larvae use a system of intercellular 'cleavage cavities' and an expanded set of
aquaporin Aquaporins, also called water channels, are channel proteins from a larger family of major intrinsic proteins that form pores in the membrane of biological cells, mainly facilitating transport of water between cells. The cell membranes of a ...
transmembrane water channels for
osmoregulation Osmoregulation is the active regulation of the osmotic pressure of an organism's body fluids, detected by osmoreceptors, to maintain the homeostasis of the organism's water content; that is, it maintains the fluid balance and the concentration o ...
in low-salinity freshwater environments during the early stages of their development.


Invasive species


Europe

Quagga mussels are presumed to have originated in the Ukrainian section of the
Black Sea The Black Sea is a marginal mediterranean sea of the Atlantic Ocean lying between Europe and Asia, east of the Balkans, south of the East European Plain, west of the Caucasus, and north of Anatolia. It is bounded by Bulgaria, Georgia, Rom ...
and probably began to spread further into eastern Europe in the 1940s. Today, they are an invasive species found throughout western Europe. In Romania, quagga mussels were first found in 2004 in the Danube River. In
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
, quagga mussels were first identified in 2005, and now populate many inland waters, such as the
Rhine–Main–Danube Canal The Rhine–Main–Danube Canal (German: ''Rhein-Main-Donau-Kanal''; also called Main-Danube Canal, RMD Canal or Europa Canal), is a canal in Bavaria, Germany. Connecting the Main and the Danube riv ...
, the
Main Main may refer to: Geography * Main River (disambiguation) **Most commonly the Main (river) in Germany * Main, Iran, a village in Fars Province *"Spanish Main", the Caribbean coasts of mainland Spanish territories in the 16th and 17th centuries ...
, and the
Rhine ), Surselva, Graubünden, Switzerland , source1_coordinates= , source1_elevation = , source2 = Rein Posteriur/Hinterrhein , source2_location = Paradies Glacier, Graubünden, Switzerland , source2_coordinates= , so ...
. They were first identified in Switzerland in 2015, and in Lake Constance in 2016, where they have since spread massively and caused considerable problems, in particular to the machinery in
waterworks Water supply is the provision of water by public utilities, commercial organisations, community endeavors or by individuals, usually via a system of pumps and pipes. Public water supply systems are crucial to properly functioning societies. Thes ...
. In 2014, the species was reported at
Wraysbury Reservoir Wraysbury Reservoir is a water supply reservoir for London, just west of the M25 near the village of Wraysbury, and directly under the western approach path of Heathrow Airport. The reservoir was begun in 1967 and completed by W. & C. French in ...
, not far from London's Heathrow Airport in the valley of the
River Thames The River Thames ( ), known alternatively in parts as the River Isis, is a river that flows through southern England including London. At , it is the longest river entirely in England and the second-longest in the United Kingdom, after the R ...
. In
Ireland Ireland ( ; ga, Éire ; Ulster Scots dialect, Ulster-Scots: ) is an island in the Atlantic Ocean, North Atlantic Ocean, in Northwestern Europe, north-western Europe. It is separated from Great Britain to its east by the North Channel (Grea ...
, the mussels were first discovered in 2021 in two lakes on the River Shannon: Lough Derg and
Lough Ree Lough Ree () is a lake in the midlands of Ireland, the second of the three major lakes on the River Shannon. Lough Ree is the second largest lake on the Shannon after Lough Derg. The other two major lakes are Lough Allen to the north, and Lou ...
. The mussel is expected to compete with existing zebra mussels and native species, becoming widespread due to its ecological tolerance and suitability to Irish climatic conditions.


North America

Zebra mussels, the first dreissenid mussel introduced in North America, rapidly spread throughout many major river systems and the
Great Lakes The Great Lakes, also called the Great Lakes of North America, are a series of large interconnected freshwater lakes in the mid-east region of North America that connect to the Atlantic Ocean via the Saint Lawrence River. There are five lak ...
, causing substantial ecological and environmental impacts. The quagga mussel was first observed in North America in September 1989, when it was discovered in
Lake Erie Lake Erie ( "eerie") is the fourth largest lake by surface area of the five Great Lakes in North America and the eleventh-largest globally. It is the southernmost, shallowest, and smallest by volume of the Great Lakes and therefore also h ...
near
Port Colborne, Ontario Port Colborne is a city in Ontario, Canada that is located on Lake Erie, at the southern end of the Welland Canal, in the Niagara Region of Southern Ontario. The original settlement, known as Gravelly Bay, dates from 1832 and was renamed after ...
. It was not identified as a distinct species until 1991. By 2021 the biomass of quaggas in the lower 4 Great Lakes was estimated to be so great as to become the primary regulator of
phosphorus Phosphorus is a chemical element with the symbol P and atomic number 15. Elemental phosphorus exists in two major forms, white phosphorus and red phosphorus, but because it is highly reactive, phosphorus is never found as a free element on Ear ...
, remetabolizing and recirculating it instead of allowing it to
sediment Sediment is a naturally occurring material that is broken down by processes of weathering and erosion, and is subsequently transported by the action of wind, water, or ice or by the force of gravity acting on the particles. For example, sa ...
ize as normal. The introduction of both dreissenid species into the Great Lakes appears to be the result of ballast water discharge from transoceanic ships that were carrying veligers, juveniles, or adult mussels. The genus ''Dreissena'' is highly polymorphic and prolific with high potential for rapid adaptation attributing to its rapid expansion and colonization. Still, other factors can aid in the spread of these species across North American waters, such as larval drift in river systems or fishing and boating activities that allow for overland transport or movement between water basins. The success of overland transport of ''Dreissena'' species depends on their ability to tolerate periods of desiccation, and results suggest that given temperate summer conditions, adult ''Dreissena'' mussels may survive 3-5 days of aerial exposure. Quaggas are prodigious water filterers, removing substantial amounts of phytoplankton and suspended particulates from the water. By removing the phytoplankton, quaggas, in turn, decrease the food source for
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
, therefore altering the food web. Impacts associated with the filtration of water include increases in water transparency, decreases in mean chlorophyll concentrations, and accumulation of pseudofeces. Water clarity increases light penetration, causing a proliferation of aquatic plants that can change species dominance and alter the entire ecosystems. The pseudofeces that are produced from filtering the water accumulate and impact the environment. As the waste particles decompose, oxygen is used up, water acidity increases (decreased pH), and toxic byproducts are produced. In addition, quagga mussels accumulate organic pollutants within their tissues to levels more than 300,000 times greater than concentrations in the environment, and these pollutants are found in their pseudofeces, which can be passed up the food chain, therefore increasing wildlife exposure to organic pollutants. Another major threat involves the fouling of native freshwater mussels. Since quaggas were discovered in Lake Michigan in 1998, plankton rings formed by the passage of storms have been eaten away by the quagga mussels, threatening the local ecosystem. Each coin-sized quagga can filter up to a liter of water per day, stripping away the plankton that for thousands of years directly and indirectly sustained the native fish. Much of that food supply has now been sucked to the lake bottom; for every pound of prey fish swimming in the lake today, an estimated three to four pounds of quaggas are clustering on the lake bed.jsonline.com: 'The lake left me. It's gone.'
13 Aug 2011
''Dreissenas ability to rapidly colonize hard surfaces causes serious economic problems. These major
biofouling Biofouling or biological fouling is the accumulation of microorganisms, plants, algae, or small animals where it is not wanted on surfaces such as ship and submarine hulls, devices such as water inlets, pipework, grates, ponds, and rivers that ...
organisms can clog water-intake structures, such as pipes and screens, therefore reducing pumping capabilities for power and
water-treatment Water treatment is any process that improves the quality of water to make it appropriate for a specific end-use. The end use may be drinking, industrial water supply, irrigation, river flow maintenance, water recreation or many other uses, includ ...
plants, costing industries, companies, and communities. Recreation-based industries and activities have also been impacted; docks,
breakwall A breakwater is a permanent structure constructed at a coastal area to protect against tides, currents, waves, and storm surges. Part of a coastal management system, breakwaters are installed to minimize erosion, and to protect anchorages, h ...
s, buoys,
boat A boat is a watercraft of a large range of types and sizes, but generally smaller than a ship, which is distinguished by its larger size, shape, cargo or passenger capacity, or its ability to carry boats. Small boats are typically found on inl ...
s, and
beach A beach is a landform alongside a body of water which consists of loose particles. The particles composing a beach are typically made from rock, such as sand, gravel, shingle, pebbles, etc., or biological sources, such as mollusc sh ...
es have all been heavily colonized. Many of the potential impacts of ''Dreissena'' are unclear due to the limited time scale of North American colonization. Nonetheless, the genus ''Dreissena'' clearly is highly polymorphic and has a high potential for rapid adaptation to extreme environmental conditions, possibly leading to significant long-term impacts on North American waters. Also, the colonization of deeper water by ''D. r. bugensis'', exposes the quagga to a new range of environmental conditions and new habitats. It causes many of the same problems (stripping life-supporting algae, damaging boats, power plants, and harbors and destroying the native mussel population) as the equally invasive zebra mussel of Russia. It is also displacing native burrowing amphipod (''Diporeia hoyi'') from the deep waters of Lake Erie. In January 2007, quagga mussels were discovered at a marina in the Nevada portion of
Lake Mead Lake Mead is a reservoir formed by the Hoover Dam on the Colorado River in the Southwestern United States. It is located in the states of Nevada and Arizona, east of Las Vegas. It is the largest reservoir in the US in terms of water capacity. L ...
, and two other lakes on the
Colorado River The Colorado River ( es, Río Colorado) is one of the principal rivers (along with the Rio Grande) in the Southwestern United States and northern Mexico. The river drains an expansive, arid watershed that encompasses parts of seven U.S. s ...
,
Lake Mohave Lake Mohave is a reservoir on the Colorado River between the Hoover Dam and Davis Dam in Cottonwood Valley defining the border between Nevada and Arizona in the United States. This 67 mile stretch of the Colorado River flows past Boulder City, ...
and
Lake Havasu Lake Havasu () is a large reservoir formed by Parker Dam on the Colorado River, on the border between San Bernardino County, California and Mohave County, Arizona, Arizona. Lake Havasu City sits on the Arizona (eastern) side of the lake with its ...
. In 2008, the threat of quagga mussels being introduced at
Lake Casitas Lake Casitas is a reservoir in Ventura County, California, built by the United States Bureau of Reclamation and completed in 1959. The project provides drinking water and water for irrigation. A secondary benefit is flood control. It was the ven ...
and Westlake Lake in California from recreational boating resulted in the lakes' banning the use of outside boats. As of March 2008, other lakes such as Castaic and Lake Cachuma are considering similar bans. In June 2008, the mussels were confirmed in
Lake Granby Lake Granby is the third largest body of water in Colorado. It was created by the erection of Granby Dam, completed in 1950, as part of the Bureau of Reclamation's Colorado-Big Thompson Project. Water from Lake Granby is pumped via the Farr Pump ...
, Colorado. Larval quagga mussels were found in the water. After 5 years of negative testing Lake Granby, Colorado has been classified as a negative waterbody for quagga mussel veligers. Quagga mussels are now in all parts of Lake Powell on the Utah and Arizona border. They were also suspected in Deer Creek Reservoir at the top of Provo Canyon in Utah, but this body of water has since been delisted as a quagga-suspected water after 3 years of negative testing.


Quagga mussels as prey

In 1994, invasive-species biologist Anthony Ricciardi determined that North American
yellow perch The yellow perch (''Perca flavescens''), commonly referred to as perch, striped perch, American perch, American river perch or preacher is a freshwater perciform fish native to much of North America. The yellow perch was described in 1814 by Sam ...
found the invasive dreissenid species palatable. In 2004, he determined that yellow perch, over the intervening 10 years, had developed an appetite for the quagga mussel. While this sounds like good news, the problem is that this feeding process introduces contaminants into the
food chain A food chain is a linear network of links in a food web starting from producer organisms (such as grass or algae which produce their own food via photosynthesis) and ending at an apex predator species (like grizzly bears or killer whales), de ...
, notably ''
Clostridium botulinum ''Clostridium botulinum'' is a Gram-positive, rod-shaped, anaerobic, spore-forming, motile bacterium with the ability to produce the neurotoxin botulinum. The botulinum toxin can cause botulism, a severe flaccid paralytic disease in humans ...
''. Redear sunfish, a specialized mollusk-eating fish, are now being stocked in the Colorado River drainage as a defense against the quaggas. As with the yellow perch, this predator–prey relationship could cause toxins and micro-organisms to move up the food chain. Although quaggas are edible for humans, eating them is not recommended due to the accumulation of toxins, pollutants, and microorganisms within the mussels' bodies.


References


External links


''Dreissena bugensis''
- Small blurb on Quagga mussel (with pic) on Anthony Ricciardi's site. Links to various papers and blurbs on other invasive species.

- A brief fact sheet on information on Quagga and Zebra Mussels.
GLANSIS Species Fact Sheet
United States Geological Survey
Species Profile - Quagga Mussel (''Dreissena bugensis'')
National Invasive Species Information Center,
United States National Agricultural Library The United States National Agricultural Library (NAL) is one of the world's largest agricultural research libraries, and serves as a national library of the United States and as the library of the United States Department of Agriculture. Located ...
. Lists general information and resources for Quagga Mussel. {{Authority control Dreissenidae Molluscs described in 1897