Invasion History

First Non-native Panama (Pacific) Tidal Record: 2008

Panama Invasion History:


Invasion history elsewhere in the world:

Charles Darwin examined specimens from Guayaquil, Ecuador and 'West Colombia’ (Darwin 1854), which may have represented populations that were carried by Spanish ships from  Europe or from the Caribbean or Atlantic South America (Carlton et al. 2011). Early shipping could also explain a 1889 collection from the Gulf of California, Mexico (Henry and McLaughlin 1975). However, it is surprising that there are few records from elsewhere on the tropical-subtropical Pacific Coast. Interestingly, this freshwater-tolerant barnacle was not reported on the Pacific coast of Panama until 2008, when it was collected at Isla Taboguilla, at the entrance of the Panama Canal (Ruiz et al. unpublished data).

Amphibalanus improvisus was first recorded from the Northwest Pacific in Tokyo Harbor in 1952. By 1968, it had spread to the Sea of Japan, where it is known from Zolotoi Rog (Golden Horn) Bay, Russia, to Kanking-Chinhan, South Korea (Kim 1992; Zvyagintsev 2003). This barnacle is now abundant in the region, especially in brackish waters. It has been recorded from Western Australia (Furlani 1996), but is not reported to have become established.

We currently consider Amphibalanus improvisus to be introduced in the Northeast Atlantic waters of Europe. It was widespread in brackish waters of England, Scotland and the Netherlands by 1854, and first reported from French Atlantic waters in 1872. It was found at a 17th century archeological site in Antwerp, Belgium (Kerckhof and Cattrijsse 2001), which could indicate that it was either a very early introduction to European waters, or that native populations were present. Carlton et al. (2011) note that Zullo and Miller (1986) argued for its Western Atlantic origin, citing the lack of verified fossils in the Eastern Atlantic and Mediterranean. Further examination of fossil and archaeological specimens may cause us to change its status to cryptogenic or native. This barnacle expanded its range into the Baltic Sea during historic times, being first reported from Kaliningrad, Russia, in 1844 and extending to the Northern Quark, Finland on the Gulf of Bothnia, by 1994 (Leppakoski and Olenin 2000). Amphibalanus improvisus occurs throughout the Mediterranean (date of first record is not yet known), mostly in brackish lagoons (Relini and Matricardi 1999; Kocak and Kucuksezgin 2000), and the Black Sea (1st record 1844, Gomiou et al. 2002), and reached the Caspian Sea by 1955 (Grigorevich et al. 2003).

Furman and Yule (1991) found that in British waters, many populations of Amphibalanus improvisus are ephemeral. In a 1985-1987 survey, they did not find this barnacle at many of the locations reported by Darwin, or at locations where it was found in the 1950s and 1970s. In some cases, such as the Thames estuary, early (pre 1854) populations apparently disappeared by the 1950s, but recolonized (Furman and Yule 1991). These fluctuations may be due to short-term weather and pollution changes.


Description

The shell of Amphibalanus improvisus is usually conical or subcylindrical. The orifice is slightly toothed, and its width is usually more than 1/2 its height. The plates have a smooth surface, with narrow longitudinal spaces (radii), narrowing to the tops of shell plates. The radii are white. Inside the operculum, the scutum has a well-developed adductor ridge on its interior face (Zullo 1979). The tergum has a blunt apex, and its spur is usually somewhat long and narrow. The spur length is about 1/3 of the length of the basal margin, and its width is about 1/5 of the basal margin (Henry and McLaughlin 1975). Amphibalanus improvisus grows up to 17 mm in diameter. It is characteristic of brackish estuarine habitats, with very low or highly variable salinity (Henry and McLaughlin 1975). Larval stages of A. improvisus are illustrated by Lang (1979; 1980).

Local ecophenotypes or genotypes are to be expected throughout the range of this species (Bacon, 1976; Henry and McLaughlin, 1975), and it may be that tropical "improvisus are a cryptic sibling species (J. T. Carlton, pers. comm., 2013).


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Arthropoda
Subphylum:   Crustacea
Class:   Maxillopoda
Subclass:   Thecostraca
Infraclass:   Cirripedia
Superorder:   Thoracica
Order:   Sessilia
Suborder:   Balanomorpha
Superfamily:   Balanoidea
Family:   Balanidae
Genus:   Amphibalanus
Species:   improvisus

Synonyms

Balanus amphitrite var. amphitrite (Stubbings, 1967)
Balanus amphitrite var. assimilis (Darwin, 1854)
Balanus var. denticulata (Stubbings, 1961)
Balanus amphitrite var. pallidus (Stubbings, 1961)
Balanus amphitrite vostokensis (Tarasov & Zevina, 1957)

Potentially Misidentified Species

Amphibalanus amphitrite
Now cosmopolitan, warm-temprate -subtropical

Amphibalanus eburneus
NW Atlantic native, widely introduced in tropical-temperate waters

Amphibalanus pallidus
Tropical Atlantic, introduced in Pacific Panama, establishment unknown

Amphibalanus reticulatus
Indo-Wesr Pacific native, now widely introduced in subtropical-tropical waters

Amphibalanus subalbidus
NW Atlantic native, introduced in Brazil and Gulf of California

Ecology

General:

Like many other barnacles, Amphibalanus improvisus, is hermaphroditic, but is capable of cross-fertilization. The fertilized eggs are brooded in the mantle cavity, sometimes for several months, and are released as nauplius larvae with three pairs of appendages (Barnes 1983). The nauplii feed in the plankton and go through five successive molts, spending four to 18 days in the water column before molting into a non-feeding cypris stage, covered with a pair of chitinous shells (Lang and Marcy 1982; Furman and Yule 1991). Cyprids swim, investigating suitable surfaces, and then settle, secreting a shell, and molting into the first juvenile barnacle stages. Juvenile and adult barnacles are filter feeders, sweeping the water with their long bristled appendages to gather phytoplankton, zooplankton, and detritus (Barnes 1983).

The Bay Barnacle, Amphibalanus improvisus, is characteristic of estuaries and brackish waters, although it can also tolerate high salinities, up to 40 PSU (Foster 1970; Furman and Yule 1991). In estuaries, it can survive weeks of exposure to freshwater, but requires salinities of at least 2 PSU for reproduction (Dineen and Hines 1992). It is typically found in lower intertidal and subtidal zones, in sheltered waters. This barnacle grows on a wide range of hard surfaces, including logs, mangroves, rocks, ship hulls, oysters, other shellfish, docks and ships' hulls.

Food:

Phytoplankton; zooplankton

Trophic Status:

Suspension Feeder

SusFed

Habitats

General HabitatCoarse Woody DebrisNone
General HabitatOyster ReefNone
General HabitatMarinas & DocksNone
General HabitatRockyNone
General HabitatVessel HullNone
General HabitatMangrovesNone
General HabitatCanalsNone
Salinity RangeLimnetic0-0.5 PSU
Salinity RangeOligohaline0.5-5 PSU
Salinity RangeMesohaline5-18 PSU
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Salinity RangeHyperhaline40+ PSU
Tidal RangeSubtidalNone
Tidal RangeLow IntertidalNone
Vertical HabitatEpibenthicNone


Tolerances and Life History Parameters

Minimum Temperature (ºC)-2Based on range, including ice-covered estuaries.
Maximum Temperature (ºC)38Maximum temperature recorded in effluent canals of a powerplant on the Patuxent River MD, with populations of A. improvisus (Nauman and Cory 1969). Maximum tolerance levels may be higher.
Minimum Salinity (‰)0Adults can survive long exposure to freshwater (Foster 1970)
Maximum Salinity (‰)40Rare at high salinities (Furman and Yule 1991), but present in the Mediterranean Sea.
Minimum Reproductive Temperature10Bousfield 1955
Maximum Reproductive Temperature30Bousfield 1955
Minimum Reproductive Salinity2Experimental, lowest tested, 28% metamorphosis, compared to 71-75% at 16-32 ppt (Furman and Yule 1991).
Maximum Reproductive Salinity40Experimental, highest tested, 45% metamorphosis, compared to 71-75% at 16-32 ppt, all at 30 C (Furman and Yule 1991).
Minimum Duration4Experimental, 30 C, 16-24 ppt, high food density (Furman and Yule 1989).
Maximum Duration1815 C, 18 days for intermittently starved larvae (Lang and Marcy 1982)
Maximum Length (mm)9Maximum adult height (Henry and McLauglin 1975)
Maximum Width (mm)17Maximum adult width (Henry and McLauglin 1975)
Broad Temperature RangeNoneCold temperate-Tropical
Broad Salinity RangeNoneOligohaline-Euhaline

General Impacts

Economic Impacts

We have not found specific reports of economic impacts for Amphibalanus improvisus in West Coast waters where it has been introduced. However, A. improvisus is one of the most abundant fouling barnacles in brackish waters of the U.S. (Carlton 1979), and is very likely a major contributor to fouling of ships, boats, harbor structures, fishing gear, and power plant intake pipes in estuaries. A variety of impacts are associated with A. improvisus worldwide, especially in regions where barnacles were previously absent, such as the Baltic and Caspian Seas.

Boating- Amphibalanus improvisus is widely reported as a common fouling organism of recreational and commercial boats and ships, and also of piers, docks, and navigational structures (Zvyagintsev 1985; Vuorinen et al. 1986; Kashin et al. 2000; Gren et al. 2009; Skolka and Preda 2010). In Skagerrak, Sweden, it was the dominant organism fouling the hulls of recreational boats, probably because it was more tolerant of hydrodynamic stress than the native Blue Mussel, Mytilus edulis, which dominated static fouling plates (Berntsson and Jonsson 2003). In Sweden, estimated costs of hull fouling by A. improvisus are 23-56 million dollars per year (Gren et al. 2009). Barnacle fouling contributes to the deterioration of piers and docks by increasing corrosion. In the Caspian Sea and in Russian harbors on the Sea of Japan, A. improvisus was an important component of fouling on harbor structures (Kashin et al. 2000; Zaitsev and Ozturk 2001; Zvyagintsev 2003).

Industry- Amphibalanus improvisus is a frequent fouler of power plants in its native and introduced range (Nauman and Cory 1969; Vuorinen et al. 1986; Zvyagintsev et al. 2003). In Sweden, estimated costs of power plant fouling by A. improvisus were 1.5-5.5 million dollars per year (Gren et al. 2009).

Fisheries- In the Caspian Sea, A. improvisus is reported to foul fishnets (Zaitsev and Ozturk 2001). More speculatively, it is thought to have adverse affects on fisheries by diverting plankton production to the benthic biomass, where it apparently constitutes a 'dead end' by having few predators. Consequently it could be decreasing fish biomass (Olenin and Leppakoski 2000).

Aesthetic- In the Baltic Sea, A. improvisus is reported to affect the recreational quality of shorelines by fouling rocks and littering beaches with its sharp shells. On the other hand, its large filter-feeding biomass increases the clarity of the water (Olenin and Leppakoski 2000).

Ecological Impacts

Specific ecological impacts of Amphibalanus improvisus have not been reported from North American waters. This species is likely to affect fouling communities in the upper regions of estuaries on the Pacific coast, since no native barnacles can tolerate very low salinities, as A. improvisus can.

Competition- In experiments conducted in the western Baltic Sea (Kiel, Germany), manipulations of Amphibalanus improvisus and the Blue Mussel, Mytilus edulis, showed that M. edulis outcompeted and replaced A. improvisus. However, when some mussels were removed, to simulate predation, the two species coexisted and out-competed other fouling community species. Amphibalanus improvisus thus has a sub-dominant role in the fouling community of the Western Baltic (Dürr and Wahl 2004). In the Caspian Sea, A. improvisus reportedly excludes A. eburneus from man-made structures (Zaitsev and Ozturk 2001).

Herbivory, Habitat Change, Trophic Cascade- In the Baltic Sea, where it is the only barnacle species present, filter-feeding by A. improvisus is thought to affect food webs by diverting planktonic production to the benthic biomass (Olenin and Leppakoski 2000). In experiments, settled A. improvisus indirectly promoted the growth of the green alga, Ulva (Enteromorpha) intestinalis, by filtering phytoplankton, remineralizing nutrients, and increasing the clarity of the water (Kotta et al. 2006).

Food/Prey- The larval stages of A. improvisus can be very abundant in the water column during the spawning period, and represents an important food resource for planktivorous fishes and other animals (Skolka and Preda 2010).

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
PAN_PAC Panama Pacific Coast 2008 Def Estab
PAN_CAR Panama Caribbean Coast Native Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
3442 Laguna 1985 1984 1984-01-01 Panama, Caribbean Sea (Colon) Native 9.3592 -79.9014
6043 Bayer et al. 1970, cited by Spivey 1976 1970 1970-01-01 Christobal Native 9.3522 -79.9044
6783 Ruiz et al., unpublished data 2008 2008-01-01 Isla Taboguilla Def 8.8000 -79.5167

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