Invasion History

First Non-native North American Tidal Record: 1986
First Non-native West Coast Tidal Record:
First Non-native East/Gulf Coast Tidal Record: 1986

General Invasion History:

Charybdis hellerii is a swimming crab with a very broad Indo-Pacific native distribution. It ranges from the Red Sea and East Africa, throughout the Indian Ocean to northern Australia and New Caledonia, and north to China and Okinawa, Japan (Stephenson et al. 1957; Spiridonov 1990; LeMaitre 1995; Galil et al. 2002). In 1925, it was collected in Haifa, Israel, in the Mediterranean Sea, which it reached through the Suez Canal (Galil et al, 2002). It is now widespread in the eastern Mediterranean Sea, and in 2003-2004, was collected in Greece and Turkey on the Aegean Sea (Yokes et al. 2007; Zenetos et al. 2008). In 1986, a specimen of C. hellerii was captured near Charleston, South Carolina (USGS Center for Aquatic Resources Studies 2008). In 1987, it was found on the northwest and southwest coast of Cuba, and on the coasts of Venezuela and Columbia. In 1995, C. hellerii was found in the Indian River Lagoon, Florida and found to be established but uncommon in that estuary (Lemaitre 1995; Dineen et al. 2001). In 1995-1996, it was collected from southern Brazil (Mantellatto et al. 1999). It now has a fairly continuous range from south of Cape Hatteras, North Carolina to Florianopolis, Brazil (Mantelatto et al. 1999; Ferreira et al. 2001; USGS Center for Aquatic Resources Studies 2008).

North American Invasion History:

Invasion History on the East Coast:

In 1986, a specimen of C. hellerii was captured in Folly Creek, near Charleston, South Carolina (USGS Nonindigenous Aquatic Species Program 2008), but was not immediately recognized as an introduced species. The first published record of C. hellerii on the US East Coast, was from Fort Pierce, Florida, in the Indian River Lagoon, in 1995. Several specimens were found, including an ovigerous female (Lemaitre 1995). Subsequently, 134 crabs were collected in St. Lucie and Sebastian inlets in the Lagoon, and Lake Worth Inlet, to the south (Dineen et al. 1997). From 2000 to 2009, C. hellerii was collected at least 17 times at 15 locations north of the Indian River Lagoon, as far north as off Portsmouth Island, ~50 km southwest of Cape Hatteras, North Carolina (in 2004, USGS Nonindigenous Aquatic Species Program 2008).

Invasion History on the Gulf Coast:

We are aware of only two records of Charybdis hellerii from the Gulf of Mexico, one from the mouth of Tampa Bay (in 2004, USGS Nonindigenous Species Program 2008; McMillen-Jackson 2008), and one from Long Key, Florida Bay (in 2009, USGS Nonindigenous Species Program 2009).

Invasion History in Hawaii:

One specimen was found within the fouling community on the hull of a ship in Pearl Harbor, Oahu in 1950, (Edmondson 1954). No further records of C. hellerii are known from Hawaii (Carlton and Eldredge 2009).

Invasion History Elsewhere in the World:

As noted above, this crab is a Lessepsian species, which entered the Mediterranean Sea through the Suez Canal, first collected in Israel in 1925 (Galil et al. 2002). It is now widespread in the eastern Mediterranean Sea, having expanded its range north and west, reaching Cyprus in 1999 (Galil et al. 2002), and Greece and Turkey on the Aegean Sea by 2003-2004 (Yokes et al. 2007; Zenetos et al. 2008).

However, the expansion of C. hellerii was much more dramatic after its introduction to the Western Atlantic, probably by ballast water. The earliest (1986-1987) records are from Charleston, South Carolina (USGS Nonindigenous Aquatic Species Program 2008), Cuba, and Venezuela (Lemaitre 1995). It was collected in Salvador, Bahia in central Brazil in 1995 (Mantellatto et al. 1999), in the Sao Paulo region in 1996 (Mantelatto et al. 1999), and at Florianopolis (-27.8 S) in 1998 (Mantelatto et al. 1999). Records from the equatorial regions of South America are a few years later (French Guiana, in 2001, Tavares et al. 2003; Macau, Rio Grande del Norte, Brazil, in 2000, Ferreira et al. 2001; Panaquatira, Maranhao, in 2005, Feres et al. 2007), but this could reflect a lower intensity of collecting. Overall, this species has been collected over a range of 62 degrees of latitude in the 20 years since its first discovery in the Western Atlantic.

Further invasions are possible. This crab was collected in the sea-chest of a ship sampled in New Zealand (Coutts and Dodgshun 2007). In Guam, C. hellerii is considered cryptogenic (Paulay et al. 2003).


Description

The Indo-Pacific swimming crab, Charybdis hellerii, has a hexagonal carpace with a width roughly 1.4 X the length. The font of the carapace bears six lateral teeth between the eyes, with six large anterolateral teeth on each side. The claws are stout, with the palms bearing five black-tipped spines on the dorsal surface. The anterior margin of the merus ('forearm') bears three prominent spines, while the carpus ('wrist') has a strong spine on the interior margin. The merus and carpus of the 5th legs each carry a strong spine on the posterior margin, and the distal segments of the legs are widened and flattened for swimming (Lemaitre 1995; Galil et al. 2002). The abdomen of both sexes has a rounded triangular shape, but is ~20% of the carapace width in adult males, and about 40% in females (Mantellato et al. 2001). The overall color is dark green, but the inner surface of the palms of the claws, and the inner surfaces of the walking legs are dark purple. The tips of the claws are white. There are whitish areas on the frontal area and the dorsal sides of the carapace (Lemaitre 1995; Galil et al. 2002). Males mature at about 35-40 mm carapace width and occasionally reach 65 mm. Females mature at about the same size, but tend to be smaller (Mantellato et al. 2001). Dineen et al. (2001) provide a description of larval development.


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Arthropoda
Subphylum:   Crustacea
Class:   Malacostraca
Subclass:   Eumalacostraca
Superorder:   Eucarida
Order:   Decapoda
Suborder:   Pleocyemata
Infraorder:   Brachyura
Superfamily:   Portunoidea
Family:   Portunidae
Genus:   Charybdis
Species:   hellerii

Synonyms

Charybdis merguiensis (de Man, 1887)

Potentially Misidentified Species

Charybdis variegata
Indo-Pacific crab, 1 specimen in Brazil (Sant'Anna et al. 2012)

Cronius ruber
A large reddish native (New Jersey-Brazil) swimming crab (Williams 1984)

Ecology

General:

Life History- In crabs of the family Portunidae, the male attends the female before molting, and carries the female around, underneath his carapace. He releases the female, allows her to molt, and then copulates with her, inserting the first pair of pleopods, carrying sperm, into the female's seminal receptacles. The eggs are fertilized internally, and then extruded as a 'sponge' or a mass of eggs, which are brooded between the abdomen and the body (Barnes 1983; Williams 1984; Dineen et al. 2001). The eggs hatch into zoeae, larvae about 1 mm long, armed with long spines, which drift in the plankton. Each zoea goes through six molts, and eventually molts into a postlarval megalopa, with prominent eyes and partially developed appendages. The megalopa is capable of crawling on the benthos and active, directed swimming. After ~44 days in the plankton, the larva settles and molts into a miniature 'first crab' which has all the features of an adult crab (Barnes 1983; Dineen et al. 2001).

Ecology- Adult Charybdis helleri use a broad range of intertidal and subtidal habitats, including natural rock, riprap, coral reefs, coral rubble, mangrove roots, seagrass beds, and patches of subtidal algae (Stephenson et al. 1957; Dineen et al. 2001; Feres et al. 2007; Felder et al. 2010). In the Bahia de Santos, Brazil, C. hellerii was most common in the shallow rocky subtidal, and the interface between rocks and the soft bottom (Sant'Anna et al. 2012).

Food:

mollusks, crustaceans, annelids, other inverts

Trophic Status:

Carnivore

Carn

Habitats

General HabitatUnstructured BottomNone
General HabitatMarinas & DocksNone
General HabitatRockyNone
General HabitatMangrovesNone
General HabitatCoral reefNone
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Tidal RangeSubtidalNone
Tidal RangeLow IntertidalNone
Vertical HabitatEpibenthicNone
Vertical HabitatNektonicNone


Tolerances and Life History Parameters

Minimum Salinity (‰)20Experimenta Data (Occhi et al. 2019); Field Data- Bahia de Santos, Brazil (Sant'Anna et al. 2012); Indian River Lagoon FL (28 PSU, Dineen et al. 2001)
Maximum Salinity (‰)40Typical Red Sea salinity
Minimum Duration44Larval release to first crab, 24 C- Dineen et al. 2002
Minimum Width (mm)37.5Mature female (Bolanos et al. 2011, Isla Margarita, Venezuela)
Maximum Width (mm)78Mature female (Indian River Lagoon, Dineen et al. 2001)
Broad Temperature RangeNoneWarm temperate-Tropical
Broad Salinity RangeNonePolyhaline-Euhaline

General Impacts

The Indo-Pacific swimming crab, Charybdis hellerii, is long-established in the Eastern Mediterranean and has recently colonized a very wide range in the Western Atlantic. Owing to its small size, it is unlikely to become a sought-after fisheries item. It is a potential predator and competitor of native crabs, and a study from Twin Cays, Belize, shows that C. hellerrii may have displaced several species of native crustaceans from shallow-water habitats (Felder et al. 2010). Impacts beyond this are not well-documented. 

Regional Impacts

CAR-IINoneEcological ImpactCompetition
Charybdis hellerii may have displaced the native crabs Mithrax spp., Callinectes spp., and the spiny lobster Panulirus which were abundant in 2002 and 1983 collections (Felder et al. 2010).
MED-VNoneEcological ImpactCompetition
Charybdis hellerii was listed as 'invasive' in the Eastern Mediterranean, implying competition with native species (Zenetos et al. 2011). However, we do not have more specific information on its impacts in this region.

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
NWP-3a None 0 Native Estab
EAS-I None 0 Native Estab
NWP-2 None 0 Native Estab
EAS-III None 0 Native Estab
AUS-I None 0 Native Estab
SP-IV None 1876 Native Estab
EAS-VI None 0 Native Estab
CIO-IV None 0 Native Estab
CIO-III None 0 Native Estab
CIO-II None 0 Native Estab
AUS-XIII None 0 Native Estab
CIO-I None 0 Native Estab
IP-1 None 0 Native Estab
AG-5 None 0 Native Estab
AG-1 None 0 Native Estab
AG-3 None 0 Native Estab
AG-4 None 0 Native Estab
RS-1 None 0 Native Estab
RS-3 None 0 Native Estab
RS-2 None 0 Native Estab
EA-III None 0 Native Estab
MED-V None 1925 Def Estab
CAR-III None 1987 Def Estab
CAR-II None 1987 Def Estab
CAR-I Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida 1987 Def Estab
SA-III None 1995 Def Estab
SA-IV None 2000 Def Estab
SA-II None 1996 Def Estab
AUS-XII None 0 Native Estab
CAR-VII Cape Hatteras to Mid-East Florida 1986 Def Estab
SP-XXI None 1950 Def Failed
CAR-VI None 2001 Def Estab
S190 Indian River 1995 Def Estab
G070 Tampa Bay 2004 Def Unk
S080 Charleston Harbor 1986 Def Estab
MED-VI None 2003 Def Estab
S030 Bogue Sound 2006 Def Estab
AUS-II None 0 Native Estab
S045 _CDA_S045 (New) 2000 Def Estab
S020 Pamlico Sound 2004 Def Estab
S196 _CDA_S196 (Cape Canaveral) 1997 Def Estab
S183 _CDA_S183 (Daytona-St. Augustine) 2002 Def Estab
S175 _CDA_S175 (Nassau) 2004 Def Estab
S170 St. Marys River/Cumberland Sound 2004 Def Estab
S125 _CDA_S125 (Ogeechee Coastal) 2003 Def Estab
S206 _CDA_S206 (Vero Beach) 2009 Def Unk
S060 Winyah Bay 2000 Def Estab
SP-XII None 0 Crypto Estab
S180 St. Johns River 2003 Def Estab
S100 St. Helena Sound 2014 Def Estab
NEA-V None 2009 Def Unk
CAR-IV None 2017 Def Estab
WA-V None 0 Native Estab
EAS-II None 0 Native Estab
SP-I None 0 Native Estab
AUS-XIII None 0 Native Estab
AUS-XII None 0 Native Estab
AUS-X None 0 Native Estab
WA-II None 2012 Def Estab
WA-I None 2011 Def Unk
S076 _CDA_S076 (South Carolina Coastal) 2019 Def Estab
S150 Altamaha River 2020 Def Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude

References

Marasinghe, M. M. K.I ; Ranatunga, R. R. M .K. P. Anil, Arga C. (2018) Settlement of non-native Watersipora subtorquata (d'Orbigny, 1852) in artificial collectors deployed in Colombo Port, Sri Lanka, BioInvasions Records 7: 7-14

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

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

Bernadochi, Ligia Coletti; Silvestri, Fausto; Turra, Alexander (2012) Composition and seasonal variation of Brachyura and Anomura (Crustacea, Decapoda) associated with brown mussel farms at Praia da Cocanha, Brazil, Brazilian Archives of Biology and Technology 55(4): 549-557

Bolanos, Juan A.; Baeza, J. Antonio; Hernandez, Jesus E.; Lira, Carlos; Lopez, Regulo (2011) Population dynamics and reproductive output of the non-indigenous crab Charybdis hellerii in the south-eastern Caribbean Sea, Journal of the Marine Biological Association of the United Kingdom 92(3): 469-474

Brockerhoff, Annette; McLay, Colin (2011) In the wrong place- Alien marine crustaceans: Distribution, biology, impacts, Springer, Dordrecht. Pp. 27-106

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

Çinar, M. E.; Noglu, M. Bilece; Özturk, B.; Katagan, T. ; Aysel, V. (2005) Alien species on the coasts of Turkey, Mediterranean Marine Science 6/2: 119-146

Cinar, Melih Ertan and 7 authors (2021) Current status (as of end of 2020) of marine alien species in Turkey, PLOS ONE 16: Published online

Çinar, Melih Ertan; Bilecenoglu, Murat; Öztürk, Bilal; Can, Alp (2006) New records of alien species on the Levantine coast of Turkey, Aquatic Invasions 1(2): 84-90

Coelho, Petrônio Alves; Santos, Maria do Carmo Ferrão (2004) [Swimmer crabs from the Una River estuary, São José da Coroa Grande, Pernambuco State, Brazil (Crustacea, Decapoda, Portunidae)] (Portuguese), Boletim Tecnico-Cientifico do CEPENE 12(1): 187-194

Coutts, Ashley D.M.; Dodgshun, Tim J. (2007) The nature and extent of organisms in vessel sea-chests: A protected mechanism for marine bioinvasions., Marine Pollution Bulletin 54: 875-886

Crafton, R. Eliot (2015) Modeling invasion risk for coastal marine species utilizing environmental and transport vector data, Hydrobiologia 746: 349-362

Cuesta, Jose A.; Almon, Bruno; Perez-Dieste, Jacinto; Trigo, Juan E.; Banon, Rafael (2015) Role of ships’ hull fouling and tropicalization process on European carcinofauna: new records in Galician waters (NW Spain), Biological Invasions Published online: <missing location>

de Almeida, Alexandre O.; Coelho, Petrônio A. (2008) Estuarine and marine brachyuran crabs (Crustacea: Decapoda) from Bahia, Brazil: checklist and zoogeographical considerations, Latin American Journal of Aquatic Research 36(2): 183-222

de Almeida, Alexandre Oliveira; Coelho, Petrônio Alves; dos Santos, José Tiago Almeida; Ferraz, Neyva Ribeiro (2006) [Estuarine Decapod Crustaceans from Ilhéus, State of Bahia], Biota Neotropica 6(2): published online

Derviche, Patrick; da Cunha Lana, Paulo (2022) The effects of the nonnative brittle star Ophiothela mirabilis Verrill, 1867 on the feeding performance of an octocoral host in a southwestern Atlantic rocky shore, Biological Invasions 24(9): 2821 - 2833

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

Dineen, Joseph; Clark, Paul F.; Hines, Anson H.; Reed, Sherry; Walton, Heather P. (2001) Life history, larval description, and natural history of Charybdis hellerii, an invasive crab in the western Atlantic, Journal of Crustacean Biology 21(3): 774-805

Eash-Loucks, Wendy E.; Kimball, Mathew E.; Petrinec, Kathryn M. (2014) Long-term changes in an estuarine mud crab community: Evaluating the impact of non-native species, Journal of Crustacean Biology 36(6): 731-738

Edmondson, C. H. (1954) Hawaiian Portunidae, Occasional Papers of Bernice P. Bishop Museum 21(12): 215-274

Ezgeta-Balic, Daria and 12 authors (2021) Competitive feeding interactions between native Ostrea edulis and non-native Crassostrea gigas with implications of introducing C. gigas into commercial aquaculture in the eastern Adriatic Sea, Marine Environmental Research 1060(105051): Published online

Felder, Darryl L. and 6 authors (2010) Obvious invaders and overlooked infauna: unexpected constituents of the decapod crustacean fauna at Twin Cays, Belize, Smithsonian Contributions to the Marine Sciences 38: 181-88

Feres, Samir Jorge Cutrim; Lopes, Ana Tereza Lyra ; Santos, Luiza de Andrade (2007) Primeiro registro de Charybdis hellerii (Milne Edwards, 1867) para o litoral maranhense-brasil. (Crustácea, Decapoda, Portunidae)., Boletim do Labratorio de Hidrobiologia 20: 77-82

Fernandes, J.M.; Rosa, D.M.; Araujo, C.C.V.; Ripoli, L.V. (2006) Primeiro registro do siri exótico Charybdis hellerii (A. Minle-Edwards, 1867) no estado do Espírito Santo, Congresso Brasileiro de Biologia Marinha <missing volume>: <missing location>

Ferreira, Alexander C.; Sankarankuty, Cheruparambil; da-Cunha, Inailson M. C.; Duarte, Fabio T. (2001) (Yet another record of Charybdis hellerii (A. Milne Edwards) (Crustacea, Decapoda) from the northeast of Brazil., Revista Brasileira da Zoologia 18(Suppl. 1): 57-358

Frick, Michael G.; Williams, Kristina L. (2006) Charybdis hellerii, a non-indigenous portunid crab from the gastrointestinal contents of a Kemp’s Ridley sea turtle (Lepidochelys kempi,) in Georgia, USA., Marine Turtle Newsletter 111: 15

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

Galil, Bella; Froglia, Carlo; Noel, Pierre (2002) CIESM atlas of exotic species in the Mediterranean: Crustaceans., In: None(Eds.) None. , Monoco. Pp. <missing location>

Grosholz, Edwin (2011) Encyclopedia of Biological Invasions, University of California Press, Berkeley and Los Angeles. Pp. 125-128

Holthuis, L.B., Gottlieb, E. (1974) An annotated list of the decapod Crustacea of the Mediterranean coast of Israel, with an appendix listing the decapoda of the eastern Mediterranean., Bulletin of the Research Council of Israel 7B: 1-126

Huang, Zongguo (Ed.) (2001) <missing title>, Krieger, Malabar, FL. Pp. <missing location>

International Commission for the Scientific Exploration of the Mediterranean Sea (CEISM) (2001) <missing title>, <missing publisher>, <missing place>. Pp. <missing location>

Kapiris, Kostas; Kataan, Tuncer; Ate, Suat A.; Conides, Alexis (2012) Review of alien decapods (Crustacea) in the Aegean Sea, Journal of the Black Sea/Mediterranean Environment 18(2): 177-187

Kathrivel. M.; Gopalakrishnan, K. N. (1974) On the occurrence of Charybdis (Charybdis) hellerii along the west coast of India., Indian Journal of Marine Science 16(1): 286-287

Katsanevakis, S.; Tsiamis, K.; Ioannou, G.; Michailidis, N.; Zenetos, A. (2009) Inventory of alien marine species of Cyprus, Mediterranean Marine Science 10(2): 109-133

Lemaitre, Rafael (1995) Charybdis hellerii (Milne Edwards, 1867), a nonindigenous portunid crab (Crustacea, Decapoda, Brachyura) discovered in the Indian River lagoon system of Florida, Proceedings of the Biological Society of Washington 108(4): 643-648

Loebmann, Daniel; Mai, Ana Cecília G.; Lee, James T. (2010) The invasion of five alien species in the Delta do Parnaíba Environmental Protection Area, Northeastern Brazil, Revista de Biologia Tropical 58(3): 909-923

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

Mantelatto, Fernando L. M., Luciano L. Diaz (1999) Extension of the known distribution of Charybdis helleri (A. Milne Edwards 1967) (Decapoda, Portunidae) along the western tropical South Atlantic., Crustaceana 72: 617-619

Mantelatto, Fernando Luis Medina; Garcia, Renata Biagi (2001) Biological aspects of the nonindigenous portunid crab Charybdis hellerii in the western tropical South Atlantic, Bulletin of Marine Science 68(3): 469-477

McMillen-Jackson, Anne L. (2008) First record of the Indo-Pacific swimming crab, Charybdis hellerii (A. Milne-Edwards, 1867) In the Gulf Of Mexico, Crustaceana 81(7): 889-894

Ozcan,Tahir; Katagan, Tuncer; Kocatas, Ahmet (2005) Brachyuran crabs from Iskenderun Bay (southeastern Turkey), Crustaceana 78(2): 237-243

Paulay, Gustan (200) Metopograpsus oceanicus (Crustacea: Brachyura) in Hawaii and Guam: another recent invasive?, Pacific Science 61(2): 295-300

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; Kropp, Roy; Ng, Peter, K. L.; Eldredge, Lucius G. (2003) The crustaceans and pycnogonids of the Mariana Islands, Micronesica 35-36: :56-513

Perry, Harriet; Yeager, David (2006) <missing title>, Gulf Coast Research Laboratory- University of Southern Mississiuppi, Ocean Springs MS. Pp. 8

Rodriguez, Gilberto; Suarez, Hector (2001) Anthropogenic dispersal of record of decapod crustaceans in aquatic environments, Interciencia 26(7): 282-288

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>

Sant’Anna, Bruno Sampaio; Watanabe, Timóteo Tadashi; Turra, Alexander; Zara, Fernando José (2011) Relative abundance and population biology of the non-indigenous crab Charybdis hellerii (Crustacea: Brachyura: Portunidae) in a southwestern Atlantic estuary-bay complex, Aquatic Invasions 7: in press

Sant’Anna, Bruno Sampaio; Watanabe, Timóteo Tadashi; Turra, Alexander; Zara, Fernando José (2012) First record of the non-indigenous portunid crab Charybdis variegata from the western Atlantic coast, Bioinvasion Records 1: in press

Simoes, Nuno; Wakida-Kusunoki, Armando T.; Cruz-Sánchez, José Luís; Alvarez, Fernando; andVillalobos-Hiriart, José Luis (2021) On the presence of Charybdis (Charybdis) hellerii (A. Milne-Edwards, 1867) on the Mexican coast of the Gulf of Mexico, BioInvasions Records 8(3): 670–674

Spiridonov, V.A. (1990) Results of the Rumphius Biohistorical Expedition to Ambon, Backhuys Publishers, Amsterdam. Pp. <missing location>

Stephenson, W.; Hudson, Joy J.; Campbell, B. (1957) The Australian portunids (Crustacea: Portunidae). II. The genus Charybdis., Australian Journal of Marine and Freshwater Research 8: 491-506

Tavares, M. R.; .Franco, A. C. S. ; . Ventura, C. R. R.; .Santos, L. N. (2021) Geographic distribution of Ophiothela brittle stars (Echinodermata: Ophiuroidea): substrate use plasticity and implications for the silent invasion of O. mirabilis in the Atlantic, Hydrobiologia 848: 2093-2103

Tavares, Marco (2011) In the wrong place- Alien marine crustaceans: Distribution, biology, impacts, Springer, Dordrecht, Netherlands. Pp. 251-268

Tavares, Marcos; Amouroux, Jean-Michel (2003) First record of the non-indigenous crab, Charybdis hellerii from French Guiana (Decapoda, Brachyura, Portunidae)., Crustaceana 76(5): 625-630

2002-2021 Invertebrate Zoology Collections Database. <missing description>

2003-2015 Nonindigenous Aquatic Species Database. Gainesville, FL. http://nas.er.usgs.gov

Wee, Desmond P. C.; Ng, Peter K. L. (1995) Swimming crabs of the genera Charybdis de Haan, 1833, and Thalamita Latreille, 1829 (Crustacea: Decapoda: Brachyura: Portunidae) from peninsular Malaysia and Singapore., Raffles Bulletin of Zoology Supplement 1.: 1-128

Williams, Austin B. (1984) Shrimps, Lobsters, and Crabs of the Atlantic Coast of the Eastern United States, Maine to Florida, Smithsonian Institution Press, Washington, DC. Pp. <missing location>

Yokes, Baki; Galil, Bella (2006) New records of alien decapods (Crustacea) from the Mediterranean coast of Turkey, with a description of a new palaemonid species, Zoosytema 28(3): 747-755

Yokes, M. Baki and 8 authors (2007) Alien crustacean decapods from the Aegean coast of Turkey., Aquatic Invasions 2(3): 162-168

Zenetos, A. and 22 authors (2011) Alien species in the Mediterranean Sea by 2010. A contribution to the application of European Union’s Marine Strategy Framework Directive (MSFD). Part I. Spatial distribution, Mediterranean Marine Science published online: <missing location>

Zenetos, A.; Vassilopoulou, V.; Salomidi, M.; Poursanidis, D. (2008) Additions to the marine alien fauna of Greek waters (2007 update)., Marine Biodiversity Records 1: e91

Zenetos, Argyro and 6 authors (2009) Aquatic alien species in Greece (2009): tracking sources, patterns and effects on the ecosystem, Journal of Biological Research-Thessaloniki 12: 135-172