Invasion History

First Non-native North American Tidal Record: 1969
First Non-native West Coast Tidal Record:
First Non-native East/Gulf Coast Tidal Record: 1969

General Invasion History:

Amphibalanus reticulatus was described from southern Japan, and is native to the Indo-Pacific region. The precise boundaries of its native region are uncertain, because of its similarity to A. amphitrite, A. venustus, and A. variegatus. It has been introduced by shipping to tropical-subtropical waters of the Eastern Pacific (Laguna 1985; Coles et al. 1999; Carlton et al., 2011), both sides of the Atlantic, and the eastern Mediterranean. 

North American Invasion History:

Invasion History on the East Coast:

In the western Atlantic, Amphibalanus reticulatus was first collected in Guyanilla, Puerto Rico in 1956 (Henry and McLaughlin 1975). It was first found to be established in Biscayne Bay, Florida, in 1969 (Moore et al. 1974). It now occurs from the St. Johns River estuary, Florida (Ruiz et al. unpublished data) through the Indian River Lagoon (McPherson et al. 1984) to Biscayne Bay and the Caribbean (Southward 1975; Bacon 1976). 

Invasion History on the Gulf Coast:

In the Gulf of Mexico, Amphibalanus reticulatus was first reported from pilings and oil platforms in Louisiana in 1972 (Thomas 1975, cited by Carlton et al. 2011), and in Apalachee Bay, Florida in 1976 (Spivey 1979). This barnacle may have been present in the Gulf as early as the 1950s, but was misidentified as A. amphitrite (Carlton et al. 2011). By 1986, it was abundant from Timbalier Bay, Louisiana to Sabine Bay, Texas (Gittings et al. 1986). It has been collected in Texas from Galveston Bay, Aransas Bay, and Corpus Christi Bay (Gittings 1985; Gittings et al. 1986; Ruiz et al., unpublished data), but it is more abundant in the more turbid waters off Louisiana (Gittings et al. 1986).

Invasion History in Hawaii:

Amphibalanus reticulatus was first collected in the Hawaiian Islands at Pearl Harbor in 1929 (Henry and McLaughlin 1975; Carlton and Eldredge 2009). It is now widespread around Oahu, and is also known from Hilo Harbor, Maui, and Kauai (Coles et al. 2004; Carlton and Eldredge 2009). This barnacle has been partially replaced by Chthamalus proteus in the intertidal zone, and is now confined to a narrow band at and just above low water (Zabin 2009).

Invasion History Elsewhere in the World:

Amphibalanus reticulatus is now established from the coast of Panama, north to Mazatlan, Mexico (Laguna 1985) and La Paz, on the Gulf of California (Carlton et al. 2011). It was collected at the Pacific entrance to the Panama Canal, suggesting that the likely invasion route is shipping (Laguna 1985). This barnacle has a broad Indo-Pacific distribution, and has been treated variously as native (Jones 1992) or introduced (Jones 2004) in Australian waters. It is introduced in Pago Pago harbor, American Samoa (Coles et al. 2003).

This barnacle entered the Mediterranean, presumably through the Suez Canal by 1956, and is established in Israel (Henry and McLaughlin 1975; Galil 2007). It has been found on ships in Mediterranean France (in 1967, Zibrowius 1991) and in Belgium (in 1999, Kerckhof and Cattrijsse 2001). In the tropical East Atlantic, it is known from Fajara, the Gambia, Freetown, Sierra Leone and Lagos, Nigeria (Stubbings 1967, cited by Henry and McLaughlin 1975; Kerckhof et al. 2010). In the western Atlantic, after A. reticulatus was first collected in Puerto Rico in 1956, it was found in Trinidad in 1965 (Henry and McLaughlin 1975), and then near the eastern entrance of the Panama Canal (Southward 1975; Spivey 1976). It was first collected in Brazil in 1990 at Recife and was found in 1996 in Rio de Janeiro state (Ferreira et al. 2009). It now occurs in six Brazilian states, Maranhao, Rio Grande do Norte, Pernambuco, Alagoa, Bahia, and Rio de Janeiro (Farrapeira 2010).


Amphibalanus reticulatus usually has a conical or subcylindrical shell, with a toothed orifice. The width of the orifice is usually more than ½ of its height. The plates have a smooth surface, with wide longitudinal spaces (radii), crossed by transverse stripes, giving a net-like appearance, with the ribs narrowing to the tops of the shell plates. The adductor ridge, on the interior face of the scutum is short and high. The tergum has a more pointed apex than A. amphitrite or A. improvisus. The tergal spur is narrow, and the spur length is about 3/10 the length of the basal margin. The spur width is about 1/4 of the basal margin (Henry and McLaughlin 1975). The shell is usually buff or white in color, with dark-purple longitudinal stripes, crossed by many alternating red-and-white transverse lines. Type specimens averaged 18 mm diameter (Henry and McLaughlin 1975). The larval stages of A. reticulatus are described by Thiyagarjan et al. (1997).

Amphibalanus reticulatus is a member of the Amphibalanus amphitrite species complex and can easily be confused with A. amphitrite, A. eburneus, A. ienustus, A. variegatus, A. subalbidus, and other closely related species (Henry and McLaughlin 1975).


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:   reticulatus


Balanus amphitrite var. cirratus (Zevina and Tarasov, 1964)
Balanus amphitrite var. tesselatus (Utinomi, 1964)

Potentially Misidentified Species

Amphibalanus variegatus

Amphibalanus venustus

Ampibalanus amphitrite



Amphibalanus reticulatus prefers saline (30-40 PSU), subtidal habitats in subtropical and tropical seas, although it has been found at sailintiies as low as 10 PSU. It is typically found in sheltered and exposed waters on a wide range of hard surfaces, including docks, pilings, floats, mangroves, rocks, ships' hulls, oysters, and other shellfish (Henry and McLaughlin 1975; Jones 1992; Farrapeira 2008) This species, 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. The nauplii feed in the plankton and go through five successive molts, spending at least 4-5 days in the water column before molting into a non-feeding cypris stage, covered with a pair of chitinous shells. 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, and gathering phytoplankton, zooplankton, and detritus (Barns 1983).


Phytoplankton; zooplankton


Crabs, snails, fishes

Trophic Status:

Suspension Feeder



General HabitatCoarse Woody DebrisNone
General HabitatOyster ReefNone
General HabitatMarinas & DocksNone
General HabitatRockyNone
General HabitatVessel HullNone
General HabitatMangrovesNone
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Tidal RangeSubtidalNone
Tidal RangeLow IntertidalNone

Tolerances and Life History Parameters

Minimum Salinity (‰)10Field salinity, Paripe River estuary, Brazil (Farrapeira 2008)
Maximum Salinity (‰)40Larval (cypris) settlement, 28 C (Thiyagarjan et al. 2002)
Minimum Reproductive Salinity20Larval (cypris) settlement, 28 C (Thiyagarjan et al. 2002)
Maximum Reproductive Salinity40Larval (cypris) settlement, 28 C (Thiyagarjan et al. 2002)
Minimum Duration526 C, from larval release to cypris (settling stage) (Thiyagarajan et al. 1997)
Maximum Width (mm)26Maximum adult width (Henry and McLaughlin 1975)
Maximum Height (mm)15Maximum adult height (Henry and McLaughlin 1975)
Broad Temperature RangeNoneWarm temperate-Tropical
Broad Salinity RangeNonePolyhaline-Euhaline

General Impacts

Economic Impacts

Amphibalanus reticulatus is a frequent fouler of ships and marine structures worldwide in warm subtropical-tropical waters (Utinomi 1970; Henry and McLaughlin 1975).

Ecological Impacts

Competition- Amphibalanus reticulatus may have replaced or excluded A. amphitrite on the coast of Louisiana (Gittings et al. 1986) and in Hawaiian waters, before it was restricted by the invasion of Chthamalus proteus (Zabin 2009). However, to our knowledge, competitive relationships of A. amphitrite and A. reticulatus have not been studied.

Regional Impacts

CAR-INorthern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern FloridaEcological ImpactCompetition
Gittings et al. (1986) suggest that A. reticulatus excludes A. amphitrite in the northwestern Gulf (Timbalier Bay, LA to the TX border/Sabine Bay), where the water is more turbid than along the southwest TX coast, where the water is clearer.
SP-XXINoneEcological ImpactCompetition
Amphibalanus amphitrite was the dominant barnacle in sheltered coastal areas of Oahu, Hawaii (Matsuda 1973, cited by Zabin 2009). It may have been replaced by A. reticulatus, before the distribution of the latter species was severely restricted by the invasion of Chthamalus proteus, which began in 1993. However, there were no studies of barnacle populations in this area between 1973 and 2001, so the nature of the interactions between the species is unknown (Zabin 2009).
HIHawaiiEcological ImpactCompetition
Amphibalanus amphitrite was the dominant barnacle in sheltered coastal areas of Oahu, Hawaii (Matsuda 1973, cited by Zabin 2009). It may have been replaced by A. reticulatus, before the distribution of the latter species was severely restricted by the invasion of Chthamalus proteus, which began in 1993. However, there were no studies of barnacle populations in this area between 1973 and 2001, so the nature of the interactions between the species is unknown (Zabin 2009).

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
NWP-3b None 0 Native Estab
NWP-3a None 0 Native Estab
NWP-2 None 0 Native Estab
EAS-I None 0 Native Estab
EAS-VI None 0 Native Estab
EAS-VII None 0 Native Estab
EAS-VIII None 0 Native Estab
CIO-I None 0 Native Estab
CIO-II None 0 Native Estab
EA-IV None 0 Crypto Estab
WA-V None 1964 Crypto Estab
CAR-I Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida 1969 Def Estab
CAR-IV None 1956 Def Estab
CAR-III None 1965 Def Estab
MED-V None 1958 Def Estab
SP-XXI None 1929 Def Estab
NEP-VIII None 1984 Def Estab
SEP-H None 1985 Def Estab
AUS-XII None 1979 Crypto Estab
AUS-IV None 1990 Crypto Estab
CAR-II None 1970 Def Estab
SP-IX None 2002 Def Estab
SA-III None 1990 Def Estab
WA-II None 1961 Def Estab
AUS-XIV None 0 Crypto Estab
S190 Indian River 1981 Def Estab
G070 Tampa Bay 2001 Def Estab
S200 Biscayne Bay 1969 Def Estab
G090 Apalachee Bay 1976 Def Estab
G240 Calcasieu Lake 1983 Def Estab
G130 Pensacola Bay 2000 Def Estab
G260 Galveston Bay 2000 Def Estab
S180 St. Johns River 2001 Def Estab
NWP-4a None 0 Native Estab
IP-1 None 0 Native Estab
AUS-II None 2000 Crypto Estab
AUS-I None 2000 Crypto Estab
G310 Corpus Christi Bay 1985 Def Estab
CAR-VII Cape Hatteras to Mid-East Florida 2002 Def Estab
SA-II None 1990 Def Estab
G210 Terrebonne/Timbalier Bays 1972 Def Estab
G220 Atchafalaya/Vermilion Bays 1986 Def Estab
G230 Mermentau River 1986 Def Estab
G250 Sabine Lake 1986 Def Estab
SA-IV None 2009 Def Estab
NEP-VII None 2006 Def Estab
WA-I None 2010 Def Estab
WA-I None 2010 Def Estab
PAN_PAC Panama Pacific Coast 1985 Def Estab
PAN_CAR Panama Caribbean Coast 1965 Def Estab
EA-III None 2018 Native Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude


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

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Carlton, James T.; Eldredge, Lucius (2009) Marine bioinvasions of Hawaii: The introduced and cryptogenic marine and estuarine animals and plants of the Hawaiian archipelago., Bishop Museum Bulletin in Cultural and Environmental Studies 4: 1-202

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

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

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Coles, S. L.; Reath, P. R.; Skelton, P. A.; Bonito, V; DeFelice; Basch, L. (2003) Introduced marine species in Pago Pago Harbor, Fagatele Bay and the National Park Coast, American Samoa., Bishop Museum Technical Report 26: 1-24

da Silva, Eder Carvalho; Barros, Francisco (2011) [Benthic macrofauna introduced in Brazil: List of marine and freshwater species and actual distribution], Oecologia Australis 15(2): 326-344

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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

Farrapeira, Cristiane Maria Rocha; de Melo,Arthur Vinícius de Oliveira Marrocos; Barbosa, Débora Ferreira; da Silva, Karla Maria Euzebio (2007) Ship hull fouling in the port of Recife, Pernambuco, Brazilian Journal of Oceanography 55(3): 207-221

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

Ferreira,Carlos Eduardo Leite; Junqueira,Andrea de Oliveira Ribeiro; Villac, Maria Célia; Rubens Mendes Lopes, Rubens Mendes (2009) Biological Invasions in Marine Ecosystems., Springer-Verlag, Berlin, Heidelberg. Pp. 459-477

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

Galil, Bella S. (2007) Seeing Red: Alien species along the Mediterranean coast of Israel., Aquatic Invasions 2(4): 281-312

Gittings, Stephen R. (1985) Notes on barnacles (Cirripedia: Thoracica) from the Gulf of Mexico., Gulf Research Reports 8(1): 35-41

Gittings, Stephen R.; Dennis, George D.; Harry, Harold W. (1986) <missing title>, Texas A&M Sea Grant, College Station TX. Pp. 1-36

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

Huang, Zongguo (Ed.), Junda Lin (Translator) (2001) Marine Species and Their Distributions in China's Seas, Krieger, Malabar, FL. Pp. <missing location>

Ignacio, Barbara L.; Julio, Luciana M.; Junqueira, Andrea O. R; Ferreira-Silva, Maria A. G. (2010) Bioinvasion in a Brazilian Bay: filling gaps in the knowledge of southwestern Atlantic biota, PLOS ONE 5(9): <missing location>

Jones, Diana S. (1992) A review of Australian fouling barnacles, Asian Marine Biology 9: 89-100

Jones, Diana S. (2004) Barnacles (Cirripedia: Thoracica) of the Dampier Archipelago, Western Australia., Records of the Western Australia Museum Suppl. 66: 121-157

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

Kerckhof, F.; Haelters, J.; Degraer, S. (2010) The barnacles Chirona (Striatobalanus) amaryllis (Darwin 1854) and Megabalanus coccopoma (Darwin 1854) (Crustacea, Cirripedia): two invasive species new to tropical West African waters, African Journal of Marine Science 32(2): 265 -268

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

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

Lira, Simone Maria de Albuquerque ; Farrapeira, Cristiane Maria Rocha (2010) Sessile and sedentary macrofauna from the Pirapama Shipwreck, Pernambuco, Brazil, Biota Neotropica 10(4): published online

Lomonaco, Cecilia; Santos, Andre S.; Christoffersen, Martin l. (2011) Effects of local hydrodynamic regime on the individual’s size in intertidal Sabellaria (Annelida: Polychaeta: Sabellariidae) and associated fauna at Cabo Branco beach, north-east Brazil, Marine Biodiversity Records 4(e76): Published online

Lopes, Rubens M. (Ed.) (2009) <missing title>, Ministry of the Environment, Brasilia, Brazil. Pp. 1-440

Marques, Antonio C. and 17 authors (2013) Rapid assessment survey for exotic benthic species in the São Sebastião Channel, Brazil, Latin American Journal of Aquatic Research 41(2): 265-285

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

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

Neves, Carolina Somaio; da Rocha, Rosana Moreira (2008) Introduced and cryptogenic species and their management in Paranaguá Bay, Brazil, Brazilian Archives of Biology and Technology 51(3): 623-633

Neves, Carolina Somao; Rocha, Rosana Moreira; Pitombo, Fabio Bettini; Roper, James J. (2007) Use of artificial substrata by introduced and cryptogenic marine species in Paranagua Bay, southern Brazil, Biofouling 23(5): 319-330

Rajagopal, S.;Nair, K.V.K.; Van Der Velde, G.; Jenner, H.A. (1997) Seasonal settlement and succession of fouling communities in Kalpakkam, east coast of India., Netherlands Journal of Aquatic Ecology 30(4): 309-325

Rocha, Rosana M.; Cangussu, Leonardo C.; Braga, Mariana P. (2010) Stationary substrates facilitate bioinvasion in Paranaguá bay in southern Brazil, Brazilian Journal of Oceanography 58(Special Issue 4): 23-28

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

Soors, Jan; Faasse, Marco; Stevens, Maarten; Verbessem, Ingrid; De Regge, Nico;Van den Bergh, Ericia (2010) New crustacean invaders in the Schelde estuary (Belgium), Belgian Journal of Zoology 140: 3-10

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

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

Spivey, Henry R. (1979) First records of Trypetesa and Megalasma (Crustacea: Cirripedia) in the Gulf of Mexico., Bulletin of Marine Science 29(4): 497-508

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

Teixeira, Rogério L.; Sá, Hélio S. (1998) [Abundance of decapod macrocrustaceans occupying the shallow waters of a tropical estuary], Revista Brasileira da Biologia 58(3): 393-404

Thiyagarajan, Vengatesen; Nair, K. V. K.; Subramonian, T.; Venugopalan, V. P. (2002) Larval settlement behaviour of the barnacle Balanus reticulatus in the laboratory., Journal of the Marine Biological Association of the United Kingdom 82: 579-582

Thiyagarjan, V.; Venugopalan, V. P.; Subranomonam, T. (1997) Larval description of Balanus reticulatus reared in the laboratory., Journal of Experimental Marine Biology and Ecology 209: 215-231

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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

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

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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

Zabin, Chela J. (2015) Patterns of adult abundance vary with recruitment of an invasive barnacle species on Oahu, Hawaii, Journal of Experimental Marine Biology and Ecology 464: 44-51

Zibrowius, Helmut (1991) Ongoing modification of the Mediterranean marine fauna and flora by the establishment of exotic species., Mesogee 51: 83-107

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