Thick Shelled River Mussel
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''Unio crassus'', the thick shelled river mussel, is a
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
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 Aquatic means relating to water; living in or near water or taking place in water; does not include groundwater, as "aquatic" implies an environment where plants and animals live. Aquatic(s) may also refer to: * Aquatic animal, either vertebrate ...
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 Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Unionidae The Unionidae are a Family (biology), family of freshwater mussels, the largest in the order Unionida, the bivalve molluscs sometimes known as river mussels, or simply as unionids. The range of distribution for this family is world-wide. It is a ...
, the river mussels.MolluscaBase eds. (2023). MolluscaBase. Unio crassus Philipsson, 1788. Accessed through: World Register of Marine Species at: https://marinespecies.org/aphia.php?p=taxdetails&id=1044116 on 2023-01-09


Taxonomy

The species was split into two species, ''Unio crassus'' and '' Unio tumidiformis'', in 2009. Various subspecies have been noted, but their validity is not always recognized. Recently, two clades and possible subspecies, ''Unio crassus crassus'' and ''Unio crassus courtillieri'', have been identified based on genetic sampling.


Geography

Its range occurs across much of Europe, excluding
Britain Britain most often refers to: * Great Britain, a large island comprising the countries of England, Scotland and Wales * The United Kingdom of Great Britain and Northern Ireland, a sovereign state in Europe comprising Great Britain and the north-eas ...
,
Italy Italy, officially the Italian Republic, is a country in Southern Europe, Southern and Western Europe, Western Europe. It consists of Italian Peninsula, a peninsula that extends into the Mediterranean Sea, with the Alps on its northern land b ...
, and the
Iberian Peninsula The Iberian Peninsula ( ), also known as Iberia, is a peninsula in south-western Europe. Mostly separated from the rest of the European landmass by the Pyrenees, it includes the territories of peninsular Spain and Continental Portugal, comprisin ...
. It extends from
France France, officially the French Republic, is a country located primarily in Western Europe. Overseas France, Its overseas regions and territories include French Guiana in South America, Saint Pierre and Miquelon in the Atlantic Ocean#North Atlan ...
in the west to the
Ural River The Ural, also known as the Yaik , is a river flowing through Russia and Kazakhstan in the continental border between Europe and Asia. It originates in the southern Ural Mountains and discharges into the Caspian Sea. At , it is the third-longes ...
basin in
Russia Russia, or the Russian Federation, is a country spanning Eastern Europe and North Asia. It is the list of countries and dependencies by area, largest country in the world, and extends across Time in Russia, eleven time zones, sharing Borders ...
and
Kazakhstan Kazakhstan, officially the Republic of Kazakhstan, is a landlocked country primarily in Central Asia, with a European Kazakhstan, small portion in Eastern Europe. It borders Russia to the Kazakhstan–Russia border, north and west, China to th ...
in the east, and historically occupied most major river systems. It is considered extirpated from the
Netherlands , Terminology of the Low Countries, informally Holland, is a country in Northwestern Europe, with Caribbean Netherlands, overseas territories in the Caribbean. It is the largest of the four constituent countries of the Kingdom of the Nether ...
, where it was last seen in 1967.


Description

''Unio crassus'' is known for its thick-walled shell. They are generally 3–7 cm in length, though some individuals reach 9–11 cm. The size and shape of the shell may vary by population or by environmental factors.


Habitat

The species lives in large streams and rivers with clear water, though it is more tolerant of silt than other threatened species. Due to its sensitivity to pollution, it is considered a
bioindicator A bioindicator is any species (an indicator species) or group of species whose function, population, or status can reveal the qualitative status of the environment. The most common indicator species are animals. For example, copepods and other sma ...
for water quality. It prefers slower-flowing waters on the banks or near roots, boulders, or other features. It is especially vulnerable to changes in water chemistry that affect local fish populations, such as nitrate and phosphate concentrations.


Ecology

Its lifespan can be up to 80 years, depending on the water temperature, with an average of 20–30 years. It often forms colonies in large streams and rivers with clear, free-flowing water. They bury themselves in the stream bed, leaving only their siphons exposed through which they inhale oxygen and food (
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 microorganisms) and expel waste. It can move several meters, sometimes farther, to find suitable habitat conditions. They are tachytictic (short-term) breeders, meaning the cycle of fertilization to juvenile takes place in one season. Males release their spermatozoa into the water in the spring, which females take in through siphoning. Females can have several broods of eggs per year. The eggs develop for a few weeks until the larvae are ready for release. Like most
Unionidae The Unionidae are a Family (biology), family of freshwater mussels, the largest in the order Unionida, the bivalve molluscs sometimes known as river mussels, or simply as unionids. The range of distribution for this family is world-wide. It is a ...
mussels, the larvae have a parasitic stage where they are required to attach and feed off a host fish. In spring and summer, the female releases around 100,000
glochidia The glochidium (plural glochidia) is a microscopic larval stage of some freshwater mussels, aquatic bivalve mollusks in the families Unionidae and Margaritiferidae, the river mussels and European freshwater pearl mussels. These larvae are t ...
into the water current. The female has a unique method of dispersing the larvae. She crawls up to the edge of the water, exposing her excurrent
aperture In optics, the aperture of an optical system (including a system consisting of a single lens) is the hole or opening that primarily limits light propagated through the system. More specifically, the entrance pupil as the front side image o ...
, and then lets loose a stream of water containing
glochidia The glochidium (plural glochidia) is a microscopic larval stage of some freshwater mussels, aquatic bivalve mollusks in the families Unionidae and Margaritiferidae, the river mussels and European freshwater pearl mussels. These larvae are t ...
(larvae). It is suggested that this spurting behavior may lure host fish that are attracted to the water disturbance. The larvae can then attach to the fish's gills.
video of spurting mussel
The glochidia attach to the gills of a fish and remain for 20–50 days while they metamorphose into juvenile mussels, after which they drop off and bury themselves in the stream bed for 1–3 years. The most frequent host fishes have been identified as bullhead (
Cottus gobio The European bullhead (''Cottus gobio'') is a freshwater fish that is widely distributed in Europe, mainly in rivers. It is a member of the family Cottidae, a type of sculpin. It is also known as the miller's thumb, freshwater sculpin, common bu ...
), minnow (
Phoxinus phoxinus ''Phoxinus phoxinus'' (known colloquially as the Eurasian minnow, minnow, Channel minnow, or common minnow) is a species of freshwater ray-finned fish belonging to the family Leuciscidae, which includes the daces, minnows and related fishes. It i ...
), chub (
Leuciscus cephalus ''Squalius cephalus'', the common chub, European chub or simply chub, is a species of freshwater ray-finned fish belonging to the family Leuciscidae, the daces, Eurasian minnows and related fishes. This species is found in Europe where it freque ...
and
Squalius cephalus ''Squalius cephalus'', the common chub, European chub or simply chub, is a species of freshwater ray-finned fish belonging to the family Leuciscidae, the daces, Eurasian minnows and related fishes. This species is found in Europe where it freq ...
), rudd (
Scardinius erythrophthalmus The common rudd (''Scardinius erythrophthalmus'') is a bentho-pelagic freshwater ray-finned fish belonging to the family Leuciscidae. This species is widely spread in Europe and central Asia, around the basins of the North, Baltic, Black, Caspi ...
), bleak (
Alburnus alburnus The bleak or common bleak (''Alburnus alburnus'') is a small freshwater coarse fish of the family Leuciscidae, which includes the minnows, daces and bleaks. The common bleak is found in Europe and Western Asia. Description The body of the ...
), nase (
Chondrostoma nasus The common nase (''Chondrostoma nasus'') is a European potamodromous cyprinid fish. It is often simply called the nase, but that can refer to any species of its genus '' Chondrostoma''. Another name is sneep. Distribution The nase is found natur ...
), stickleback (
Gasterosteus aculeatus The three-spined stickleback (''Gasterosteus aculeatus'') is a fish native to most inland and coastal waters north of 30°N. It has long been a subject of scientific study for many reasons. It shows great morphological variation throughout its ra ...
), ide (
Leuciscus idus The ide (''Leuciscus idus''), or orfe, is a species of freshwater ray-finned fish belonging to the family Leuciscidae. The ide is found in larger rivers, ponds, and lakes across Northern Europe and Asia. It has been introduced outside its nativ ...
) and perch (
Perca fluviatilis The European perch (''Perca fluviatilis''), also known as the common perch, redfin perch, big-scaled redfin, English perch, Euro perch, Eurasian perch, Eurasian river perch, Hatch, poor man's rockfish or in Anglophone parts of Europe, simply the ...
). It is not able to metamorphose on roach (
Rutilus rutilus The roach, or rutilus roach (''Rutilus rutilus''), also known as the common roach, is a fresh- and brackish-water fish of the family Cyprinidae, native to most of Europe and western Asia. Fish called roach can be any species of the genera ''Rut ...
) and sterlet (
Acipenser ruthenus The sterlet (''Huso ruthenus'') is a relatively small species of sturgeon from Eurasia native to large rivers that flow into the Black Sea, Azov Sea, and Caspian Sea, as well as rivers in Siberia as far east as Yenisei. Populations migrating b ...
). The most suitable host fish can vary by locality and population. For example, in the
Danube The Danube ( ; see also #Names and etymology, other names) is the List of rivers of Europe#Longest rivers, second-longest river in Europe, after the Volga in Russia. It flows through Central and Southeastern Europe, from the Black Forest sou ...
drainage, European chub (
Squalius cephalus ''Squalius cephalus'', the common chub, European chub or simply chub, is a species of freshwater ray-finned fish belonging to the family Leuciscidae, the daces, Eurasian minnows and related fishes. This species is found in Europe where it freq ...
) is considered the most suitable host for ''U. crassus''. Invasive species such as round goby (
Neogobius melanostomus The round goby (''Neogobius melanostomus'') is a euryhaline bottom-dwelling species of fish of the family Gobiidae. It is native to Central Eurasia, including the Black Sea and the Caspian Sea. Round gobies have established large non-native popul ...
) and rainbow trout (
Oncorhynchus mykiss The rainbow trout (''Oncorhynchus mykiss'') is a species of trout native to cold-water tributaries of the Pacific Ocean in North America and Asia. The steelhead (sometimes called steelhead trout) is an anadromous (sea-run) form of the coastal ...
) were also unable to keep larvae alive.


Use by humans

Freshwater mussels were consumed by prehistoric people in Central Europe. Archaeological evidence from shell collections in
Hungary Hungary is a landlocked country in Central Europe. Spanning much of the Pannonian Basin, Carpathian Basin, it is bordered by Slovakia to the north, Ukraine to the northeast, Romania to the east and southeast, Serbia to the south, Croatia and ...
as far back as the
Neolithic The Neolithic or New Stone Age (from Ancient Greek, Greek 'new' and 'stone') is an archaeological period, the final division of the Stone Age in Mesopotamia, Asia, Europe and Africa (c. 10,000 BCE to c. 2,000 BCE). It saw the Neolithic Revo ...
period indicate that several mussel species were gathered from a nearby river and consumed raw as an important supplementary food source. ''Unio crassus'' appeared to be a preferred species, likely because of its relatively high meat content. '' Unio'' shells used as tools, including ''U. crassus'', have been found at archaeological sites dating to the
5th millennium BC The 5th millennium BC spanned the years (5000 BC – 4001 BC) (c. 7 ka to c. 6 ka), that is, inclusive of 5000 BC but exclusive of 4000 BC. It is impossible to precisely date events that happened around the time of this millennium ...
. ''Unio crassus'' was also one of several mussels used to make ornamental buttons between 4200-3800 BC in central Europe. Widespread occurrence of these ornaments, from
Denmark Denmark is a Nordic countries, Nordic country in Northern Europe. It is the metropole and most populous constituent of the Kingdom of Denmark,, . also known as the Danish Realm, a constitutionally unitary state that includes the Autonomous a ...
to
Romania Romania is a country located at the crossroads of Central Europe, Central, Eastern Europe, Eastern and Southeast Europe. It borders Ukraine to the north and east, Hungary to the west, Serbia to the southwest, Bulgaria to the south, Moldova to ...
, indicates trade or cross-cultural exchange of knowledge. The white hue of the
nacre Nacre ( , ), also known as mother-of-pearl, is an organicinorganic composite material produced by some molluscs as an inner shell layer. It is also the material of which pearls are composed. It is strong, resilient, and iridescent. Nacre is ...
was highly sought after. Freshwater
mother-of-pearl Nacre ( , ), also known as mother-of-pearl, is an organicinorganic composite material produced by some molluscs as an inner shell layer. It is also the material of which pearls are composed. It is strong, resilient, and iridescent. Nacre is ...
was used in ornaments through the
Middle Ages In the history of Europe, the Middle Ages or medieval period lasted approximately from the 5th to the late 15th centuries, similarly to the post-classical period of global history. It began with the fall of the Western Roman Empire and ...
. The species was used as livestock fodder in Central Europe in the 19th century. Today, it is still used as fodder for poultry in Romania, and for human consumption in some areas.


Use as a host

In addition to being a parasite in its larval stage, the thick-shelled river mussel also acts as a host for the endangered
European bitterling The European bitterling (''Rhodeus amarus'') is a temperate freshwater ray-finned fish belonging to the family Acheilognathidae, the bitterlings. It originates in Europe, ranging from the Rhone River basin in France to the Neva River in Russia. ...
(''Rhodeus amarus''). The bitterling's eggs must develop in the gills of freshwater mussels. Bitterling egg-infested mussels may have reduced growth rates.


Threats and conservation

During the 20th century, the thick shelled river mussel declined in
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
by as much as 50% due to deteriorating water quality,
habitat fragmentation Habitat fragmentation describes the emergence of discontinuities (fragmentation) in an organism's preferred environment (habitat), causing population fragmentation and ecosystem decay. Causes of habitat fragmentation include geological proces ...
, host fish limitation, canalisation of rivers, and introduction of the
muskrat The muskrat or common muskrat (''Ondatra zibethicus'') is a medium-sized semiaquatic rodent native to North America and an introduced species in parts of Europe, Asia, and South America. The muskrat is found in wetlands over various climates ...
(''Ondatra zibethicus'') to European rivers. A 2020 study identified habitat destruction, predation by muskrat, and lack of host fish as the major factors for decline. ''U. crassus'' was noted to be more tolerant of silt in the water than other threatened species. The most stable populations are found in Russia and northern Europe. A project in Poland successfully used artificial breeding methods including release of gravid females, release of glochidia-infested fish, and in vitro-raised juveniles, to double the population of the mussel in the Biała Tarnowska river. Long-term conservation of the species will require changes in river management practices, controlling pollution and invasive species, restoration of river basins, and further outreach and education. More research is needed to identify priority populations for monitoring.


Status by country

Its native distribution is Europe and
Western Asia West Asia (also called Western Asia or Southwest Asia) is the westernmost region of Asia. As defined by most academics, UN bodies and other institutions, the subregion consists of Anatolia, the Arabian Peninsula, Iran, Mesopotamia, the Armenian ...
. It is mentioned in annexes II and IV of the European Union
Habitats Directive The Habitats Directive (more formally known as Council Directive 92/43/EEC on the Conservation of natural habitats and of wild fauna and flora) is a directive adopted by the European Community in 1992 as a response to the Berne Convention. The ...
. Globally, it is ranked endangered on the
IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is an inventory of the global conservation status and extinction risk of biological ...
. Local statuses are as follows: * Austria - critically endangered * Albania - vulnerable * Belarus - vulnerable * Belgium - strictly protected * Bulgaria - protected * Croatia - endangered *
Czech Republic The Czech Republic, also known as Czechia, and historically known as Bohemia, is a landlocked country in Central Europe. The country is bordered by Austria to the south, Germany to the west, Poland to the northeast, and Slovakia to the south ...
– endangered Its conservation status in 2004-06 was unfavourable (U2) according to a report for the
European Commission The European Commission (EC) is the primary Executive (government), executive arm of the European Union (EU). It operates as a cabinet government, with a number of European Commissioner, members of the Commission (directorial system, informall ...
in accordance with the Habitats Directive. * Denmark – believed to have been extirpated until rediscovered in Odense River in 2003. The Odense River population is estimated to number 3000 individuals and subsequently it has been rediscovered from
Suså River The Suså or Susåen is a river in Denmark. It is the largest drainage system on the island of Zealand and the fifth largest in Denmark as a whole. Geography The source of the Suså is in Tingerup Tykke, a woodland south of Rønnede. Its fi ...
where the population size is unknown (might also survive in a couple of other rivers, but this remains unconfirmed). * Finland – in southern Finland. Vulnerable. * France * Germany – critically endangered (''vom Aussterben bedroht''), strictly protected In Germany this bivalve has disappeared from 90% of its former range. * Hungary - protected and rare * Kazakhstan - least concern * Latvia - vulnerable and protected * Lithuania - sometimes reported as "Extinct in the wild", but populations exist in the
Neman River Neman, Nemunas or Niemen is a river in Europe that rises in central Belarus and flows through Lithuania then forms Lithuania–Russia border, the northern border of Kaliningrad Oblast, Russia's western exclave, which specifically follows its s ...
drainage. * Netherlands – locally extinct. In the
Netherlands , Terminology of the Low Countries, informally Holland, is a country in Northwestern Europe, with Caribbean Netherlands, overseas territories in the Caribbean. It is the largest of the four constituent countries of the Kingdom of the Nether ...
it has not been seen alive after 1968 and is most likely
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
in that country. * Poland – endangered * Romania - endangered * Russia - least concern * Slovakia – vulnerable and protected * Sweden – endangered * Switzerland - critically endangered


References


External links


The Mussel Project Web Site

''Fauna Europaea''
{{Taxonbar , from=Q798236
crassus Marcus Licinius Crassus (; 115–53 BC) was a Roman general and statesman who played a key role in the transformation of the Roman Republic into the Roman Empire. He is often called "the richest man in Rome". Wallechinsky, David & Walla ...
Bivalves of Asia Bivalves of Europe Freshwater bivalves Bivalves described in 1788 Habitats Directive species