Amphibalanus eburneus

Invasion History

First Non-native North American Tidal Record: 2000
First Non-native West Coast Tidal Record: 2000
First Non-native East/Gulf Coast Tidal Record:

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General Invasion History:

Amphibalanus eburneus is native to the Western Atlantic, from the southern Gulf of Maine to Panama, Colombia and Venezuela. It ranges further south, to Uruguay and Argentina (Henry and McLaughlin 1975; Young 1994), where it may be cryptogenic. Based on its range, this barnacle is moderately tolerant of brackish waters and cold temperatures.  Amphibalanus eburneus may have reached the Pacific through the Panama Canal. It was collected on the Pacific coast of northern Colombia in 1924 (USNM 59208, U.S. National Museum of Natural History 2013) and is now common on the Pacific coast of Panama (Laguna 1985). It was collected in the Gulf of California, Mexico, by 1959 (Henry and McLaughlin 1975). In 2000, an established population was reported for the first time in U.S. Pacific waters, in the Colorado Lagoon, Long Beach, California (Cohen et al. 2002; Carlton, personal communication), and in 2010, it was found at two marinas in San Francisco Bay, California (Foss 2011; Ruiz et al. unpublished data). Based on its broad tolerances, this species has the potential to greatly expand its range on the Pacific coast.

North American Invasion History:

Invasion History on the West Coast:

In July 2000, A. eburneus was collected in the Colorado Lagoon, part of Alamitos Bay in Long Beach, California. This population is well-established (Cohen et al. 2002; Cohen et al. 2005). Amphibalanus eburneus was established at the Pacific end of the Panama Canal and at several locations on the Mexican coast by 1964 (Matsui et al. 1964; Henry and McLaughlin 1975). The California population may have been transported from Mexican harbors by yachts.

Invasion History in Hawaii:

Amphibalanus eburneus was first collected in the Hawaiian Islands in Pearl Harbor, Oahu, in 1929 (Henry and McLaughlin 1975). On Oahu, it has been collected in Honolulu Harbor (in 1947, Henry and McLaughlin 1975) and Kaneohe Bay (in 1959, Henry and McLaughlin 1975). It is also established in Maui (in 1962, Matsui et al. 1964) and Kauai (in 2002, Coles et al. 2004). Amphibalanus eburneus is most abundant in brackish water, and is rare outside harbors and estuaries (Zabin et al. 2009). Carlton and Eldredge (2009) review its history in Hawaii and provide additional details. 

Invasion History Elsewhere in the World:

Amphibalanus eburneus has been widely introduced around the world by shipping, and has invaded the Northeast Atlantic, the Indian Ocean, Northwestern Pacific and the Northeastern Pacific (Barnes and Barnes 1972; Henry and McLaughlin 1975; Utinomi 1975; Laguna 1985). On the Atlantic Coast, it became established in the Bay of Biscay, France by 1940 (Barnes and Barnes 1972; Golletquer et al. 2002). Further north, it was first collected in Dutch waters as early as 1895, but sporadically appeared and disappeared, and does not seem to be established there (Wolff 2005). Amphibalanus eburneus has become established in the seas of southern Europe, entering the Mediterranean by 1863, and reaching the Black Sea by 1895 (Relini and Matricardi 1999; Kocak and Kucuksezgin 2000; Gomiou et al. 2002; Innocenti 2006). This barnacle has colonized the Caspian Sea via canals and ballast water (Grigorevich et al. 2003), but has also invaded oceanic islands, including Bermuda (in 1962, Henry and McLaughlin 1975) and the Azores (Southward 1998).

Amphibalanus eburneus was collected in Manila, the Philippines, before 1916 (Pilsbry 1916), but its expansion into the Northwest Pacific was not documented until after World War II. It appeared in Tokyo Bay in 1950 (Utinomi 1970), and eventually spread north to Vladivostok, Russia and south to Hong Kong (Utinomi 1975; Rainbow 1990, Kim 1992; Zvyaginstsev 2003). Amphibalanus eburneus was collected at Enewetak Atoll in the Marshall Islands in 1967 (Zullo et al. 1972), possibly brought there during World War II or during post-war atomic bomb tests. It was first collected on Guam in 2000, where it was found in an estuary and near freshwater seeps (Paulay et al. 2002; Paulay and Ross 2003). It has also become established at several points along trade routes between Europe and Asia, including the Suez Canal (in 2003, Emara and Belal 2004) and in Mumbai and Madras, India (Daniel 1955 in Matsui et al. 1964; Henry and McLaughlin 1975).

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Description

Amphibalanus eburneus has a shell which varies from conical to cylindrical, depending on the amount of crowding. The orifice is round or slightly toothed, and its width is usually more than ½ its height. The plates have wide longitudinal spaces (radii), narrowing towards the top of the shell plates, while the tops (summits) of the shell plates are thick and rough (Henry and McLaughlin 1975). Inside the operculum, the scutum has thick growth ridges. The tergum has a blunt apex, and a broad spur, with a length only a little greater than its width. The basal margin of the tergum curves inward beside the spur, and then curves outward to form a protuberance with prominent ridges, forming a jagged edge (Henry and McLaughlin 1975). This barnacle can grow to 40 mm diameter and 30 mm height (Henry and McLaughlin 1975), but adults more typically reach 25 mm basal diameter (Gosner 1978). It is characteristic of sheltered estuarine habitats, and tolerates considerable salinity variation (Henry and McLaughlin 1975). The larval stages are described and illustrated in Lang (1979) and Lang (1980).

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Taxonomy

Ecology

General:

Amphibalanus eburneus, like many other barnacles, 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). Broods of A. eburneus in culture ranged from ,1000 ro 13,000 eggs (El-Komy and Kajihara 1991). 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 (Scheltema et al. 1982). 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 that gather phytoplankton, zooplankton, and detritus. Amphibalanus eburneus has seven larval stages, as in other Thoracica: a non-feeding nauplius I, feeding nauplius stages II-VI, and a nonfeeding cyprid, the settling stage (Costlow and Boukhout 1957). Larval development period is based on laboratory experiments with larvae from Woods Hole, Massachusetts. The longest larval development, 18 days, was observed at 20ºC with a low food concentration, the shortest, four days, at 30ºC with high food availability (Scheltema and Williams 1982).

Based on its native range, Amphibalanus eburneus survives in estuaries prone to some winter ice cover, and also survives warm tropical temperatures (Henry and McLaughlin 1975). Its requirement for a temperature of ~20+ºC for larval development (Scheltema et al. 1992) may limit its range. This is suggested by its sporadic appearances in shallow, warm habitats at the northern edges of its range in the Sea of Japan (Zvyaginstsev 2003), the Netherlands (Wolff 2005) and Atlantic France (Barnes and Barnes 1972). Carlton et al. (2011) predict that with climate change, A. eburneus will move north into Maine. Amphibalanus eburneus can be found in marine and estuarine environments from the shallow subtidal to 37 m. Individuals settle on mangroves, mollusk shells, wood, rocks, harbor installations and ships (Wells 1966; Henry and McLaughlin 1975; Laguna 1985).

In a survey of barnacles in the upper and middle Chesapeake Bay, Amphibalanus eburneus was collected at salinities as low as 8 PSU, but was the least abundant species (Kennedy and de Cosimo 1983). This barnacle is most abundant in lower portions of the Bay (Ruiz et al., unpublished data). In a Trinidad mangrove swamp, A. eburneus occurred and reproduced at 6-40 ppt (Bacon 1971). Settlement of cyprids from Chesapeake Bay occurred at salinities of 5-35 ppt (Dineen and Hines 1994). In the Loxahatchee River estuary, Indian Lagoon, Florida, A. eburneus was common at sites with average salinities of 22-32 ppt, and rare at average salinities as low as 19 ppt (SEM ± 3 ppt) (McPherson et al. 1984).

Food:

Phytoplankton

Competitors:

Other barnacles

Trophic Status:

Suspension Feeder

SusFed

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Habitats

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

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	-2	Based on occurrence in estuaries with winter ice cover.
Minimum Salinity (‰)	6	Experimental, lowest tested Trinidad (Bacon 1971)
Maximum Salinity (‰)	70	Field, Laguna Madre TX (Simmons 1957)
Minimum Reproductive Temperature	20	Experimental. larval survival, lowest tested (Scheltema et al. 1982).
Maximum Reproductive Temperature	30	Experimental. larval survival, highest tested (Scheltema et al. 1982).
Minimum Reproductive Salinity	5	Settlement of cyprids in laboratory experiments (Dineen and Hines 1994).
Maximum Reproductive Salinity	45	Field, Laguna Madre TX (Simmons 1957); Successful development, at 40 PSU highest salinity tested (Bacon 1976).
Minimum Duration	4	Experimental, larval development to cypris, 30 C, high food (Scheltema et al. 1982).
Maximum Duration	18	Experimental, larval development to cypris, 20 C, low food (Scheltema et al. 1982).
Maximum Length (mm)	30	Maximum adult height (Henry and McLaughlin 1975)
Maximum Width (mm)	40	Maximum basal height (Henry and McLaughlin 1975)
Broad Temperature Range	None	Cold temperate-Tropical
Broad Salinity Range	None	Mesohaline-Euhaline

General Impacts

Economic Impacts

We have not found specific reports of economic impacts for Amphibalanus eburneus. However, A. eburneus is an abundant fouling species in its native range on the East and Gulf Coasts of the U.S. (Visscher 1927; Moore and Frue 1959; Utinomi 1970; Relini and Matricardi 1999; Kocak and Kucuksezgin 2001) and it can be a major contributor to growth/biofouling on the surfaces of ships and harbor structures. It is also a common fouling organism in oyster beds, and a potential competitor with oysters for space and food (White and Wilson 1996). Hull fouling by barnacles and other organisms has costly impacts for shipping lines by increasing fuel costs, decreasing maneuverability, and clogging internal seawater piping (Visscher 1927; Haderlie 1984). Barnacles also greatly contribute to fouling of navigational buoys and coastal power station intakes (Haderlie 1984).

Ecological Impacts

In its native waters on the Atlantic and Gulf coasts, A. eburneus is considered a potential fouling organism and competitor of the Eastern Oyster (Crassostrea virginica) (White and Wilson 1996). In the southern Caspian Sea, a mass population boom of A. eburneus is reported to have adversely affected molluscs and hydroids, due to competition for space and planktonic food (Zaitsev and Ozturk 2001).

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Regional Impacts

CASP	Caspian Sea		Ecological Impact	Competition
'In the Southern Caspian Sea, mass development of Amphibalanus eburneus led to a considerable decrease in the biomass of native species of molluscs and hydroids. Under the conditions of food shortage, it prevents the development of other organisms in the fouling. This competition is particularly severe in the larval stage. The larvae of Amphibalanus sometimes reached up to 90% of all plankton in some areas of the Caspian Sea.' (Zaitsev and Ozturk 2001).

References

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

Amor, Kounofi-Ben; Rifi, M.; Ghanem, R.; Draief, I.; Zouali, J.; Souissi, J. Ben (2016) Update of alien fauna and new records of Tunisian marine fauna, Mediterranean Marine Science 17(1): 124-143

Ates, A. Suat; Katagan, Tuncer; Sezgin, Murat; Özcan, Tahir (2013) Exotic crustaceans of the Turkish coast, Arthropods 2(1): 20-25

Bacon, P. R. (1971) The maintenance of a resident population of Balanus eburneus in relation to salinity fluctuations in a Trinidad mangrove swamp, Journal of Experimental Marine Biology and Ecology 6: 187-198

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

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, H.; Barnes, Margaret (1972) Some Cirripedes of the French Atlantic Coast., Journal of Experimental Marine Biology and Ecology 8: 187-194

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

Bishop, Marcus W. (1951) Distribution of barnacles by ships., Nature 167(4248): 531

Bonvechio, Kimberly I.; Barthel, Brandon; Carroll, Jessica (2018) Health and Genetic Structure of the American Eel in Florida, Southeastern Naturalist 17(3): 483-455
https://doi.org/10.1656/058.017.0311

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

California Academy of Sciences 2005-2015 Invertebrate Zoology Collection Database. <missing URL>



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

Celis, Antonio; Rodríguez-Almaráz, Gabino; Álvarez, Fernando (2007) [The shallow-water thoracican barnacles (Crustacea) of Tamaulipas, Mexico], Revista Mexicana de Biodiversidad 78: 325-337

Chainho, Paula and 20 additional authors (2015) Non-indigenous species in Portuguese coastal areas, lagoons, estuaries, and islands, Estuarine, Coastal and Shelf Science <missing volume>: <missing location>

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

Cohen, Andrew N. and 12 authors (2002) Project report for the Southern California exotics expedition 2000: a rapid assessment survey of exotic species in sheltered coastal waters., In: (Eds.) . , Sacramento CA. Pp. 1-23

Coles, S. L.; DeFelice, R. C.; Eldredge, L. G.; Carlton, J. T. (1999b) Historical and recent introductions of non-indigenous marine species into Pearl Harbor, Oahu, Hawaiian Islands., Marine Biology 135(1): 147-158

Coles, S. L.; Reath, P. R.; Longenecker, K.; Bolick, Holly; Eldredge, L. G. (2004) <missing title>, Hawai‘i Community Foundation and the U. S. Fish and Wildlife Service, Honolulu. Pp. 1-187

Cornelio, Michele; Manzoni, Alberto (1999) Caratterizzazione stagionale degli insediamenti di organismi macrobentonici su substrati sperimentali nel bacino centrale della laguna di Venezia., Bollettino del Museo Civico di Storia Naturale di Venezia 49: 135-144

Costlow, John D., Jr.; Bookhout, C. G. (1957) Larval development of Balanus eburneus in the laboratory, Biological Bulletin 112(3): 313-324

de Montaudouin, Xavier; Sauriau, Pierre-Guy (2000) Contributions to a synopsis of marine species richness in the Pertuis-Charentais Sea with new insights into the soft-bottom macrofauna of the Marennes-Oleron Bay, Cahiers de Biologie Marine 41: 181-222

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 F.; Hines, Anson H. (1994) Larval settlement of the polyhaline barnacle Balanus eburneus: cue interactions and comparisons with two estuarine congeners., Journal of Experimental Marine Biology and Ecology 179: 223-234

El-Komy, M. M.; Kajihara, T. (1991) Breeding and moulting of barnacles under rearing conditions, Marine Biology 108: 83-89

Eldredge, L. G.; Smith, C. M. (2001) Introduced marine species of Hawaii, Bishop Museum Technical Report 21: 1-60

Emara, Ahmed; Belal, Aisha (2004) Marine fouling in Suez Canal, Egypt, Egyptian Journal of Aquatic Research 30A: 189-206

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 (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

Farrapeira, Cristiane Maria Rocha; Ferreira, Gledson Fabiano de Araujo; Tenório, Deusinete de Oliveira (2010) Intra regional transportation of a tugboat fouling community between the ports of Recife and Natal, northeast Brazil, Brazilian Journal of Oceanography 58(IV SBO): 1-14

Foss, Stephen (2011) <missing title>, California Department of Fish and Game, Office of Spill Prevention and Response, Sacramento. Pp. 54

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

Gosner, Kenneth L. (1978) A field guide to the Atlantic seashore., In: (Eds.) . , Boston. Pp. <missing location>

Gould, Augustus A. (1841) <missing title>, Folsom, Wells, and Thurston, Cambridge. Pp. <missing location>

Gould, Augustus A. (1870) <missing title>, Wright and Potter, State Printers, Boston. Pp. <missing location>

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

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

Hardwick-Witman, Morgan; Mathieson, Arthur C. (1983) Intertidal macroalgae and macroinvertebrates: seasonal and spatial abundance patterns along an estuarine gradient., Estuarine Coastal and Shelf Science 16: 113-129

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

Hidu, Herbert (1978) Setting of estuarine invertebrates in Delaware Bay, New Jersey, related to intertidal-subtidal gradients, Internationale Revue der Gesamten Hydrobiologie 53(5): 637-661

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

Honma, Y.; Nakajima, J. (1973) Studies on gonad maturity in some marine invertebrates - 8.Structure of reproductive and related organs of a foreign barnacle, Balanus eburneus, that appears in Lake Kamo-ko, Sado Island., Bulletin of the Japanese Society of Scientific Fisheries 39(4): 357-365

Innocenti, G. (2006) Collections of the Natural History Museum, Zoological Section «la Specola» of the University of Florence. XXIII. Crustacea, class Maxillopoda, subclass Thecostraca, infraclass Cirripedia, Atti della Societa Toscana di Scienze Naturali Memorie, series B. 113: 1-11

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

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

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

Kim, Jeongho; Ubagan, Michael; Kwon, Soyeon; Kim, l-Hoi; Shin, Sook (2022) Variation in genetics, morphology, and recruitment of the invasive barnacle Amphibalanus eburneus (Gould, 1841) in the southern Korean peninsula, PeerJ 10: Published online
DOI 10.7717/peerj.14002

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

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>

Laguna, Jorge E. (1990) Shore barnacles (Cirripedia, Thoracica) and a revision of their provincialism and transition zones in the tropical Eastern Pacific, Bulletin of Marine Science 46(2): 406-424

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

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

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>

Matsui, Tetsui; Shane, Gillian; Newman, William (1964) On Balanus eburneus (Cirripedia, Thoracica) in Hawaii., Crustaceana 7: 141-145

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

Mook, David (1983) Indian River fouling organisms, a review, Florida Scientist 26(3/4): 162-167

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

Nelson, W. G.; Savercool, T. E.; Rodda, Neth and J. R. (1994) A comparison of the fouling community development on stabilized oil-ash and concrete reefs., Bulletin of Marine Science 55(2-3): 1303-1315

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

Norris, D. R. (1991) Recruitment, growth, and survivorship of Balanus eburneus with respect to settlement, surface, and season, Journal of the Elisha Mitchell Scientific Society 107(3): 123-130

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

Paulay, Gustav; Kirkendale, Lisa; Lambert, Gretchen; Meyer, Chris (2002) Anthropogenic biotic interchange in a coral reef ecosystem: a case study from Guam, Pacific Science 56(2): 403-422

Paulay, Gustav; Ross, Arnold (2003) An annotated checklist of the shallow water Cirripedia of Guam, Micronesica 35-36: 303-314

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

Pestana, L. B.; Dias, G. M.; Marques, A. C . (2021) Spatial and temporal diversity of non-native biofouling species associated with marinas in two Angolan bays, African Journal of Marine Science 42(4): 413-422

Pilsbry, Henry A. (1889) New and little-known American molluscs. No. 1., Proceedings of the Academy of Natural Sciences of Philadelphia 41: 81-89

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

Prezant, Robert S.; Toll, Ronald B.; Rollins, Harold B.; Chapman, Eric J. (2002) Marine macroinvertebrate diversity of St. Catherines Island, Georgia., American Museum Novitates 3367: 1-31

Rainbow, P. S. (1990) The marine flora and fauna of Hong Kong and southern China, 2 Hong Kong University Press, Hong Kong, China.. Pp. 567-568.

Ramadan, Sh. E.; Kheirallah, A. M.; Abdel-salam, Kh. M. (2006) Marine fouling community in the Eastern harbour of Alexandria, Egypt compared with four decades of previous studies, Mediterranean Marine Science 7/2: 19-29

Reise, K.; Gollasch, S.; Wolff, W.J. (1999) Introduced marine species of the North Sea coasts., Helgoländer Meeresuntersuchungen 52: 219-234

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

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

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.

Scheltema, Rudolf S.; Williams, Isabelle P. (1982) Significance of temperature to larval survival and length of development in Balanus eburneus (Crustacea: Cirripedia), Marine Ecology Progress Series 9: 43-49

Simmons, Ernest G. (1957) An ecological survey of the Upper Laguna Madre of Texas, Publications in Marine Science of the University of Texas 4(2): 156-200

Southward, A. J. (1975) Intertidal and shallow water Cirripedia of the Caribbean., Studies of the Fauna of Curacao and other Caribbean Islands 150: 1-153

Southward, A. J. (1998) New observations on barnacles (Crustacea: Cirripedia) of the Azores regions., Arquipelago. Life and Marine Sciences 16A: 11-27

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

Torres, Paulo; Costa, Ana Cristina; Dionisio, Maria Ana (2012) New alien barnacles in the Azores and some remarks on the invasive potential of Balanidae, Helgoland Marine Research 66: 513-522

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

Van Syoc (2009) Marine Biodiversity of Costa Rica, In Attached CD Springer, Berlin. Pp. 219-223

Van Syoc, Robert J. (1992) Living and fossil populations of a western Atlantic barnacle, Balanus subalbidus Henry, 1974, in the Gulf of California region, Proceedings of the San Diego Society of Natural History 12: 1-7

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

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

Zabin, Chela J. (2009) Battle of the barnacle newcomers: niche compression in invading species in Kaneohe Bay, Oahu, Hawaii, Marine Ecology Progress Series 381: 175-182

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.

Zullo, Victor A.; Beach, Dea B.; Carlton, James T. (1972) New barnacle records (Cirripedia, Thoracica), Proceedings of the California Academy of Sciences 39(6): 65-74

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

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