Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide


Charybdis hellerii



Charybdis hellerii


  • Last modified
  • 06 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Preferred Scientific Name
  • Charybdis hellerii
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Crustacea
  •         Class: Malacostraca
  • Summary of Invasiveness
  • C. hellerii is a portunid crab which is widely distributed in the Indo-West Pacific. It was introduced to the Mediterranean in the early 1920s via the Suez Canal (...

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C. hellerii collected from a depth of 4 m on muddy sand beneath a stone. Datca, Turkey. June 2003.
CaptionC. hellerii collected from a depth of 4 m on muddy sand beneath a stone. Datca, Turkey. June 2003.
CopyrightMehmet Baki Yokes
C. hellerii collected from a depth of 4 m on muddy sand beneath a stone. Datca, Turkey. June 2003.
AdultC. hellerii collected from a depth of 4 m on muddy sand beneath a stone. Datca, Turkey. June 2003.Mehmet Baki Yokes
C. hellerii from a depth of 7 m on a rocky bottom. Tekirova, Antalya, Turkey.  August 2001.
CaptionC. hellerii from a depth of 7 m on a rocky bottom. Tekirova, Antalya, Turkey. August 2001.
CopyrightHasan Yokes
C. hellerii from a depth of 7 m on a rocky bottom. Tekirova, Antalya, Turkey.  August 2001.
AdultC. hellerii from a depth of 7 m on a rocky bottom. Tekirova, Antalya, Turkey. August 2001.Hasan Yokes
C. hellerii from a depth of 2 m on a rocky bottom. Arsuz, Iskenderun, Turkey. October 2006.
CaptionC. hellerii from a depth of 2 m on a rocky bottom. Arsuz, Iskenderun, Turkey. October 2006.
CopyrightMehmet Baki Yokes
C. hellerii from a depth of 2 m on a rocky bottom. Arsuz, Iskenderun, Turkey. October 2006.
AdultC. hellerii from a depth of 2 m on a rocky bottom. Arsuz, Iskenderun, Turkey. October 2006.Mehmet Baki Yokes


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Preferred Scientific Name

  • Charybdis hellerii (A. Milne-Edwards, 1867)

Other Scientific Names

  • Charybdis (Charybdis) helleri
  • Charybdis (Goniosoma) merguiensis
  • Charybdis hellery
  • Charybdis merguiensis
  • Goniosoma hellerii A. Milne-Edwards, 1873
  • Goniosoma merguiense
  • Goniosoma merguiensis

International Common Names

  • English: Indo-Pacific swimming crab; spiny hands

Local Common Names

  • Japan: ishigani
  • Philippines: kasag

Summary of Invasiveness

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C. hellerii is a portunid crab which is widely distributed in the Indo-West Pacific. It was introduced to the Mediterranean in the early 1920s via the Suez Canal (Steinitz, 1929). In the late 1980s, it was further introduced to the western Atlantic via ballast waters from ships coming from eastern Mediterranean ports (Campos and Türkay, 1989; Galil and Zenetos, 2002). Populations with high densities of adults, young and ovigerous females confirm that C. hellerii is well established in the eastern Mediterranean and western Tropical Atlantic (Mantelatto and Dias, 1999; Galil et al., 2002) and in some locations it has become more abundant than the local species (Carqueija, 2000). Although C. hellerii is accepted as an “invasive species” by some authors (Dineen et al., 2001), it has not yet been shown that its successful establishment and spread threatens native ecosystems, habitats or species. Possible impacts on native communities and the economy need to be quantified.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Arthropoda
  •             Subphylum: Crustacea
  •                 Class: Malacostraca
  •                     Subclass: Eumalacostraca
  •                         Order: Decapoda
  •                             Family: Portunidae
  •                                 Genus: Charybdis
  •                                     Species: Charybdis hellerii

Notes on Taxonomy and Nomenclature

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Synonymy is given according to Wee and Ng (1995); misspellings commonly encountered in the literature are also included.


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Carapace hexagonal, convex, pilose; all anterior carapace ridges present and granular, none behind epibranchials; six prominent frontal teeth: two inner orbitals plus four blunt submedians, medians eliptical, on lower plane, projecting beyond submedians, laterals acutely triangular, separated from submedians by deep V-shaped notch; inner supraorbitallobe broadly triangular, outer infraorbitallobe with convex border; anterolateral margin with six sharp, black, brown or red-tipped teeth, separated by deep notches, anteriormost tooth smallest. Basal antennal article expanded, excluding antennal flagellum from orbit. Chelipeds massive and unequal, surface finely pubescent; anterior border of merus with three spines and a spinule at distal end; carpus with strong spine on interior margin and three carinae on external surface; chela bearing four spines on superior surface, single spine near carpal articulation, lower surface smooth; merus and carpus of fifth leg bearing spine on posterior margin, posterior margin of propodus denticulate, dactyl ovate, first male pleopod tapering, distally spinose, curved distally; colour mottled brownish-grey, chelipeds with spines distally brown, fingers distally black with white tips (Wee and Ng, 1995; Galil et al., 2002). Maximum carapace length for males is about 8 cm (Wee and Ng, 1995; Dineen et al., 2001) and for ovigerous females it is 5.7 cm (Mantelatto and Garcia, 2001).

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes


ChinaPresentPresent based on regional distribution.
-FujianWidespreadNativeDai and Yang, 1991Xiamen, Liuwutien
-GuangdongWidespreadNativeDai and Yang, 1991
-GuangxiWidespreadNativeDai and Yang, 1991
-Hong KongWidespreadNativeApel and Spiridonov, 1998Tolo Harbour in 1986
IndiaPresentPresent based on regional distribution.
-Andaman and Nicobar IslandsWidespreadNativeAlcock, 1899
IndonesiaPresentPresent based on regional distribution.
-Irian JayaPresentLeene, 1938Salawati Island
-JavaWidespreadNativeLeene, 1938
-MoluccasWidespreadNativeSpiridonov, 1999Ambon
-Nusa TenggaraWidespreadNativeStephenson, 1972a; Leene, 1938Labuhanbajo, East Nusa Tenggara
-SumatraWidespreadNativeLeene, 1938
IranWidespreadNativeApel and Spiridonov, 1998Collected in Bushire in 1937 and Quism Island, Strait of Hormuz in 1938
IsraelPresentIntroducedSteinitz, 1929Samples were collected in 1924
JapanPresentPresent based on regional distribution.
-KyushuWidespreadNativeUrita, 1926Kagoshima Bay and Tanegashima
-Ryukyu ArchipelagoWidespreadNativeMinei, 1971; Sakai, 1976Chinen Peninsula, Okinawa
KuwaitWidespreadNativeApel and Spiridonov, 1998Off Kuwait in 1995 and Ras al-Jlay’ah in 1998
LebanonPresentIntroducedShiber, 1981
MalaysiaPresentPresent based on regional distribution.
-Peninsular MalaysiaWidespreadNativeWee and Ng, 1995Pontian, Johor
MyanmarWidespreadNativeChopra and Das, 1937; Apel and Spiridonov, 1998Mergui Achipelago in 1886
OmanWidespreadNativeApel and Spiridonov, 1998
PakistanWidespreadNativeApel and Spiridonov, 1998Manora Island, Karachi in 1966
PhilippinesWidespreadNativeApel and Spiridonov, 1998Cebu in 1983
SingaporeWidespreadNativeWee and Ng, 1995Siglap, Changi point, Tuas
Sri LankaWidespreadNativeAlcock, 1899
SyriaWidespreadIntroducedKuznetsov et al., 1993
TaiwanWidespreadNativeChou et al., 1999Kaohsiung, Changhua and Chiku lagoon
ThailandWidespreadNativeStephenson, 1972a
TurkeyPresentIntroducedKocatas, 1981; Yokes and Galil, 2006; Yokes et al., 2007Iskenderun Bay
United Arab EmiratesPresent, few occurrencesApel and Spiridonov, 1998One carapace was only found in Ras al-Khaimah and 5 specimens were collected in Fujairah Port in 1995


DjiboutiWidespreadNativeNobili, 1906
EgyptWidespreadNativeRüppell, 1830; Balss, 1936Red Sea and Tor, Sinai
MadagascarWidespreadNativeLenz, 1905; Crosnier, 1962Mahajanga
MozambiqueWidespreadNativeBarnard, 1950; Apel and Spiridonov, 1998Delagoa Bay
SomaliaWidespreadNativeVannini and Innocenti, 2000Sar Uanle
South AfricaWidespreadNativeBarnard, 1950Natal

North America

USAPresentPresent based on regional distribution.
-FloridaWidespreadIntroducedLemaitre, 1995; McMillen-Jackson, 2008Indian River lagoon system
-GeorgiaPresent, few occurrencesIntroducedFrick and Williams, 2006Live specimens have not yet been recorded in Georgia; however, remains of C. hellerii were found in the gastrointestinal tract of a dead Kemp’s Ridley Sea Turtle Lepidochelys kempii found on the beach of Wassaw Island. C. hellerii is abundantly found on the Atlantic coast of Florida, South Carolina and North Carolina. Most probably it also inhabits the coasts of Georgia
-HawaiiPresent, few occurrencesIntroducedEdmondson, 1954Edmondson found one male specimen among the fouling on the hull of a ship in dry dock in Pearl Harbour Navy Yard. The ship had seen service in Guam sometime previous to its defouling; thus Edmondson indicated that the crab might have been transported to Pearl Harbour as a juvenile. No further records (Coles et al., 1997, 2009)
-North CarolinaWidespreadIntroducedPortsmouth Island and Bogue Sound in Carteret
-South CarolinaWidespreadIntroducedFirst specimen was collected in Folly River near Charleston in 1986, but not identified until 2001 (McMillen-Jackson, 2008). Succesively recorded in Charleston and Georgetown

Central America and Caribbean

CubaWidespreadIntroducedGómez and Martínez-Iglesias, 1990Four specimens from Bahia de Gibara and one specimen from Bahia Cienfuegos were collected in 1987

South America

BrazilPresentPresent based on regional distribution.
-AlagoasWidespreadIntroducedCalado, 1996
-BahiaWidespreadIntroducedAlmeida et al., 2006One specimen was collected in Rio Cachoeira, Ilhéus in 2005
-CearaWidespreadIntroducedBezerra and Almeida, 2005
-MaranhaoWidespreadIntroducedFeres et al., 2007Specimens were collected in Panaquatira, Araçagy, Ponta d'Areia and Jacamim
-ParanaWidespreadIntroducedFrigotto and Serafim-Junior, 2007Four males were collected in Baía de Guaratuba
-PernambucoWidespreadIntroducedCoelho and Santos, 2003Two males and one female were collected in Tamandaré Bay
-PiauiWidespreadIntroducedLima-Júnior et al., 2008One female specimen was collected in the estuarine system of Macapá Beach, Luis Correia
-Rio de JaneiroWidespreadIntroducedTavares and Mendonça, 1996Recorded in Guanabara Bay from 1995 to 1996
-Rio Grande do NorteWidespreadIntroducedFerreira et al., 2001Seven specimens were collected in Galinhos and Macau estuaries
-Santa CatarinaWidespreadIntroducedMantelatto and Dias, 1999Six males were collected in April 1998 in South Caieira da Barra, Florianopolis
-Sao PauloWidespreadIntroducedMantelatto and Dias, 1999Forty-five males and thirty-one females were collected from March to August 1996 in Ubatuba Bay
ColombiaWidespreadIntroducedCampos and Türkay, 1989Samples were collected in 1987
French GuianaWidespreadIntroducedTavares and Amouroux, 2003One male specimen was collected in Rémiré Beach, Cayenne in 2001
VenezuelaWidespreadIntroducedHernandez and Bolanos, 1995First recorded in Isla de Margarita, 1987


CyprusWidespreadIntroducedKatsanevakis et al., 2009First observed in 1999
GreecePresentIntroducedKirmitzoglou et al., 2006Recorded in Kastellorizo Island and Rhodes in 2004


AustraliaPresentPresent based on regional distribution.
-Australian Northern TerritoryWidespreadNativeStephenson et al., 1957Sir Edward Pellew Island group and Darwin
-QueenslandWidespreadNativeStephenson et al., 1957; Stephenson et al., 1957Endeavour, Cooktown, Cairns, Port Denison, Townsville, Moreton Bay
-Western AustraliaWidespreadNativeStephenson et al., 1957Recorded between C. Bossut and Broome and also in Ninety Mile Beach
New CaledoniaWidespreadNativeMilne Edwards, 1867Type locality of the species

History of Introduction and Spread

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C. hellerii is widely distributed in the Indo-West Pacific. It also inhabits the Red Sea. The first Mediterranean record was from Palestine as Charybdis (Goniosoma) merguiensis, based on the specimens collected in 1924-1925 (Steinitz, 1929). The Suez Canal was suggested to be the most likely mode of introduction (Galil et al., 2002). The crab was rapidly dispersed to other eastern Mediterranean countries by larval transport with local currents and successively recorded in Egypt (Balss, 1936), Turkey (Kocatas, 1981; Yokes and Galil, 2006; Yokes et al., 2007), Lebanon (Shiber, 1981), Syria (Kuznetsov et al., 1993), Greece (Kirmirtzoglou et al., 2006) and Cyprus (Katsanevakis et al., 2009). The population density decreases from east to west on the Mediterranean coast of Turkey and also from south to north in the Aegean Sea, suggesting that water temperature is the limiting factor in its spread (B Yokes, Halic University, Turkey, personal communication, 2010).

C. hellerii has not yet been recorded in the central or eastern Pacific, except one record from Pearl Harbour, Hawaii (Edmondson, 1954). One male specimen was found among fouling on the hull of a ship in dry dock in the Pearl Harbour Navy Yard, Oahu. The ship had seen service in Guam sometime after it had last been defouled; thus the crab may have been transported to Pearl Harbour as a juvenile. Surveys of alien species have been routinely conducted in harbours and lagoons of Oahu, but no further specimens have been observed since (Cole et al., 1997; 2009).
C. hellerii has also been introduced to the western Tropical Atlantic and the first records were from the Caribbean Sea. In 1987 and 1988, specimens were collected in Colombia (Bahia de Portete, Guajira and Bahia Chengue, Magdalena) (Campos and Türkay, 1989); in Cuba (Bahia de Gibara, NE Cuba and Bahia Cienfuegos, S Cuba) (Gomez and Martinez-Iglesias, 1990) and in Venezuela (Isla de Margarita, Golfo de Cariaco) (Hernandez and Bolanos, 1995). Although the publication years of these records are different, all the specimens were collected in 1987-1988, suggesting that this species had already dispersed throughout the entire Caribbean before it was first noticed. Moreover, in the oldest record from South Carolina, five juveniles were collected from the Folly River near Charleston in 1986, but were not identified until 2001 (McMillen-Jackson, 2008; U.S. Geological Survey, Nonindigenous Aquatic Species, Specimen ID # 164602). In 1995, C. hellerii was collected in the Indian River lagoon system, Florida (Lemaitre, 1995), and it was subsequently recorded from the continental United States as follows: In Florida it was reported from Amelia Island (2000), the intracoastal waterway (2002), Matanzas River Inlet (2002), Fort George Inlet (2003), Nassau River (2004), Anna Maria Island (2004), Tolomato River (2004), Bellis River (2004), Palm Coast (2004), Amelia River (2004), Long Key (2009) and Ponce de Leon Inlet (2009); in North Carolina from Carteret (Portsmouth Island in 2004 and Bogue Sound in 2006 and 2008); and in South Carolina from Georgetown, North Inlet Estuary and Winyah Bay (2000), and from Charleston, West bank of Town Creek (2002) (U.S. Geological Survey, Nonindigenous Aquatic Species, 2009; McMillen-Jackson, 2008). Although the crab is abundantly found on the Atlantic coasts of Florida, South Carolina and North Carolina, live specimens have not yet been collected in Georgia. Remains of C. hellerii were, however, found in the gastrointestinal tract of a Kemp’s Ridley sea turtle Lepidochelys kempii found dead on the beach of Wassaw Island, suggesting that C. hellerii may also inhabit the coasts of Georgia (Frick and Williams, 2006).
After it was first observed in the Caribbean Sea and along the coast of the continental United States, C. hellerii appeared to have dispersed southwards and was successively recorded on the coasts of Brazil: Alagoas (Calado, 1996); Rio de Janeiro (Tavares and Mendonça, 1996); Sao Paulo (Mantelatto and Dias, 1999); Rio Grande do Norte (Ferreira et al., 2001); Pernambuco (Coelho and Santos, 2003); Santa Catarina (Mantelatto et al., 2004); Ceara (Bezerra and Almeida, 2005); Bahia (Almeida et al., 2006); Maranhao (Feres et al., 2007) and Piaui (Lima-Junior et al., 2008).
Shipping from eastern Mediterranean ports is thought to have been the method of introduction of C. hellerii into the western Atlantic Ocean in the late 1980s (Campos and Türkay, 1989; Galil and Zenetos, 2002).

Risk of Introduction

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C. hellerii has a long larval life (44 days) which is much more than the transit time of ships crossing the Atlantic Ocean. This characteristically long larval period clearly enhances the potential of the species for long-distance dispersal via ballast water or in hull fouling (Dineen et al., 2001). Founder populations are easily established because of the crab's rapid growth and short generation time. It is proposed that the western Atlantic population was introduced from the eastern Mediterranean via ballast water (Galil and Zenetos, 2002). One individual of C. hellerii was found in the sea-chest of a fishing vessel in New Zealand (Dodgshun and Coutts, 2003). The record for Hawaii was of fouling on the hull of a ship in dry dock (Edmondson, 1954). Thus, C. hellerii has proved to have a high potential for further introductions to new locations via shipping.


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C. hellerii has been reported from a variety of habitats. It can be found on soft bottoms, such as sand and/or mud flats with scattered stones (Spiridonov, 1999). It has also been recorded from hard substrata such as under rocks and among live corals (Lemaitre, 1995), from rock pools (Mustaquim and Rabbani, 1976), from Thalassia meadows and on mangrove roots (Rhizophora mangle) (Campos and Türkay, 1989). Juveniles have been found in association with the bryozoan Schizoporella unicornis (Mantelatto and Souza-Carey, 1998). The species can be found from the intertidal zone down to a depth of 51 m (Lemaitre, 1995).

Habitat List

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Mangroves Principal habitat Natural
Mud flats Principal habitat Natural
Intertidal zone Principal habitat Natural
Estuaries Principal habitat Natural
Lagoons Principal habitat Natural
Inshore marine Principal habitat Natural
Coral reefs Principal habitat Natural
Benthic zone Principal habitat Natural

Biology and Ecology

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C. hellerii matures at a size smaller than that of most other Charybdis spp. In the laboratory, one female reached maturity at a size of 77 mm in 12 months (Dineen et al., 2001). C. hellerii is thought to be capable of storing sperm for at least five months and it can produce at least six broods per year (Dineen et al., 2001). Fecundity in C. hellerii is high and ranges from 22,550 to 3,200,000 eggs per brood depending on the size of the female (Sumpton, 1990; Siddiqui and Ahmed, 1992; Lemaitre, 1995). Zoeal development occurs through six stages and takes 40 days, which is more than twice as long as the 15 days reported for Charybdis callianassa (Greenwood and Fielder, 1980) and significantly longer than the 23 days reported for Charybdis acuta (Kurata and Omi, 1969). With an additional 4 days of the megalopa stage, total larval development is completed in 44 days, which is also longer than the 30-day larval period for C. acuta (Kurata and Omi, 1969; Dineen et al., 2001).

C. hellerii has a generalized carnivorous diet and can exploit a variety of food resources opportunistically (Dineen et al., 2001).

Latitude/Altitude Ranges

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Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
37 19

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Lepidochelys kempii Predator Adult not specific Frick and Williams, 2006
Octopus vulgaris Predator Adult not specific Sampaio and Rosa, 2006

Notes on Natural Enemies

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Large groupers (Epinephelus spp. and Mycteroperca spp.) may be potential predators of C. hellerii (Sampaio and Rosa, 2006). It has been reported that Octopus vulgaris also feeds on C. hellerii on reefs in northeastern Brazil (Sampaio and Rosa, 2006). Remains of an adult male of C. hellerii, together with the remains of six native crab species, were found in the gastrointestinal tract of a Kemp’s Ridley Sea Turtle (Lepidochelys kempi) in Georgia, USA (Frick and Williams, 2006), suggesting that turtles may be predators of C. hellerii.

Environmental Impact

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It has not yet been shown that the successful establishment and spread of C. hellerii threatens native ecosystems, habitats or species. However, C. hellerii  is a territorial omnivore which often feeds on other small crabs. Thus an established population may adversely affect local crab populations. Also Charybdis hellerii could potentially compete for food and habitat with native brachyuran crabs; for example there is concern in Florida that the fishery for Callinectes sapidus could be negatively impacted (Dineen, 2001).

C. hellerii is a potential host of the White Spot Syndrome Virus (WSSV), which naturally infects Charybdis spp. as well as a number of other species of decapods (Joint Subcommittee on Aquaculture, 1997; Tavares and Amouroux, 2003). The virus can be transferred from natural environments to aquaculture facilities and cause lethal outbreaks. WSSV viruses have been reported to decimate crops of penaeid shrimps (Joint Subcommittee on Aquaculture, 1997). Possible impacts on native communities and local economies remain to be quantified.

Risk and Impact Factors

Top of page Invasiveness
  • Has a broad native range
  • Abundant in its native range
  • Highly adaptable to different environments
  • Is a habitat generalist
  • Capable of securing and ingesting a wide range of food
  • Fast growing
  • Has high reproductive potential
Impact mechanisms
  • Competition - monopolizing resources
  • Predation
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally
  • Difficult to identify/detect as a commodity contaminant
  • Difficult/costly to control


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C. hellerii is of commercial value in South-East Asia (Lemaitre, 1995) and has been cultivated in the Philippines (Southeast Asian Fisheries Development Centre Annual Report, 1977); however, in the Mediterranean and Atlantic it currently does not have any market value.


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Alcock AW, 1899. Materials for a carcinological fauna of India. No. 4. The Brachyura Cyclometopa. Part 2. A revision of the Cyclometopa with an account of the families Portunidae, Cancridae and Corystidae. Journal of the Asiatic Society of Bengal, Calcutta, 68(part 2(1)):1-104.

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Apel M; Spiridonov VA, 1998. Taxonomy and zoogeography of the portunid crabs (Crustacea: Decapoda: Brachyura: Portunidae) of the Arabian Gulf and adjacent waters. Fauna of Arabia, 17:159-331.

Balss H, 1936. Decapoda. The fishery grounds near Alexandria. VII. Fish. Research Direct. Not. and Mem., Cairo, 15:1-67.

Barnard KH, 1950. Descriptive catalogue of South African Decapod Crustacea (crabs and shrimps). Annals of the South African Museum, 38:1-837.

Bezerra LEA; Almeida AO, 2005. [English title not available]. (Primeiro registro da espécie Indo-Pacífica Charybdis hellerii (A. Milne-Edwards, 1867) (Crustacea: Decapoda: Portunidae) para o litoral do Estado do Cearà.) Tropical Oceanography, 33(1):33-38.

Calado TCS, 1996. [English title not available]. (Registro de Charybdis hellerii (Milne Edwards, 1867) em àguas do litoral brasileiro (Decapoda: Portunidae).) Bol. Est. Ciên. Mar, Maceió, 9:175-180.

Campbell BM; Stephenson W, 1970. The sublittoral Brachyura (Crustacea: Decapoda) of Moreton Bay. Memoirs of the Queensland Museum, 15(4):235-301.

Campos NH; Türkay M, 1989. On a record of Charybdis helleri from the Caribbean coast of Colombia (Crustacea: Decapoda: Portunidae). Senckenbergiana maritima, 20(3/4):119-123.

Carqueija CRG, 2000. [English title not available]. (Situação atual e impactos da introdução da espécie exótica Charybdis hellerii (Decapoda, Brachyura, Portuinidae) na costa da Bahia.) In: Resumos do I. Congresso Brasileiro Sobre Crustàceos, October 16-20, São Pedro, SP, 2000. unpaginated.

Chopra BN, 1935. Further notes on Crustacea Decapoda in the Indian Museum. VIII. On the Decapod Crustacea collected by the Bengal Pilot Service off the mouth of the river Hooghly. Brachygnatha (Oxyrhyncha and Brachyrhyncha). Record of the Indian Museum, 37(4):463-514.

Chopra BN; Das KN, 1937. Further notes on Crustacea Decapoda in the Indian Museum. IX. On three collections of crabs from Tavoy and Mergui Archipelago. Record of the Indian Museum, Calcutta, 39(4):377-434.

Chou WR; Lai SH; Fang LS, 1999. Benthic crustacean communities in waters of Southwestern Taiwan and their relationships to environmental characteristics. Acta Zoologica Taiwanica, 10(1):25-33.

Coelho PA; Santos MCF, 2003. [English title not available]. (Ocorrência de Charybdis hellerii (Milne Edwards, 1867) (CRUSTACEA, DECAPODA, PORTUNIDAE) no litoral de Pernambuco.) Boletim Técnico Cientifico CEPENE, 11(1):167-173.

Cole SL; Bolick H; Hauk B; Montgomery A, 2009. Ten year resurveys of the biodiversity of marine communities and introduced species in Pearl Harbor, Honolulu Harbor and Ke'ehi Lagoon, O'ahu, Hawaii. Ten year resurveys of the biodiversity of marine communities and introduced species in Pearl Harbor, Honolulu Harbor and Ke'ehi Lagoon, O'ahu, Hawaii. Honolulu, Hawaii: Bishop Museum Press, 215 pp. [Bishop Museum Technical Report No. 48.]

Cole SL; DeFelice RC; Eldredge LG; Carlton JT, 1997. Biodiversity of marine communities in Pearl Harbor, Oahu, Hawaii with observations on introduced exotic species. Biodiversity of marine communities in Pearl Harbor, Oahu, Hawaii with observations on introduced exotic species. Honolulu, Hawaii: Bishop Museum Press, 237 pp. [Bishop Museum Technical Report No. 10.]

Crosnier A, 1962. [English title not available]. (Crustacés décapodes Portunidae.) Faune de Madagascar, 16:1-154.

Dai A; Yang S, 1991. Crabs of the China Seas. Beijing: China Ocean Press, 608 pp.

Dineen J, 2001. Species name: Charybdis hellerii. Fort Pierce, Florida, USA: Smithsonian Marine Station at Fort Pierce.

Dineen JE; Clark PF; Hines AH; Reed SA; Walton HP, 2001. Life history, larval description, and natural history of Charybdis hellerii (Decapoda, Brachyura, Portunidae), an invasive crab in the western Atlantic. Journal of Crustacean Biology, 21:774-805.

Dodgshun T; Coutts A, 2003. Opening the lid on sea chests. Seafood New Zealand, 11:35.

Edmondson CH, 1954. Hawaiian Portunidae. Occasional Papers of the Bernice P Bishop Museum, Honolulu, 21(12):217-274.

Feres SJC; Lopes ATL; Andrade Santos L de, 2007. [English title not available]. (Primeiro Registro de Charybdis Hellerii (Milne Edwards, 1867) Para O Litoral Maranhense-Brasil. (Crustàcea, Decapoda, Portunidae).) Boletim Do Laboratório De Hidrobiologia, 20:77-82.

Ferreira AC; Sankarankuty C; Cunha IMC; Duarte FT, 2001. Yet another record of Charybdis hellerii (A. Milne-Edwards) (Crustacea, Decapoda) from the Northeast of Brazil. Revista Brasileira de Zoologia, 18(supp. 1):357-358.

Fourmanoir P, 1954. [Crabs from the west coast of Madagascar]. (Crabes de la côte ouest de Madagascar.) Le Naturaliste Malgasche, 6(1/2):1-16.

Frick M; Williams K, 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.

Frigotto SF; Serafim-Junior M, 2007. First record of Charybdis hellerii (Milne Edwards, 1867) (Crustacea) in the littoral State of Parana. Estud. Biol, 29(67):227-230.

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06/01/10 Original text by:

Baki Yokes, Halic University, Turkey

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