Quagga Mussel
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The quagga mussel (''Dreissena bugensis'') is a species (or subspecies) of
freshwater mussel Freshwater bivalves are molluscs of the order Bivalvia that inhabit freshwater ecosystems. They are one of the two main groups of freshwater molluscs, along with freshwater snails. The majority of bivalve molluscs are saltwater species that l ...
, an aquatic
bivalve Bivalvia () or bivalves, in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class (biology), class of aquatic animal, aquatic molluscs (marine and freshwater) that have laterally compressed soft bodies enclosed b ...
mollusk Mollusca is a phylum of protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000  extant species of molluscs are recognized, making it the second-largest animal phylum after Arthropoda. The ...
in the family Dreissenidae. It has an average lifespan of 3 to 5 years. The species is indigenous to the Dnipro River drainage of
Ukraine Ukraine is a country in Eastern Europe. It is the List of European countries by area, second-largest country in Europe after Russia, which Russia–Ukraine border, borders it to the east and northeast. Ukraine also borders Belarus to the nor ...
, and is named after the
quagga The quagga ( or ) (''Equus quagga quagga'') is an extinct 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 mtDNA ...
, an extinct subspecies of African
zebra Zebras (, ) (subgenus ''Hippotigris'') are African equines with distinctive black-and-white striped coats. There are three living species: Grévy's zebra (''Equus grevyi''), the plains zebra (''E. quagga''), and the mountain zebra (''E. ...
, 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 spanning the Canada–United States border. The five lakes are Lake Superior, Superior, Lake Michigan, Michigan, Lake Huron, H ...
of North America where it was brought by overseas shippers that use the
Saint Lawrence Seaway The St. Lawrence Seaway () is a system of rivers, locks, canals and channels in Eastern Canada and Northern United States that permits oceangoing vessels to travel from the Atlantic Ocean to the Great Lakes of North America, as far inland ...
.


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 ( ) 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 also has the shortest avera ...
that is pale or completely white. The shell has a rounded carina and a convex ventral side. The quagga mussel resembles the
zebra mussel The zebra mussel (''Dreissena polymorpha'') is a small freshwater mussel, an Aquatic animal, aquatic bivalve mollusk in the family Dreissenidae. The species originates from the lakes of southern Russia and Ukraine, but has been accidentally Intro ...
, 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 A hinge is a mechanical bearing that connects two solid objects, typically allowing only a limited angle of rotation between them. Two objects connected by an ideal hinge rotate relative to each other about a fixed axis of rotation, with all ...
. 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 Filter feeders are aquatic animals that acquire nutrients by feeding on organic matters, food particles or smaller organisms (bacteria, microalgae and zooplanktons) suspended in water, typically by having the water pass over or through a s ...
; it uses its
cilia The cilium (: cilia; ; in Medieval Latin and in anatomy, ''cilium'') is a short hair-like membrane protrusion from many types of eukaryotic cell. (Cilia are absent in bacteria and archaea.) The cilium has the shape of a slender threadlike proj ...
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 Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater Aquatic ecosystem, ecosystems. The name comes from the Greek language, Greek words (), meaning 'plant', and (), mea ...
,
zooplankton Zooplankton are the heterotrophic component of the planktonic community (the " zoo-" prefix comes from ), having to consume other organisms to thrive. Plankton are aquatic organisms that are unable to swim effectively against currents. Consequent ...
,
algae Algae ( , ; : alga ) is an informal term for any organisms of a large and diverse group of photosynthesis, photosynthetic organisms that are not plants, and includes species from multiple distinct clades. Such organisms range from unicellular ...
, 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 taxono ...
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 fluid, serous and muc ...
, 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 Dioecy ( ; ; adj. dioecious, ) is a characteristic of certain species that have distinct unisexual individuals, each producing either male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproduction is ...
(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 ( ; see also other names) is the second-longest river in Europe, after the Volga in Russia. It flows through Central and Southeastern Europe, from the Black Forest south into the Black Sea. A large and historically important riv ...
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 ...
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 ...
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 sea, marginal Mediterranean sea (oceanography), mediterranean sea 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 bound ...
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 lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...
, quagga mussels were first identified in 2005, and now populate many inland waters, such as the Rhine–Main–Danube Canal, the Main, and the
Rhine The Rhine ( ) is one of the List of rivers of Europe, major rivers in Europe. The river begins in the Swiss canton of Graubünden in the southeastern Swiss Alps. It forms part of the Swiss-Liechtenstein border, then part of the Austria–Swit ...
. They were first identified in
Switzerland Switzerland, officially the Swiss Confederation, is a landlocked country located in west-central Europe. It is bordered by Italy to the south, France to the west, Germany to the north, and Austria and Liechtenstein to the east. Switzerland ...
in 2015, and in
Lake Constance Lake Constance (, ) refers to three bodies of water on the Rhine at the northern foot of the Alps: Upper Lake Constance (''Obersee''), Lower Lake Constance (''Untersee''), and a connecting stretch of the Rhine, called the Seerhein (). These ...
in 2016, where they have since spread massively and caused considerable problems, in particular to the machinery in waterworks. In 2014, the species was reported at Wraysbury Reservoir, not far from London's Heathrow Airport in the valley of the
River Thames The River Thames ( ), known alternatively in parts as the The Isis, River Isis, is a river that flows through southern England including London. At , it is the longest river entirely in England and the Longest rivers of the United Kingdom, s ...
. In
Ireland Ireland (, ; ; Ulster Scots dialect, Ulster-Scots: ) is an island in the North Atlantic Ocean, in Northwestern Europe. Geopolitically, the island is divided between the Republic of Ireland (officially Names of the Irish state, named Irelan ...
, the mussels were first discovered in 2021 in two lakes on the
River Shannon The River Shannon ( or archaic ') is the major river on the island of Ireland, and at in length, is the longest river in the British Isles. It drains the Shannon River Basin, which has an area of , – approximately one fifth of the area of I ...
: Lough Derg and
Lough Ree Lough Ree (), translated to English as ''King's Lake'' or ''King Lake'', is a lake in the midlands of Republic of Ireland, Ireland, the second of the three major lakes on the River Shannon. Lough Ree is the second largest lake on the Shannon af ...
. 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 spanning the Canada–United States border. The five lakes are Lake Superior, Superior, Lake Michigan, Michigan, Lake Huron, H ...
, 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 ( ) 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 also has the shortest avera ...
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 Biomass is a term used in several contexts: in the context of ecology it means living organisms, and in the context of bioenergy it means matter from recently living (but now dead) organisms. In the latter context, there are variations in how ...
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; it has Chemical symbol, symbol P and atomic number 15. All elemental forms of phosphorus are highly Reactivity (chemistry), reactive and are therefore never found in nature. They can nevertheless be prepared ar ...
, remetabolizing and recirculating it instead of allowing it to
sediment Sediment is a solid material that is transported to a new location where it is deposited. It occurs naturally and, through the processes of weathering and erosion, is broken down and subsequently sediment transport, transported by the action of ...
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. They have as of August 2024 been for the first time found inland at the small freshwaters of Geneva Lake in Wisconsin. Quaggas are prodigious water filterers, removing substantial amounts of
phytoplankton Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater Aquatic ecosystem, ecosystems. The name comes from the Greek language, Greek words (), meaning 'plant', and (), mea ...
and suspended particulates from the water. By removing the phytoplankton, quaggas, in turn, decrease the food source for
zooplankton Zooplankton are the heterotrophic component of the planktonic community (the " zoo-" prefix comes from ), having to consume other organisms to thrive. Plankton are aquatic organisms that are unable to swim effectively against currents. Consequent ...
, therefore altering the food web. Impacts associated with the filtration of water include increases in water transparency, decreases in mean
chlorophyll Chlorophyll is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words (, "pale green") and (, "leaf"). Chlorophyll allows plants to absorb energy ...
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 plants, costing industries, companies, and communities. Recreation-based industries and activities have also been impacted;
dock The word dock () in American English refers to one or a group of human-made structures that are involved in the handling of boats or ships (usually on or near a shore). In British English, the term is not used the same way as in American Engl ...
s, breakwalls,
buoy A buoy (; ) is a buoyancy, floating device that can have many purposes. It can be anchored (stationary) or allowed to drift with ocean currents. History The ultimate origin of buoys is unknown, but by 1295 a seaman's manual referred to navig ...
s,
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 or capacity, its shape, or its ability to carry boats. Small boats are typically used on inland waterways s ...
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 (geology), rock, such as sand, gravel, shingle beach, shingle, pebbles, etc., or biological s ...
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 () is one of the principal rivers (along with the Rio Grande) in the Southwestern United States and in northern Mexico. The river, the List of longest rivers of the United States (by main stem), 5th longest in the United St ...
,
Lake Mohave Lake Mohave is a reservoir on the Colorado River between the Hoover Dam and Davis Dam in Cottonwood Valley (Arizona/Nevada), Cottonwood Valley defining the border between Nevada and Arizona in the United States. This stretch of the Colorado Rive ...
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. Lake Havasu City sits on the Arizonan side of the lake with its Californian coun ...
. In 2008, the threat of quagga mussels being introduced at Lake Casitas 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, 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 or preacher is a freshwater perciform fish native to much of North America. The yellow perch was described in 1814 by Samuel Latham Mitchill fr ...
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, often starting with an autotroph (such as grass or algae), also called a producer, and typically ending at an apex predator (such as grizzly bears or killer whales), detritivore (such as ...
, notably ''
Clostridium botulinum ''Clostridium botulinum'' is a Gram-positive bacteria, gram-positive, Bacillus (shape), rod-shaped, Anaerobic organism, anaerobic, endospore, spore-forming, Motility, motile bacterium with the ability to produce botulinum toxin, which is a neurot ...
''.
Redear sunfish The redear sunfish (''Lepomis microlophus''), also known as the shellcracker, Georgia bream, cherry gill, chinquapin, improved bream, and sun perch, is a freshwater fish in the family Centrarchidae and is native to the southeastern United States ...
, 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.


See also

* '' Dreissena rostriformis distincta''


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. Locate ...
. Lists general information and resources for Quagga Mussel. {{Authority control Dreissenidae Molluscs described in 1897