Amphibalanus improvisus

Invasion History

First Non-native North American Tidal Record: 1853
First Non-native West Coast Tidal Record: 1853
First Non-native East/Gulf Coast Tidal Record:

---

General Invasion History:

Amphibalanus improvisus was first described by Darwin (1854), using specimens collected from England, Scotland, the United States, the West Indies, Argentina, 'West Colombia' and Ecuador. Most of the specimens were found 'attached to wood, shells, rocks, ships' bottoms' (Darwin 1854). It is native to the Atlantic and Gulf Coasts of North America, based on the occurrence of fossils, and ranges south through the Caribbean to South America. Orensanz et al. (2002) consider this barnacle to be cryptogenic in Argentina and Uruguay. It was apparently a very early invader to Europe (possibly in the 1600s), Ecuador, and San Francisco Bay (both by the 1850s).  It spread rapidly through Europe, colonizing the brackish Baltic, Black, and Caspian Seas. After World War II, it was found in Northwest Pacific waters, from Vladivostok to the East China Sea.

North American Invasion History:

Invasion History on the West Coast:

In the Northeast Pacific, Amphibalanus improvisus was first reported from San Francisco Bay in 1853. It was transported in ship fouling from the East Coast of the U.S. or Europe, but oyster transplants and coastal shipping were likely also responsible for subsequent introductions and movements along the coast. A specimen was collected from the Gulf of California in 1889. Other records of established populations in West Coast estuaries are more recent, but likely reflect temporal sampling bias, and this species was almost certainly present in all of these locations for decades earlier (J.T. Carlton, pers. comm., 2013): Willapa Bay,Washington (1955); Esquimalt Lagoon, British Columbia (1956); Puerto Vallarta, Mexico (1960), Columbia River Estuary (1965), Coos Bay, Oregon (1970), and Elkhorn Slough, California (1998) (Henry and McLaughlin, 1975; Carlton 1979; Wasson et al. 2001). The spotty nature of these records probably reflects the preference of this species for sheltered, brackish estuaries. A recent survey of National Estuarine Research Reserves found new records for A. improvisus in the Tijuana Estuary (southern California), a small boat harbor at the mouth of the Quillayute River on Washington's Olympic coast, and Padilla Bay, Washington (deRivera et al. 2005a), but we do not yet know whether these records represent established populations. There are a number of records from marine harbors which do not appear to have resulted in subsequent populations: San Diego Bay (1939); Los Angeles-Long Beach (1932); and Monterey Bay (1916, 1954) (Carlton 1979).

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

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 Habitat	Coarse Woody Debris	None
General Habitat	Oyster Reef	None
General Habitat	Marinas & Docks	None
General Habitat	Rocky	None
General Habitat	Vessel Hull	None
General Habitat	Mangroves	None
General Habitat	Canals	None
Salinity Range	Limnetic	0-0.5 PSU
Salinity Range	Oligohaline	0.5-5 PSU
Salinity Range	Mesohaline	5-18 PSU
Salinity Range	Polyhaline	18-30 PSU
Salinity Range	Euhaline	30-40 PSU
Salinity Range	Hyperhaline	40+ PSU
Tidal Range	Subtidal	None
Tidal Range	Low Intertidal	None
Vertical Habitat	Epibenthic	None

---

---

Tolerances and Life History Parameters

Minimum Temperature (ºC)	-2	Based on range, including ice-covered estuaries.
Maximum Temperature (ºC)	38	Maximum 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 (‰)	0	Adults can survive long exposure to freshwater (Foster 1970)
Maximum Salinity (‰)	40	Rare at high salinities (Furman and Yule 1991), but present in the Mediterranean Sea.
Minimum Reproductive Temperature	10	Bousfield 1955
Maximum Reproductive Temperature	30	Bousfield 1955
Minimum Reproductive Salinity	2	Experimental, lowest tested, 28% metamorphosis, compared to 71-75% at 16-32 ppt (Furman and Yule 1991).
Maximum Reproductive Salinity	40	Experimental, highest tested, 45% metamorphosis, compared to 71-75% at 16-32 ppt, all at 30 C (Furman and Yule 1991).
Minimum Duration	4	Experimental, 30 C, 16-24 ppt, high food density (Furman and Yule 1989).
Maximum Duration	18	15 C, 18 days for intermittently starved larvae (Lang and Marcy 1982)
Maximum Length (mm)	9	Maximum adult height (Henry and McLauglin 1975)
Maximum Width (mm)	17	Maximum adult width (Henry and McLauglin 1975)
Broad Temperature Range	None	Cold temperate-Tropical
Broad Salinity Range	None	Oligohaline-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 Impacts

B-XII	None		Economic Impact	Industry
In the Baltic Sea, where it is the only barnacle species present, A. improvisus is reported to contribute to hull fouling of ships and the fouling of power plants (Vuorinen et al. 1986). Costs of fouling of power plants in Swedish waters were estimated at 10-38 SK (=1.5-5.5 million dollars) per year (Gren et al. 2009).
B-XII	None		Economic Impact	Shipping/Boating
In the Baltic Sea, where it is the only barnacle species present, A. improvisus is reported to contribute to hull fouling of ships and the fouling of power plants (Vuorinen et al. 1986). Costs of fouling of recreational boats in Swedish waters, due to A. improvisus were estimated at 122-331 million Swedish kroner, SK, equal to 8-49 million dollars, per year. Fouling of commercial vessels was estimated at 33-49 SK, equal to 5-7 million dollars, per year, for a total cost of 23-56 million dollars per year (Gren et al. 2009).
B-XI	None		Economic Impact	Industry
In the Baltic Sea, where it is the only barnacle species present, A. improvisus is reported to contribute to hull fouling of ships and the fouling of power plants (Vuorinen et al. 1986). Costs of fouling of powerplants in Swedish waters were estimated at 10-38 SK (=1.5-5.5 million dollars) per year (Gren et al. 2009).
B-XI	None		Economic Impact	Shipping/Boating
In the Baltic Sea, where it is the only barnacle species present, A. improvisus is reported to contribute #to hull fouling of ships and the fouling of power plants (Vuorinen et al. 1986). Fouling of commercial vessels was estimated at 33-49 SK, equal to 5-7 million dollars, per year, for a total cost of 23-56 million dollars per year (Gren et al. 2009). Costs of fouling of recreational boats in Swedish waters, due to A. improvisus, were estimated at 122-331 million Swedish kroner, SK, equal to 8-49 million dollars, per year.
B-X	None		Economic Impact	Industry
In the Baltic Sea, where it is the only barnacle species present, A. improvisus is reported to contribute to hull fouling of ships and the fouling of power plants (Vuorinen et al. 1986). Costs of fouling of powerplants in Swedish waters were estimated at 10-38 SK (=1.5-5.5 million dollars) per year (Gren et al. 2009).
B-X	None		Economic Impact	Shipping/Boating
In the Baltic Sea, where it is the only barnacle species present, A. improvisus is reported to contribute to hull fouling of ships and the fouling of power plants (Vuorinen et al. 1986). Costs of fouling of recreational boats in Swedish waters, due to A. improvisus, were estimated at 122-331 million Swedish kroner, SK, equal to 8-49 million dollars, per year. Fouling of commercial vessels was estimated at 33-49 SK, equal to 5-7 million dollars, per year, for a total cost of 23-56 million dollars per year (Gren et al. 2009).
B-IX	None		Economic Impact	Industry
In the Baltic Sea, where it is the only barnacle species present, A. improvisus is reported to contribute to hull fouling of ships and the fouling of power plants (Vuorinen et al. 1986).
B-IX	None		Economic Impact	Shipping/Boating
In the Baltic Sea, where it is the only barnacle species present, A. improvisus is reported to contribute to hull fouling of ships and the fouling of power plants (Vuorinen et al. 1986).
B-VII	None		Economic Impact	Industry
In the Baltic Sea, where it is the only barnacle species present, Amphibalanus improvisus is reported to contribute to hull fouling of ships and the fouling of power plants (Vuorinen et al. 1986).
B-VII	None		Economic Impact	Shipping/Boating
In the Baltic Sea, where it is the only barnacle species present, Amphibalanus improvisus is reported to contribute to hull fouling of ships and the fouling of power plants (Vuorinen et al. 1986).
NWP-4a	None		Economic Impact	Industry
In Russian waters of the Sea of Japan, A. improvisus is an abundant fouling organism of power plants, docks, and ships in Zolotoi Rog (Golden Horn) Bay in the vicinity of Vladivostok (Zvyagintsev 1985; Kashin et al. 2000; Zvyagintsev et al. 2003).
NWP-4a	None		Economic Impact	Shipping/Boating
In Russian waters of the Sea of Japan, A. improvisus is an abundant fouling organism of power plants, docks, and ships in Zolotoi Rog (Golden Horn) Bay in the vicinity of Vladivostok (Zvyagintsev 1985; Kashin et al. 2000; Koryakova et al. 2002; Zvyagintsev et al. 2004).
B-III	None		Ecological Impact	Competition
In experiments in the western Baltic Sea (Kiel, Germany), manipulations of A. 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).
B-VII	None		Ecological Impact	Herbivory
In experiments, settled Amphibalanus improvisus promoted the growth of the green alga, Enteromorpha intestinalis, by filtering phytoplankton, remineralizing nutrients, and increasing the clarity of the water (Kotta et al. 2006).
B-VII	None		Ecological Impact	Habitat Change
In experiments, settled Amphibalanus improvisus promoted the growth of the green alga, Enteromorpha intestinalis, by filtering phytoplankton, remineralizing nutrients, and increasing the clarity of the water (Kotta et al. 2006). In the central Baltic, A. improvisus was classified as having some habitat impacts, by fouling macroalgae (Bostrom and Bonsdorff 1997, cited by Zaiko et al. 2011).
B-VII	None		Ecological Impact	Trophic Cascade
In experiments, settled Amphibalanus improvisus promoted the growth of the green alga, Enteromorpha intestinalis, by filtering phytoplankton, remineralizing nutrients, and increasing the clarity of the water (Kotta et al. 2006).
B-I	None		Economic Impact	Shipping/Boating
In the Skagerrak, Sweden, A. improvisus was the dominant organism fouling the hulls of recreational boats, probably because it was more tolerant of hydrodynamic stress than the Blue Mussel, Mytilus edulis, which dominated static fouling plates (Berntsson and Jonsson 2003).
CASP	Caspian Sea		Ecological Impact	Competition
'There is a competition between A. improvisus and A. eburneus, as a result of that A. improvisus lives on the hulls of ships and hydrotechnical constructions, but A. eburneus inhabits bottom hard substrates.' (Zaitsev and Ozturk 2001).
CASP	Caspian Sea		Economic Impact	Industry
'Amphibalanus increases the corrosion of metallic constructions in the sea including oil pipes.' (Zaitsev and Ozturk 2001).
CASP	Caspian Sea		Economic Impact	Fisheries
'By covering fish nets Amphibalanus diminishes their catchability and floatation.' (Zaitsev and Ozturk 2001).
CASP	Caspian Sea		Economic Impact	Shipping/Boating
'Mass fouling of Amphibalanus on piers considerably increases wave loading.'(Zaitsev and Ozturk 2001).
MED-IX	None		Ecological Impact	Competition
Skolka and Preda (2010) suggest that the success and abundance of A. improvisus has prevented the settlement of other barnacle species, including introduced A. eburneus, A. amphitrite, and native Balanus crenatus (Skolka and Preda 2010).
MED-IX	None		Ecological Impact	Food/Prey
'Larval stages represent a valuable food resource' (Skolka and Preda 2010).
MED-IX	None		Economic Impact	Shipping/Boating
'Fouling organisms, like barnacles (A. improvisus or the polychaete Ficopomatus enigmaticus), are harmful because their biomass increases the fuel consumption of ships. Such organisms also increase the corrosion rate of metallic submerged structures' (Skolka and Preda 2010).
B-VI	None		Economic Impact	Shipping/Boating
Costs of fouling of recreational boats in Swedish waters, due to A. improvisus, were estimated at 122-331 million Swedish kroner, SK, equal to 18-49 million dollars, per year. Fouling of commercial vessels was estimated at 33-49 SK, equal to 5-7 million dollars, per year, for a total cost of 23-56 million dollars per year (Gren et al. 2009).
B-VI	None		Economic Impact	Industry
Costs of fouling of power plants in Swedish waters were estimated at 10-38 SK (=1.5-5.5 million dollars) per year (Gren et al. 2009).
B-V	None		Economic Impact	Shipping/Boating
Costs of fouling of recreational boats in Swedish waters, due to A. improvisus, were estimated at 122-331 million Swedish kroner, SK, equal to 18-49 million dollars, per year. Fouling of commercial vessels was estimated at 33-49 SK, equal to 5-7 million dollars, per year, for a total cost of 23-56 million dollars per year (Gren et al. 2009).
B-V	None		Economic Impact	Industry
Costs of fouling of power plants in Swedish waters were estimated at 10-38 SK (=1.5-5.5 million dollars) per year (Gren et al. 2009).
B-IV	None		Economic Impact	Shipping/Boating
Costs of fouling of recreational boats in Swedish waters, due to A. improvisus, were estimated at 122-331 million Swedish kroner, SK, equal to 18-49 million dollars, per year. Fouling of commercial vessels was estimated at 33-49 SK, equal to 5-7 million dollars, per year, for a total cost of 23-56 million dollars per year (Gren et al. 2009).
B-IV	None		Economic Impact	Industry
Costs of fouling of power plants in Swedish waters were estimated at 10-38 SK (=1.5-5.5 million dollars) per year (Gren et al. 2009).
B-IX	None		Ecological Impact	Competition
In the Gulf of Finland, A. improvisus was classified as having moderate community impacts (Zaiko et al. 2011).
B-IX	None		Ecological Impact	Habitat Change
In the Gulf of Finland, A. improvisus was classified as having some habitat impacts (Zaiko et al. 2011).
B-IX	None		Ecological Impact	Trophic Cascade
In the Gulf of Finland, A. improvisus was classified as having moderate ecosystem impacts (Zaiko et al. 2011).
B-VII	None		Ecological Impact	Competition
In the Curonian Lagoon, A. improvisus was classified as having some community impacts (Zaiko et al. 2011).
B-XI	None		Ecological Impact	Habitat Change
In the Gulf of Bothnia , A. improvisus was classified as having some habitat impacts, by fouling eelgrass and algae (Bostrom and Bonsdorff 1997; Raberg and Kautsky 2007, cited by Zaiko et al. 2011).
B-XII	None		Ecological Impact	Habitat Change
In the Gulf of Bothnia, A. improvisus was classified as having some habitat impacts, by fouling eelgrass and algae (Bostrom and Bonsdorff 1997 and Raberg and Kautsky 2007, cited by Zaiko et al. 2011).
B-VIII	None		Ecological Impact	Competition
In the Gulf of Bothnia, A. improvisus was classified as having some community impacts (Zaiko et al. 2011).
B-VIII	None		Ecological Impact	Habitat Change
In the Gulf of Riga, A. improvisus was classified as having some habitat impacts (Zaiko et al. 2011).
CASP	Caspian Sea		Ecological Impact	Habitat Change
Fouling by Amphibalanus improvisus reduces the growth rate of the cockle Cerastoderma glaucum in the souther Caspian Sea. Another, deeper-burrowing bivalve, Adacna vitrea was not colonized by A. improvisus (Mirzajan et al. 2016).
P090	San Francisco Bay		Ecological Impact	Competition
Transplant experiments indicate that competition with Amphibalanus improvisus may limit growth and abundance of the introduced bryozoan Conopeum chesapeakensis at the less saline end of Carquinez Strait in the San Francisco estuary (Benicia, ~3.3 PSU, Newcomer et al. 2018).
NEP-V	Northern California to Mid Channel Islands		Ecological Impact	Competition
Transplant experiments indicate that competition with Amphibalanus improvisus may limit growth and abundance of the introduced bryozoan Conopeum chesapeakensis at the less saline end of Carquinez Strait in the San Francisco estuary (Benicia, ~3.3 PSU, Newcomer et al. 2018).
CA	California		Ecological Impact	Competition
Transplant experiments indicate that competition with Amphibalanus improvisus may limit growth and abundance of the introduced bryozoan Conopeum chesapeakensis at the less saline end of Carquinez Strait in the San Francisco estuary (Benicia, ~3.3 PSU, Newcomer et al. 2018)., Transplant experiments indicate that competition with Amphibalanus improvisus may limit growth and abundance of the introduced bryozoan Conopeum chesapeakensis at the less saline end of Carquinez Strait in the San Francisco estuary (Benicia, ~3.3 PSU, Newcomer et al. 2018).

References

Orth, Donald J. (2010) Socrates Opens a Pandora’s Box of Northern Snakehead Issues, American Fisheries Society Symposiums 89: 203-221

Shalovenko, N. N. (2020) Tendencies of Invasion of Alien Zoobenthic Species into the Black Sea, Russian Journal of Biological Invasions 11(2): . 164–171

Aarnio, Katri; Törnroos, Anna; Björklund, Charlotta; Bonsdorff, Erik (2015) Food web positioning of a recent coloniser: the North American Harris mud crab Rhithropanopeus harrisii (Gould, 1841) in the northern Baltic Sea, Aquatic Invasions 10: In press

Aasen, Geir; Sweetnam, Dale A.; Lynch. Lisa M. (1998) Establishment of the wakasagi, Hypomesus nipponensis in the Sacramento-San Joaquin estuary., California Fish and Game 84(1): 31-35

Aladin, Nikolai V.; Plotnikov, Igor S.; Filipov, Andrei A. (2002) Invasive aquatic species of Europe: Distribution, impacts, and management., Kluwer Academic Publishers, Dordrecht. Pp. 351-359

Artuz, M. Levent 2005 First record of two marine invaders.. <missing URL>



Asakura, A. (1992) Recent introductions of marine benthos into Tokyo Bay (review): Process of invasion into an urban ecosystem with discussion on the factors inducing their successful introduction, Journal of the Natural History Museum and Institute, Chiba 2(1): 1-14

Ashton, Gail; Davidson, Ian; Ruiz, Gregory (2014) Transient small boats as a long-distance coastal vector for dispersal of biofouling organisms, Estuaries and Coasts 37: 1572-1581

Bacon, Peter R. (1976) The Cirripedia of Trinidad, Studies of the Fauna of Curacao and other Caribbean Islands 163: 1-55

Bagaveeva, E. V.; Zvyagintsev, A. Yu. (2000) Introduction of polychaetes, Hydroides elegans (Haswell), Polydora limicola Annenkova, and Pseudopotamilla occelata Moore to the Northwestern part of the East Sea, Ocean Research 22(1): 25-36

Baldwin, Andy; Leason, Diane (2016) Potential Ecological impacts of Emerald Ash Borer on Maryland's Eastern Shore, In: None(Eds.) None. , <missing place>. Pp. <missing location>

Bancila. Raluca I.; Skolka, Marius; Ivanova, Petya; Surugiu, Victor; Stefanova, Kremena; Todorova. Valentina; Zenetos, Argyro (2022) Alien species of the Romanian and Bulgarian Black Sea coast: state of knowledge, uncertainties, and needs for future research, Aquatic Invasions 17: Published online

Barnes, Robert D. (1983) Invertebrate Zoology, Saunders, Philadelphia. Pp. 883

Berntsson, Kent; Jonsson, Per R. (2003) Temporal and spatial patterns in recruitment and succession of a temperate marine fouling assemblage: a comparison of static panels and boat hulls during the boating season, Biofouling 19(3): 187-195

Bingham, Brian L.; Young, Craig M. (1991) Larval behavior of the ascidian Ecteinascidia turbinata Herdman; an in situ experimental study of the effects of swimming on dispersal, Journal of Experimental Marine Biology and Ecology 145: 189-204

Bishop, M. W. H. (1947) Establishment of an immigrant barnacle in British coastal waters, Nature 159(4041): 501-502

Bishop, M. W.; Crisp, D. J.; Fischer-Piette, E.; Prenant, M. (1957) Sur l’écologie des Cirripèdes de la côte atlantique française., Bulletin de l'Institut Océanographique, Monaco 1099: 1-12

Bloch. Philip; Novak, Grant (2011) The 2011 International Conference on Ecology & Transportation, International Conference on Ecology & Transportation, <missing place>. Pp. 400-410

Bousfield, E. L. (1953) The distribution and spawning seasons of barnacles on the Atlantic coast of Canada., Bulletin of the National Museum of Canada 132: 112-154

Bousfield, E. L. (1955) Ecological control of the occurrence of barnacles in the Miramichi estuary., Bulletin of the National Museum of Canada 137: 1-69

Box, Antonio; Sureda, Antoni; Tauler, Pere;Terrados, Jorge; Marba, Nuria; Pons, Antoni Deudero, Salud (2010) Seasonality of caulerpenyne content in native Caulerpa prolifera and invasive C. taxifolia and C. racemosa var.cylindracea in the western Mediterranean Sea, Botanica Marina 53: 367-375

Branscomb, E. Sanford (1976) Proximate causes of mortality determining the distribution and abundance of the barnacle Balanus improvisus in Chesapeake Bay., Chesapeake Science 17(4): 281-288

Branstrator, Donn K.; Brown, Meghan E.;Shannon, Lyle J.; Thabes, Marte; Heimgartner, Katie (2006) Range expansion of Bythotrephes longimanus in North America: evaluating habitat characteristics in the spread of an exotic zooplankter., Biological Invasions 8: 1367-1379

Bumbeer, Janaína de Araújo; da Rocha, Rosana Moreira (2012) Detection of introduced sessile species on the near shore continental shelf in southern Brazil, Zoologia 29(2): 126-134

Buschbaum, Christian; Lackschewitz, Dagmar; Reise, Karsten (2012) Nonnative macrobenthos in the Wadden Sea ecosystem, Journal of Ocean Management 68: 89-101

California Department of Fish and Wildlife (2014) Introduced Aquatic Species in California Bays and Harbors, 2011 Survey, California Department of Fish and Wildlife, Sacramento CA. Pp. 1-36

Carlton, James T. (1979) History, biogeography, and ecology of the introduced marine and estuarine invertebrates of the Pacific Coast of North America., Ph.D. dissertation, University of California, Davis. Pp. 1-904

Carlton, James T. (1989) <missing title>, <missing publisher>, <missing place>. Pp. <missing location>

Carlton, James T.; Zullo, Victor A. (1969) Early records of the barnacle Balanus improvisus Darwin from the Pacific coast of North America, Occasional Papers of the California Academy of Sciences 75: 1-6

Carlton, James T.; Newman, William A.; Pitombo, Fábio Bettini (2011) In the wrong place- Alien marine crustaceans: Distribution, biology, impacts, Springer, Dordrecht. Pp. 159-213

Chavanich, S.; Tan, L. T.; Vallejo, B.; Viyakarn, V. (2010) Report on the current status of marine non-indigenous species in the Western Pacific region, Intergovernmental Oceanographic Commission, Subcommission for the Western Pacific, Bangkok, Thailand. Pp. 1-61

Chepalyga, A. L.; Tarasov, A. G. (1997) Invasion of the Atlantic Species into the Caspian Sea, Oceanology 37(2): 237-244

Cohen, A. N. and 11 authors (2005) Rapid assessment survey for exotic organisms in southern California bays and harbors, and abundance in port and non-port areas., Biological Invasions 7: 995-1002

Cohen, Andrew N. and 10 authors (2005) <missing title>, San Francisco Estuary Institute, Oakland CA. Pp. <missing location>

Cohen, Andrew N. and 22 authors (2001) <missing title>, Washington State Department of Natural Resources, Olympia. Pp. <missing location>

Cohen, Andrew N.; Carlton, James T. (1995) Nonindigenous aquatic species in a United States estuary: a case study of the biological invasions of the San Francisco Bay and Delta, U.S. Fish and Wildlife Service and National Sea Grant College Program (Connecticut Sea Grant), Washington DC, Silver Spring MD.. Pp. <missing location>

Crandall, Marc. E. (1977) Epibenthic Invertebrates of Croton Bay in the Hudson River, New York Fish and Game Journal 24(2): 178-196

DAISIE (Delivering Alien Invasive Species Inventories to Europe) (2009) Handbook of alien species in Europe, Springer, Dordrecht, Netherlands. Pp. 269-374

Darwin, Charles (1854) A Monograph on the Sub-class Cirripedia: With Figures of All the Species: The Balanidæ (or Sessile Cirripedes); the Verrucidæ, Etc, The Ray Society, London. Pp. <missing location>

Darwin, Charles (1859) <missing title>, J. Murray, London. Pp. <missing location>

De Mesel, Ilse; Kerckhof, Francis; Norro, Alain; Rumes, Bob; Degraer, Steven (2015) Succession and seasonal dynamics of the epifauna community on offshore wind farm foundations and their role as stepping stones for non-indigenous species, Hydrobiologia 756: 37-50

de Rivera, Catherine, and 27 authors (2005) Broad-scale non-indigenous species monitoring along the West Coast in National Marine Sanctuaries and National Estuarine Research Reserves report to National Fish and Wildlife Foundation, National Fish and Wildlife Foundation, Washington, D.C.. Pp. <missing location>

Dean, Thomas A.; Bellis, Vincent J. (1975) Seasonal and spatial distribution of epifauna in the Pamlico River Estuary, North Carolina, Journal of the Elisha Mitchell Scientific Society 91(1): 1-12

del Monaco, Carlos; Capelo, Juan (2000) [The genera Balanus, Chthamalus and Tetraclita from the coast of Nueva Esparta, Venezuela], Memoria de la Fundacion La Salle de Ciencias Naturales 154: 77-197

Dineen, Joseph Jr.; Hines, Anson H. (1992) Interactive effects of salinity and adult extract upon settlement of the estuarine barnacle Balanus improvisus., Journal of Experimental Marine Biology and Ecology 156: 239-252

Durr, Simone; Wahl, Martin (2004) Isolated and combined impacts of the blue mussels Mytilus edulis and barnacles, Balanus improvius, on structure and diversity of a fouling community., Journal of Experimental Marine Biology and Ecology 306: 181-195

Dziubinska, Anna; Janas, Urszula (2007) Submerged objects: a nice place to live and develop. Succession of fouling communities in the Gulf of Gdansk, Southern Baltic, Oceanological and Hydrobiological Studies 36(4): 65-78

Eaton, Lawrence (1994) Preliminary survey of benthic macroinvertebrates of Currituck Sound, North Carolina, Journal of the Elisha Mitchell Scientific Society 110(3/4): 121-129

Faasse, Marco (2012) The exotic isopod Synidotea in the Netherlands and Europe, A Japanese or American invasion (Pancrustacea: Isopoda)?, Nederlandse Faunistiche Mededelingen 108: 103-106

Farrapeira, Christiane M. R. (2010) Barnacles (Crustacea:Cirripedia) of the estuarine and marine areas of the port of Recife (Pernambuco, Brazil), Cahiers de Biologie Marine 50: 199-129

Farrapeira, Cristiane Maria Rocha (2008) [Cirripedia Balanomorpha of Paripe River estuary (Itamaracá Island, Pernambuco, Brazil).], Biota Neotropica 8(3): 31-39

Farrapeira, Cristiane Maria Rocha (2010) Shallow water Cirripedia of the northeastern coast of Brazil: The impact of life history and invasion on biogeography, Journal of Experimental Marine Biology and Ecology 392: 210-219

Foster, B. A. (1970) Responses and acclimation to salinity in the adults of some balanomorph barnacles, Philosophical Transactions of the Royal Society of London B 256(810): 377-400

Foster, B. A., Willan, R. C. (1979) Foreign barnacles transported to New Zealand on an oil platform., New Zealand Journal of Marine and Freshwater Research 13(1): 143-149

Furlani, Dianne M. (1996) A guide to the introduced marine species in Australian waters., In: (Eds.) . , Hobart, Australia. Pp. <missing location>

Furman, E. R.; Yule, A. B. (1991) Estuaries and coasts: Spatial and temporal intercomparisons., Olsen and Olsen, Fredensborg, Denmark. Pp. Pp. 273-276.

Furman, Eeva (1989) Enzyme genetic variation in Balanus improvisus Darwin (Crustacea, Cirripedia) in the Baltic Sea., Ophelia 30(1): 35-45

Furman, Eeva (1990) Geographical variation of Balanus improvisus in biochemical and morphometric characters., Journal of the Marine Biological Association of the United Kingdom 70: 721-740

Gartner, Heidi N.; Murray, Cathryn Clarke; Frey, Melissa A.; Nelson, Jocelyn C.; Larson, Kristen J.; Ruiz, Gregory M.; Therriault, Thomas W. (2016) Non-indigenous invertebrate species in the marine fouling communities of British Columbia, Canada, BioInvasions Records <missing volume>: <missing location>

Gislen, Torsten (1950) {On the invasion and distribution of Balanus improvisus along the coast of Sweden.], Fauna och Flora 45: 32-37

Godhe, Anna, Wallentinus, Inger (1999) <missing title>, Nordic Council of Ministers, Copenhagen. Pp. 140-183

Gomiou, Marian-Traian; Alexandrov, Boris; Shadrin, Nikolai; Zaitsev, Yuvenaly (2002) The Black Sea- a recipient, donor, and transit area for alien species., In: Leppakoski, E.; Gollasch, S.; Olenin, S.(Eds.) Invasive aquatic species of Europe: Distribution, impacts, and management.. , Dordrecht. Pp. 341-350

Goulletquer, Philippe; Bachelet, Guy; Sauriau, Pierre; Noel, Pierre (2002) Invasive aquatic species of Europe: Distribution, impacts, and management, Kluwer Academic Publishers, Dordrecht. Pp. 276-290

Graham, Herbert W.; Gay, Helen (1945) Seasonal attachment and growth of sedentary marine organisms at Oakland, California., Ecology 26(4): 375-386

Gren, Ing-Marie; Isacs, Lina; Carlsson, Mattias (2009) Costs of alien invasive species in Sweden, Ambio 38(3): 135-140

Grigorovich, Igor A.; Therriault, Thomas W.; MacIsaac, Hugh J. (2003) History of aquatic invertebrate invasions in the Caspian Sea., Biological Invasions 5: 103-115

Henry, Dora P.; McLaughlin, Patsy A. (1975) The barnacles of the Balanus amphitrite complex (Cirripedia, Thoracica)., Zoologische Verhandelingen 141: 1-203

Holm, Eric R. (2012) Barnacles and biofouling, Integrative and Comparative Biology Published online: <missing location>

Holmes, Samuel; Callaway, Ruth (2021) Fouling communities and non-native species within five ports along the Bristol Channel, South Wales, UK, Estuarine Coastal and Shelf Science 252(1107295): Publiished online

Holopainen, Reetta; Lehtiniemi, Maiju; Meier, H. E. Markus; Albertsson, Jan; Gorokhova, Elena; Kotta, Jonne; Viitasalo, Markku (2016) Impacts of changing climate on the non-indigenous invertebrates in the northern Baltic Sea by end of the twenty-first century, Biological Invasions Published online: <missing location>

Houghton, D.R. (1971) Barnacles and fouling., In: Gareth Jones, E.B. & Eltringham, S.K.(Eds.) Marine borers, fungi and fouling organisms of wood. , Paris. Pp. <missing location>

Iwasaki, Keiji (2006) Assessment and Control of Biological Invasion Risks., Shoukadoh Book Sellers,and IUCN, Kyoto and Gland, Switzerland. Pp. 104-112

Jablonska-Barna, Izabela; Rychter, Agata; Kruk, Marek (2013) Biocontamination of the western Vistula Lagoon (south-eastern Baltic Sea, Poland), Oceanologia 53(3): 751-763

Jazdzewski, Kryzysztof; Grabowski, Michal (2011) In the wrong place- Alien marine crustaceans: Distribution, biology, impacts, Springer, Dordrecht, Netherlands. Pp. 323-344

Jensen, Kathe R.; Knudsen, Jorgen (2005) A summary of alien marine benthic invertebrates in Danish waters., Oceanological and Hydrobiological Studies 34 (suppl. 1): 137-161

Jones, Diana S.; Hewitt, Melissa A.; Sampey, Alison (2000) Checklist of the Cirripedia of the South China Sea, Raffles Bulletin of Zoology Supplement 8: 233-307

Kashin, I. A.; Zvyagintsev, A. Yu;. Maslennikov, S. I. (2000) Fouling of hydrotechnical structures in western Peter the Great Bay, Sea of Japan., Russian Journal of Marine Biology 26(2): 89-97

Kasymov, A.G. (1982) The role of Azov-Black Sea invaders in the productivity of the Caspian Sea benthos, Internationale Revue der Gesamten Hydrobiologie 67(4): 533-541

Kennedy, Victor S.; DiCosimo, Jane (1983) Subtidal distribution of barnacles (Cirripedia: Balanidae) in Chesapeake Bay, MD., Estuaries 6(2): 95-101

Kerckhof, Francis; Haelters, Jan; Gollasch, Stephan G. (2007) Alien species in the marine and brackish ecosystem: the situation in Belgian waters., Aquatic Invasions 2(3): 243-257

Kerckhof, Francis; Cattrijsse, Andre (2001) Exotic Cirripedia (Balanomorpha) from buoys off the Belgian Coast., Senckenbergiana Maritima 31(2): 245-254

Kim, I-H. (1992) Invasion of foreign barnacles into Korean waters, Korean Journal of Systematic Zoology 8(2): 163-176

Kocak, F. ; Kucuksezgin, F. (2000) Sessile fouling organisms and environmental parameters in the marinas of the Turkish Aegean coast., Indian Journal of Marine Science 29: 149-157

Korn, O.M. (1999) Distribution of cirripede larvae in Nakhodka Bay, Sea of Japan., Russian Journal of Marine Biology 25(5): 395-402

Koryakova, M. D.; Nikitin, V. M.; Zvyagintsev, A. Yu.; Belogurova, L. S. (2002) Effect of polluted harbor waters on fouling and biocorrosion of steel, Russian Journal of Marine Biology 28(2): 127-131

Kotta, Jonne and 6 authors (2006) Ecological consequences of biological invasions: three invertebrate case studies in the north-eastern Baltic Sea., Helgoland Journal of Marine Research 60: 106-112

Koukouras, Anathasios; Matsa, Athanasia (1998) The thoracican cirriped fauna of the Aegean Sea: New information, checklist of the Mediterranean species, faunal comparisons., Senckenbergiana Maritima 28(4/6): 133-142

Laguna, Jorge (1985) <missing title>, M.S. Thesis, University of California, San Diego. Pp. <missing location>

Lang, W. H. (1980) Cirripedia: balanomorph nauplii of the NW Atlantic shores, Fiches D’Identification du Zooplancton 163: 1-6

Lang, William H. (1979) Larval development of shallow water barnacles of the Carolinas (Cirripedia: Thoracica) with keys to naupliar stages., NOAA Technical Report NMFS Circular 421: 1-39

Lang, William H.; Marcy, Martha (1982) Some effects of early starvation on survival and development of barnacle nauplii Balanus improvisus, Journal of Experimental Marine Biology and Ecology 60: 63-70

Larsen, Peter F. (1985) The benthic fauna associated with the oyster reefs of the James River estuary, Virginia, U. S. A., Internationale Revue der Gesamten Hydrobiologie 70(9): 707-814

Laudien, Jurgen; Wahl, Martin (1999) Indirect effects of epibiosis on host mortality: Seastar predation on differently fouled mussels, Marine Ecology 20(1): 24-36

Leppakoski, Erkki; Olenin, Sergei (2000) Non-native species and rates of spread: lessons from the brackish Baltic Sea., Biological Invasions 2: 151-163

Levings, C. D.; Cordell, J. R.; Ong, S.; Piercey, G. E. (2004) The origin and identity of invertebrate organisms being transported to Canada's Pacific coast by ballast water., Canadian Journal of Fisheries and Aquatic Science 61: 1-11

Li, Xinzhengl Shenz. Anglv; Xu, zhaou (2008) Report on some shrimos from dontou Island, Zhejiang, China (Decapoda, Natantia, Crustaceana (Leiden) <missing volume>: <missing location>

Liu, Wenliang; Liang, Xiaoli ; Zhu, Xiaojing (2015) A new record and mitochondrial identification of Synidotea laticauda Benedict, 1897 (Crustacea: Isopoda: Valvifera: Idoteidae) from the Yangtze Estuary, China, Zootaxa 4294: 371-380

Llansó, Roberto J.; Sillett, Kristine; Scott, Lisa (2011) <missing title>, Versar, Inc., Columbia MD. Pp. <missing location>

Lozano-Cortés, Diego F.; Londoño-Cruz, Edgardo (2013) Checklist of barnacles (Crustacea; Cirripedia: Thoracica) from the Colombian Pacific, Marine Biodiversity 43: 463-471

Lu, L.; Levings, C. D.; Piercey, G. E. (2007) Preliminary investigation on aquatic invasive species of marine and estuarine macroinvertebrates on floating structures in five British Columbia harbours, Canadian Manuscript Report of Fisheries and Aquatic Sciences 2814: 2-30

Lutaenko, Konstantin A.; Furota,Toshio; Nakayama, Satoko; Shin, Kyoungsoon; Xu, Jing (2013) <missing title>, Northwest Pacific Action Plan- Data and Information Network Regional Activity Center, Beijing, China. Pp. <missing location>

Luther, A. (1950) On the invasion and distribution of Balanus improvisus along the Swedish coast, Fauna och Flora 45: 155-160

McPherson, Benjamin, Sonntag, Wayne H.; Sabanskas, Maryann (1984) Fouling community of the Loxahatchee River estuary, Florida, 1980-81, Estuaries 7(2): 149-157

Meng, Stefan; trahl, Jaqueline; \ Bormer, Andreas; Krienke, Kay; Zettler, Michael L.; Wroznya, Claudia (2024) The bay barnacle Amphibalanus improvisus (Darwin, 1854) in the Pleistocene of Europe? A review of Pleistocene Balanidae of northern Central Europe, Journal of Quaternary Science <missing volume>: Published online
DOI: 10.1002/jqs.3603

Minchin, Dan; Nunn, Julia (2014) The invasive brown alga Undaria pinnatifida (Harvey) Suringar, 1873 (Laminariales: Alariaceae), spreads northwards in Europe, BioInvasions Records 3: in press

Montalvo-Urgel, Hugo; Sánchez, Alberto J.; Florido, Rosa; Macossay-Cortez, Alberto A. (2010) [List of crustaceans distributed in submerged woody debris in the tropical wetlands of Pantanos de Centla, southern Gulf of Mexico], Revista Mexicana de Biodiversidad 81: S121- S131

Moore, H. B.; Frue, A. C. (1959) The settlement and growth of Balanus improvisus, B. eburneus, and B. amphitrite in the Miami area., Bulletin of Marine Science of the Gulf and Caribbean 9(4): 421-440

Moore, Hilary B.; Albertson, Helen D.; Miller, Sigmund M. (1974) Long-term changes in the settlement of barnacles in the Miami area., Bulletin of Marine Science 24: 86-100

Naser, Murtada D.;Rainbow, Philip S.; Clark, Paul F.; Yasser, Amaal Gh.; Jones, Diana S. (2015) The barnacle Amphibalanus improvisus (Darwin, 1854), and the mitten crab Eriocheir: one invasive species getting off on another!, BioInvasions Records 4: In press

Nasrolahi, Ali; Pansch, Christian; Lenz, Mark; Wahl, Martin (2012) Being young in a changing world: how temperature and salinity changes interactively modify the performance of larval stages of the barnacle Amphibalanus improvisus, Marine Biology 159: 331-340

Nauman, J. W.; Cory, R. L. (1969) Thermal additions and epifaunal organisms at Chalk Point, Maryland., Chesapeake Science 10(3-4): 218-226

Nehring, Stefan (2006) Four arguments why so many alien species settle into estuaries, with special reference to German River Elbe., Helgoland Marine Research 60: 127-134

Newman, William A. (2007) The Light and Smith Manual: Intertidal Invertebrates from Central California to Oregon, University of California Press, Berkeley CA. Pp. 475-481

Noel, Pierre Y. (2011) In the wrong place- Alien marine crustaceans: Distribution, biology, impacts, Springer, Dordrecht, Netherlands. Pp. 345-375

Norris, James N. (2010) Marine Algae of the northern Gulf of California: Chlorophyta and Phaeophyceae, Smithsonian Contributions to Botany 94: 1276

Nunn, Julia; Minchin, Dan 2013 Marine non-native invasive species in Northern Ireland. <missing URL>



Olenin, S., Leppakoski, E. 2000-2016 Inventory of Baltic Sea alien species. <missing URL>



Orensanz, Jose Maria and 14 other authors (2002) No longer the pristine confines of the world ocean: a survey of exotic marine species in the southwestern Atlantic, Biological Invasions 4(1-2): 115-143

Ortiz, Osvaldo Gomez (1987) Dos neuvos registros de cirripedios (Crustacea) para Cuba., Revista da Investicaciones Marinas 8(3): 99-100

Pansch, Christian; Schlegel, Peter; Havenhand, Jonathan (2013) Larval development of the barnacle Amphibalanus improvisus responds variably but robustly to near-future ocean acidification, ICES Journal of Marine Science 70(4): 805-811

Partaly, E. M. (1979) Ecology of bryozoan foulings in the Sea of Azov, Soviet Journal of Marine Biology 5(3): 213-216

Partaly, Ye. M. (2003) Peculiarities of successions in biocenoses of fouling in the Sea of Azov, Hydrobiological Bulletin 39(1): 103-112

Pavlova, E. V.; Murina, V. V.; Kemp, R. B.; Wilson, J. G. ; Parchevsky, V. P. (2007) Annual dynamics of abundance, biomass and survival of meroplankton in Sevastopol Bay, Black Sea, Morskii Ekologicheskii Zhurnal 6(2): 63-78

Pearce, John B. (1974) Invertebrates of the Hudson River estuary, Annals of the New York Academy of Sciences 250: 137-173

Pecarevic, M.; Mikus, J.; Cetinic, A. Bratos; Dulcic, J.; Calic, M. (2013) Introduced marine species in Croatian waters (Eastern Adriatic Sea), Mediterranean Marine Science 14(1): 224-237

Pilsbry, Henry A. (1916) The sessile barnacles contained in the collections of the U.S. National Museum, including a monograph of the American species., United States National Museum Bulletin 93: 1-366

Pitombo, F. B. (2004) Phylogenetic analysis of the Balanidae (Cirripedia, Balanomorpha)., Zoologica Scripta 33(3): 261-276

Pyefinch, K. A. (1950) Notes on the ecology of ship-fouling organisms., Journal of Animal Ecology 19: 29-35

Reish, Donald J.; Gerlinger, Thomas V.; Ware, Robert R. (2018) Comparison of the polychaetous annelids populations on suspended test panels in Los Angeles Harbor in 1950-1951 with the populations in 2013-2014, Bulletin of the Southern California Academy of Sciences 17(1): 82-90

Relini, G; Matricardi, G, (1999) Cirripedi toracici delle Lagune Di Orbetello., Atti della Societa Toscana di Scienze Naturali, series B. 86: 55-57

Riedel, Frank; Audzijonyte, Asta; Mugue, Nikolai (2006) Aliens associating with Caspian Sea endemic bivalves., Biological Invasions 8: 1067-1071

Rodríguez-Almaraz, Gabino A.; García-Madrigal, María del Socorro (2014) [Aquatic Invasive Species in Mexico], Comisión Nacional para el Conocimiento y Uso de la Biodiversidad, <missing place>. Pp. 337-371

Rubal, Marcos; Maldre, Kelli; ,Sousa-Pinto, Isabel; Veiga, Puri (2021) Current distribution and abundance of Austrominius modestus (Darwin, 1854) and other non-indigenous barnacles along the northern coast of Portugal, Regional Studies in Marine Science 41(101586): Published online

Ruiz, Gregory M.; Geller, Jonathan (2018) Spatial and temporal analysis of marine invasions in California, Part II: Humboldt Bay, Marina del Re, Port Hueneme, and San Francisco Bay, Smithsonian Environmental Research Center & Moss Landing Laboratories, Edgewater MD, Moss Landing CA. Pp. <missing location>

Ruiz, Gregory; Geller, Jonathan (2021) Spatial and temporal analysis of marine invasions: supplemental studies to evaluate detection through quantitative and molecular methodologies, Marine Invasive Species Program, California Department of Fish and Wildlife, Sacramento CA. Pp. 153 ppl.

Scuchert, Peter (2010) The European athecate hydroids and their medusae (Hydrozoa, Cnidaria): Capitata Part 2, Revue Suisse de Zoologie 117(3): 337-355

Shalaeva, E. A.; Lisitiskaya, E. V. (2004) Seasonal dynamics of the number of larvae of common fouling organisms in Balaklava Bay (Black Sea)., Russian Journal of Marine Biology 30(6): 371-378

Skolka, Marius; Preda, Cristina (2010) Alien invasive species at the Romanian Black Sea coast: present and perspectives, Travaux du Muséum National d’Histoire Naturelle «Grigore Antipa» 53: 443-467

Sliwa, Cathryn; Migus, Sasha; McEnnulty, Felicity; Hayes, Keith R. (2009) Biological Invasions in Marine Ecosystems., Springer-Verlag, Berlin Heidelberg. Pp. 425-437

Spivey, H. R. (1976) The cirripeds of the Panama Canal, Corrosion and Marine Fouling 1(1): 43-50

Sylvester, Francisco and 8 authors (2011) Hull fouling as an invasion vector: can simple models explain a complex problem?, Journal of Applied Ecology 48: 415-423

Sytsma, Mark D.; Cordell, Jeffrey R.; Chapman, John W.; Draheim, Robyn, C. (2004) <missing title>, Center for Lakes and Reservoirs, Portland State University, Portland OR. Pp. <missing location>

Thompson, Michelle Lynne (1993) <missing title>, M.S. Thesis, College of William and Mary, Williamsburg VA. Pp. <missing location>

U.S. National Museum of Natural History 2002-2021 Invertebrate Zoology Collections Database. http://collections.nmnh.si.edu/search/iz/



Utinomi, Huzio (1970) Studies on the Cirripedian Fauna of Japan.IX., Distributional Survey of Thoracic Cirripeds in the Southeastern part of the Japan Sea, Publications of the Seto Marine Biological Laboratory 17(5): 339-372

Vuorinen, Ilppo; Laihonen, Pasi; Lietzén, Erkki (1986) Distribution and abundance of invertebrates causing fouling in power plants on the Finnish coast, Memoranda Societatis pro Fauna et Flora Fennica 62: 123-125

Ware, Chris et al. (2015) Biological introduction risks from shipping in a warming Arctic, Journal of Applied Ecology Published online: <missing location>

Wasson, Kerstin; Zabin, C. J.; Bedinger, L.; Diaz, M. C.; Pearse J. S. (2001) Biological invasions of estuaries without international shipping: the importance of intraregional transport, Biological Conservation 102: 143-153

Wells, Harry W. (1966) Barnacles of the northeastern Gulf of Mexico., Quarterly Journal of the Florida Academy of Sciences 29(4): 81-95

Wittfoth, Anne K. J.; Zettler, Michael L. (2013) The application of a Biopollution Index in German Baltic estuarine and lagoon waters, Management of Biological Invasions 4: in press

Wolff, W. J. (2005) Non-indigenous marine and estuarine species in the Netherlands., Zoologische Verhandelingen 79(1): 1-116

Wonham, Marjorie J.; Carlton, James T. (2005) Trends in marine biological invasions at local and regional scales: the Northeast Pacific Ocean as a model system, Biological Invasions 7: 369-392

Woods Hole Oceanographic Institution, United States Navy Dept. Bureau of Ships (1952) Marine fouling and its prevention., United States Naval Institute., Washington, D.C.. Pp. 165-206

Wrange, Anna-Lisa; André, Carl; Lundh, Torbjörn; Lind, Ulrika; ; Blomberg, Anders; Jonsson, Per J.; Havenhand, Jon N. (2014) Importance of plasticity and local adaptation for coping with changing salinity in coastal areas: a test case with barnacles in the Baltic Sea, BMC Evolutionary Biology 14(156): published online

Wrange, Anna-Lisa; Charrier, Gregory; Thonig, Anne; Rosenblad, Magnus Alm; Blomberg, Anders; Havenhand, Jonathan N.; Jonsson, Per R.; André, Carl (2016) The story of a hitchhiker: population genetic patterns in the invasive barnacle Balanus (Amphibalanus) improvisus Darwin 1854, PLOS ONE 11(1): e0147082.

Yamanishi, Ryohei; Yokoyama, Hisashi; Ariyama, Hiroyukii (1991) Distributional and seasonal changes of intertidal attaching organisms in relation to water quality along the brackish reaches of the Yodo River., Occasional Papers from the Osaka Museum of Natural History 2(7): 83-96

Young, Paulo S. (1994) Superfamily Balanoidea Leach (Cirripedia, Balanomorpha) from the Brazilian coast., Boletim do Museo Nacional (Zoologia) 356: 1-36

Young, Paulo S. 1998 Maxillopoda. Thecostraca.. <missing URL>



Zaiko, Anastasija; Lehtiniemi, Maiju; Narscius, Aleksas; Olenin, Sergej (2011) Assessment of bioinvasion impacts on a regional scale: a comparative approach, Biological Invasions 13: 1739-1765

Zaitsev, Yuvenali; Ozturk, Bayram (2001) <missing title>, Turkish Marine Research Foundation Publication, <missing place>. Pp. 1-265

Zevina, G. B.; Goryn, A. N. (1971) Invasion of Balanus improvisus and B. eburneus into the Sea of Japan., Zoologicheskii Zhurnal 50(5): 771-773

Zevina, G. B.; Poltarukha, O. P. (1999) [Barnacles of the Black Sea), Bulletin of the Moscow Society of Naturalists 104(1): 30-39.

Zevina, G. B.; Zvyagintsev, A. Yu.; Negashev, C. Z. (1992) <missing title>, [Institute of Marine Biology], Vladivostok. Pp. <missing location>

Zullo, Victor A. (1966) Thoracic cirripedia from the continental shelf of South Carolina USA., Crustaceana 11(3): 229-244

Zullo, Victor A. (1979) Marine flora and fauna of the Northeastern United States. Arthropoda: Cirripedia, NOAA Technical Report NMFS Circular 425: 1-29

Zullo, Victor A., Miller, William, III (1986) Barnacles (Cirripedia: Balanidae) from the lower Pleistocene James City formation, North Carolina coastal plain, with the description of a new species of Balanus da costa, Proceedings of the Biological Society of Washington 99(4): 717-730

Zullo, Victor August (1963) <missing title>, University of California, Berkeley CA. Pp. 1-368

Zvyaginstev, A. Yu.; Radashevsky, V. I.; Ivin, V. V.; Kashin, I. A.; Gorodkov, A. N. (2011) Nonindigenous species in the far-eastern seas of Russia, Russian Journal of Biological Invasions 2(1): 164-182

Zvyagintsev, A. Yu . (1985) Succession of fouling communities of a ship during one navigation in Peter the Great Bay (the Sea of Japan), Soviet Journal of Marine Biology 21(3): 134-140

Zvyagintsev, A. Yu;. Korn, O. M.; Kulikova, V. A. (2004) Seasonal dynamics of pelagic larvae and settling of the fouling organisms in conditions of thermal pollution., Russian Journal of Marine Biology 30(4): 266-277

Zvyagintsev, A. Yu. (2003) Introduction of species into the Northwestern Sea of Japan and the problem of marine fouling., Russian Journal of Marine Biology 29(Suppl. 1:): 10-21

Zvyagintsev, Yu. A. (1992) Seasonal changes of epifauna on valves of giant oysters in Amur Bay, Sea of Japan, Russian Journal of Marine Biology 17(2): 71-76

---