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''Styela clava'' is a solitary, subtidal ascidian
tunicate Tunicates are marine invertebrates belonging to the subphylum Tunicata ( ). This grouping is part of the Chordata, a phylum which includes all animals with dorsal nerve cords and notochords (including vertebrates). The subphylum was at one time ...
. It has a variety of common names such as the stalked sea squirt, clubbed tunicate, Asian tunicate, leathery sea squirt, or rough sea squirt. As its common names suggest, ''S. clava'' is club-shaped with an elongated oval body and a long peduncle for attaching to a substrate. Although native to the northwestern waters of the
Pacific Ocean The Pacific Ocean is the largest and deepest of Earth's five Borders of the oceans, oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean, or, depending on the definition, to Antarctica in the south, and is ...
, since the 1900s, ''S. clava'' has become an increasingly successful
invasive species An invasive species is an introduced species that harms its new environment. Invasive species adversely affect habitats and bioregions, causing ecological, environmental, and/or economic damage. The term can also be used for native spec ...
outside of its native range. It is edible.


Morphology

''S. clava'' is a solitary tunicate. Including both the club-shaped body and peduncle, larger specimen of ''S. clava'' can have a maximum length of around 130 mm (5.1 in) and smaller specimen only reaching 30 mm (1.2 in) in length. Smaller specimen tend to have no distinct peduncle. As described by some of its common names, ''S. clava'' has a tough, wrinkled or irregularly grooved skin and comes in two variations of color dependent on size. Larger specimen have a light brown body and a darker brown peduncle while smaller specimen are yellow-brown.


Habitat

''S. clava'' is a marine
invertebrate Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordata, chordate s ...
animal. Adults are entirely sessile, growing attached to hard subtidal substrata as deep as 25 meters (82 ft). They can be found on virtually any hard surface such as rocks, buoys, pilings and shells of mussels. ''S. clava'' is predominant in the
littoral zone The littoral zone, also called litoral or nearshore, is the part of a sea, lake, or river that is close to the shore. In coastal ecology, the littoral zone includes the intertidal zone extending from the high water mark (which is rarely flood ...
, preferring sheltered localities free of strong wave action and floating objects, making artificial surfaces in harbors and marinas exceptional habitat. It is a hardy species that can live in a wide range of temperatures from -2 °C to 27 °C and can tolerate high salinity waters (26% - 28% salinity) that would be lethal to other tunicate species.


Reproduction

Like most tunicates, ''S. clava'' is hermaphroditic and produces short-lived
pelagic The pelagic zone consists of the water column of the open ocean and can be further divided into regions by depth. The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or water column between the sur ...
lecithotrophic larvae. They reproduce externally via broadcast spawning, and the reproductive period is highly dependent on
sea surface temperature Sea surface temperature (or ocean surface temperature) is the ocean temperature, temperature of ocean water close to the surface. The exact meaning of ''surface'' varies in the literature and in practice. It is usually between and below the sea ...
s reaching a critical temperature threshold, between 16 °C and 20°C. The reproductive period can vary from 4 to 10 months depending on location. Along the Californian coast in the United States, the reproduction period occurs for 4 months from June to September, while in Denmark and England, the reproduction period is also 4 months but occurs from July to October.


Native Range

''S. clava'' is native to the northwestern waters of the Pacific, particularly the seas and coasts of Japan and Korea, to Siberia, and as far south as the coasts off Shanghai, China.


Invasions

Outside of its native range, ''S. clava'' has proven to be an increasingly successful invasive species due to physiological adaptations and environmental tolerances. ''S. clava's'' thick tunic, relative to native tunicates, provides better protection from possible predators and helps prevent
desiccation Desiccation is the state of extreme dryness, or the process of extreme drying. A desiccant is a hygroscopic (attracts and holds water) substance that induces or sustains such a state in its local vicinity in a moderately sealed container. The ...
. It can withstand subzero to 23 °C sea temperatures and high salinity water, giving it strong tolerance to environmental changes in water. The lack of a natural predator already gives ''S. clava'' an advantage over native tunicates, but their large size as well allows them to outcompete other filter-feeding species such as oysters or mussels for food and substrate space.


Range

Since the mid-1900s, ''S. clava'' has been unintentionally introduced globally to temperate coastal waters outside of its native range. It has successfully established stable populations on both coasts of North America, Europe, New Zealand, Australia, and Argentina. The earliest sighting of ''S. clava'' outside of its natural range was on the United States’ west coast in Californian coastal waters in the early-1900s. Since then, the invasive tunicate has spread as far south as Baja, Mexico and as far north as Vancouver Island, Canada. ''S. clava'' populations in North America’s Atlantic waters is believed to have been introduced around the 1970s. In the mid-1900s, the next sighting of ''S. clava'' was recorded in European waters in Britain. In the span of 25 years, ''S. clava'' populations have expanded their range in the coastal waters of the United Kingdom and to mainland Europe. The current European countries with established ''S. clava'' populations are England, Ireland, Belgium, Netherlands, Denmark, France, Portugal and Spain.


Vectors

For each region there are multiple pathways by which ''S. clava'' could have been introduced. Although not definitive, ''S. clava's'' new population distribution is most likely due to
anthropogenic Anthropogenic ("human" + "generating") is an adjective that may refer to: * Anthropogeny, the study of the origins of humanity Anthropogenic may also refer to things that have been generated by humans, as follows: * Human impact on the enviro ...
vectors. The introduction of ''S. clava'' was most likely the result of direct introduction from Japan through shellfish transfers or ship fouling. In Europe, the source of introduction is believed to be from the hulls of military ships returning from the
Korean War The Korean War (25 June 1950 – 27 July 1953) was an armed conflict on the Korean Peninsula fought between North Korea (Democratic People's Republic of Korea; DPRK) and South Korea (Republic of Korea; ROK) and their allies. North Korea was s ...
. As waters are becoming warmer due to
climate change Present-day climate change includes both global warming—the ongoing increase in Global surface temperature, global average temperature—and its wider effects on Earth's climate system. Climate variability and change, Climate change in ...
, there is concern of habitat expansion into previously unoccupied waters.


Effects

Successful introduction and establishment of populations outside of its natural range can cause dramatic changes in the structure and composition of
benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from the Ancient Greek word (), meaning "the depths". ...
communities. It is dominating fouling communities, leading to population declines in other filter-feeding species, leading to lower
biodiversity Biodiversity is the variability of life, life on Earth. It can be measured on various levels. There is for example genetic variability, species diversity, ecosystem diversity and Phylogenetics, phylogenetic diversity. Diversity is not distribut ...
. ''S. clava'' is a solitary species, but with optimal conditions can reach high densities, up to 500 - 1,000 individuals, fouling man-made substrates resulting in boat and fishing gear difficulties. They also pose a threat to aquaculture, which is being seen in European waters. In Bassin de Thau, France ( Étang de Thau), ''S. clava'' is becoming a management problem as they pose a threat to oyster and mussel farming by outcompeting the shellfish for food and substrate space.


References


External links


National Introduced Marine Pest Information System of Australia: Leathery Sea Squirt

National Centre for Aquatic Biodiversity & Biosecurity of New Zealand: Invasive Sea Squirt Fact Sheet
* ttps://www.invasivespeciesinfo.gov/aquatic/invertebrates/clubbed-tunicate Species Profile - Clubbed Tunicate (''Styela clava'') 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 Clubbed Tunicate. {{Taxonbar, from=Q3023683 Styelidae Animals described in 1881 Taxa named by William Abbott Herdman