- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat List
- Biology and Ecology
- Latitude/Altitude Ranges
- Natural enemies
- Notes on Natural Enemies
- Pathway Causes
- Pathway Vectors
- Environmental Impact
- Risk and Impact Factors
- Uses List
- Links to Websites
- Distribution Maps
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PicturesTop of page
IdentityTop of page
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 InvasivenessTop of page
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 TreeTop of page
- 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 NomenclatureTop of page
Synonymy is given according to Wee and Ng (1995); misspellings commonly encountered in the literature are also included.
DescriptionTop of page
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 TableTop of page
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.Last updated: 10 Feb 2022
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Egypt||Present, Widespread||Native||Red Sea and Tor, Sinai|
|Mozambique||Present, Widespread||Native||Delagoa Bay|
|Somalia||Present, Widespread||Native||Sar Uanle|
|South Africa||Present, Widespread||Native||Natal|
|China||Present||Present based on regional distribution.|
|-Fujian||Present, Widespread||Native||Xiamen, Liuwutien|
|Hong Kong||Present, Widespread||Native||Tolo Harbour in 1986|
|India||Present||Present based on regional distribution.|
|-Andaman and Nicobar Islands||Present, Widespread||Native|
|Indonesia||Present||Present based on regional distribution.|
|-Irian Jaya||Present||Salawati Island|
|-Lesser Sunda Islands||Present, Widespread||Native||Labuhanbajo, East Nusa Tenggara|
|-Maluku Islands||Present, Widespread||Native||Ambon|
|Iran||Present, Widespread||Native||Collected in Bushire in 1937 and Quism Island, Strait of Hormuz in 1938|
|Israel||Present||Introduced||Samples were collected in 1924|
|Japan||Present||Present based on regional distribution.|
|-Kyushu||Present, Widespread||Native||Kagoshima Bay and Tanegashima|
|-Ryukyu Islands||Present, Widespread||Native||Chinen Peninsula, Okinawa|
|Kuwait||Present, Widespread||Native||Off Kuwait in 1995 and Ras al-Jlay’ah in 1998|
|Malaysia||Present||Present based on regional distribution.|
|-Peninsular Malaysia||Present, Widespread||Native||Pontian, Johor|
|Myanmar||Present, Widespread||Native||Mergui Achipelago in 1886|
|Pakistan||Present, Widespread||Native||Manora Island, Karachi in 1966|
|Philippines||Present, Widespread||Native||Cebu in 1983|
|Singapore||Present, Widespread||Native||Siglap, Changi point, Tuas|
|Sri Lanka||Present, Widespread||Native|
|Taiwan||Present, Widespread||Native||Kaohsiung, Changhua and Chiku lagoon|
|United Arab Emirates||Present, Few occurrences||One carapace was only found in Ras al-Khaimah and 5 specimens were collected in Fujairah Port in 1995|
|Cyprus||Present, Widespread||Introduced||First observed in 1999|
|Greece||Present||Introduced||Recorded in Kastellorizo Island and Rhodes in 2004|
|Cuba||Present, Widespread||Introduced||Four specimens from Bahia de Gibara and one specimen from Bahia Cienfuegos were collected in 1987|
|United States||Present||Present based on regional distribution.|
|-Florida||Present, Widespread||Introduced||Indian River lagoon system|
|-Georgia||Present, Few occurrences||Introduced||Live 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|
|-Hawaii||Present, Few occurrences||Introduced||Edmondson 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 Carolina||Present, Widespread||Introduced||Portsmouth Island and Bogue Sound in Carteret|
|-South Carolina||Present, Widespread||Introduced||First specimen was collected in Folly River near Charleston in 1986, but not identified until 2001 (McMillen-Jackson, 2008). Succesively recorded in Charleston and Georgetown|
|Australia||Present||Present based on regional distribution.|
|-Northern Territory||Present, Widespread||Native||Sir Edward Pellew Island group and Darwin|
|-Queensland||Present, Widespread||Native||Endeavour, Cooktown, Cairns, Port Denison, Townsville, Moreton Bay|
|-Western Australia||Present, Widespread||Native||Recorded between C. Bossut and Broome and also in Ninety Mile Beach|
|New Caledonia||Present, Widespread||Native||Type locality of the species|
|Brazil||Present||Present based on regional distribution.|
|-Bahia||Present, Widespread||Introduced||One specimen was collected in Rio Cachoeira, Ilhéus in 2005|
|-Maranhao||Present, Widespread||Introduced||Specimens were collected in Panaquatira, Araçagy, Ponta d'Areia and Jacamim|
|-Parana||Present, Widespread||Introduced||Four males were collected in Baía de Guaratuba|
|-Pernambuco||Present, Widespread||Introduced||Two males and one female were collected in Tamandaré Bay|
|-Piaui||Present, Widespread||Introduced||One female specimen was collected in the estuarine system of Macapá Beach, Luis Correia|
|-Rio de Janeiro||Present, Widespread||Introduced||Recorded in Guanabara Bay from 1995 to 1996|
|-Rio Grande do Norte||Present, Widespread||Introduced||Seven specimens were collected in Galinhos and Macau estuaries|
|-Santa Catarina||Present, Widespread||Introduced||Six males were collected in April 1998 in South Caieira da Barra, Florianopolis|
|-Sao Paulo||Present, Widespread||Introduced||Forty-five males and thirty-one females were collected from March to August 1996 in Ubatuba Bay|
|Colombia||Present, Widespread||Introduced||Samples were collected in 1987|
|French Guiana||Present, Widespread||Introduced||One male specimen was collected in Rémiré Beach, Cayenne in 2001|
|Venezuela||Present, Widespread||Introduced||First recorded in Isla de Margarita, 1987|
History of Introduction and SpreadTop of page
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).
Risk of IntroductionTop of page
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.
HabitatTop of page
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 ListTop of page
|Littoral||Mud flats||Principal habitat||Natural|
|Littoral||Intertidal zone||Principal habitat||Natural|
|Marine||Inshore marine||Principal habitat||Natural|
|Marine||Coral reefs||Principal habitat||Natural|
|Marine||Benthic zone||Principal habitat||Natural|
Biology and EcologyTop of page
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).
Latitude/Altitude RangesTop of page
|Latitude North (°N)||Latitude South (°S)||Altitude Lower (m)||Altitude Upper (m)|
Natural enemiesTop of page
Notes on Natural EnemiesTop of page
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.
Pathway CausesTop of page
Pathway VectorsTop of page
Environmental ImpactTop of page
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 FactorsTop of page
- 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
- Competition - monopolizing resources
- Highly likely to be transported internationally accidentally
- Difficult to identify/detect as a commodity contaminant
- Difficult/costly to control
UsesTop of page
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.
Uses ListTop of page
Human food and beverage
- Meat/fat/offal/blood/bone (whole, cut, fresh, frozen, canned, cured, processed or smoked)
ReferencesTop of page
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.
Almeida AO; Coelho PA; Santos JTA dos; Ferraz NR, 2006. [English title not available]. (Crustàceos decàpodos estuarinos de Ilhéus, Bahia, Brasil.) Biota Neotropica, 6(2):unpaginated. http://www.biotaneotropica.org.br/v6n2/en/abstract?inventory+bn03406022006
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.
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.
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.]
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Dineen J, 2001. Species name: Charybdis hellerii. Fort Pierce, Florida, USA: Smithsonian Marine Station at Fort Pierce. http://www.edu/irlspec/charyb_heller
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.
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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.
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.
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Wee DPC; Ng PKL, 1995. Swimming crabs of the genera Charybdis de Haan, 1833, and Thalamita Latreille, 1829 (Crustacea: Decapoda: Brachyura: Portunidae) from Peninsular Malaysia and Singapore. The Raffles Bulletin of Zoology, Supplement 1:1-128.
Zarenkov NA, 1968. On the crabs of the genus Charybdis de Haan from the collections of Soviet-expedition in the Pacific and Indian Oceans. Vestnik Moskovskogo Universiteta, Ser. Biologia, Pochvovedenie, 1968(2):32-38.
Alcock A W, 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.
Almeida A O, Coelho P A, Santos J T A dos, Ferraz N R, 2006. Estuarine decapod crustaceans of the Ilhéus, Bahia, Brazil. (Crustàceos decàpodos estuarinos de Ilhéus, Bahia, Brasil.). Biota Neotropica. 6 (2), unpaginated. http://www.biotaneotropica.org.br/v6n2/en/abstract?inventory+bn03406022006
Apel M, Spiridonov V A, 1998. Taxonomy and zoogeography of the portunid crabs (Crustacea: Decapoda: Brachyura: Portunidae) of the Arabian Gulf and adjacent waters. In: Fauna of Arabia, Vol. 17. Basle, Switzerland: Pro Entomologia. 159-331.
Bezerra L E A, Almeida A O, 2005. First record of the Indo-Pacific species Charybdis hellerii (A. Milne-Edwards, 1867) (Crustacea: Decapoda: Portunidae) on the littoral coast of Cearà. (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.
CABI, Undated. Compendium record. Wallingford, UK: CABI
CABI, Undated a. CABI Compendium: Status inferred from regional distribution. Wallingford, UK: CABI
CABI, Undated b. CABI Compendium: Status as determined by CABI editor. Wallingford, UK: CABI
Calado T C S, 1996. Presence of Charybdis hellerii (Milne Edwards, 1867) in the littoral waters of Brazil (Decapoda: Portunidae). (Registro de Charybdis hellerii (Milne Edwards, 1867) em àguas do litoral brasileiro (Decapoda: Portunidae).). Bol. Est. Ciên. Mar, Maceió. 175-180.
Chopra B N, Das K N, 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.
Coelho P A, Santos M C F, 2003. Occurrence of Charybdis hellerii (Milne Edwards, 1867) (CRUSTACEA, DECAPODA, PORTUNIDAE) on the littoral of Pernambuco. (Ocorrência de Charybdis hellerii (Milne Edwards, 1867) (CRUSTACEA, DECAPODA, PORTUNIDAE) no litoral de Pernambuco.). Boletim Técnico Cientifico CEPENE. 11 (1), 167-173.
Feres S J C, Lopes A T L, Andrade Santos L de, 2007. First record of Charybdis Hellerii (Milne Edwards, 1867) from the littoral of Maranhense - Brazil. (Crustacea, Decapoda, Portunidae). (Primeiro Registro de Charybdis Hellerii (Milne Edwards, 1867) Para O Litoral Maranhense-Brasil. (Crustàcea, Decapoda, Portunidae).). Boletim Do Laboratório De Hidrobiologia. 77-82.
Ferreira A C, Sankarankuty C, Cunha I M C, Duarte F T, 2001. Yet another record of Charybdis hellerii (A. Milne-Edwards) (Crustacea, Decapoda) from the Northeast of Brazil. Revista Brasileira de Zoologia. 18 (Suppl. 1), 357-358.
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. 15.
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Hernandez G, Bolanos J, 1995. Additions to the anomuran and brachyuran fauna of north eastern Venezuela. In: The Crustacean Society Summer Meeting, May 25-27, 1995 [The Crustacean Society Summer Meeting, May 25-27, 1995.], unpaginated.
Kirmitzoglou I, Kitsos M S, Thessalou-Legaki M, Tselepides A, Koukouras A, 2006. Investigation of the progress and possible expansion of the limits of the lessepsian migratory current regarding Decapoda (Crustacea). In: Proceedings of the 10th International Congress on the Zoogeography and Ecology of Greece and adjacent regions, Patra, Greece, 26-30 June 2006 [Proceedings of the 10th International Congress on the Zoogeography and Ecology of Greece and adjacent regions, Patra, Greece, 26-30 June 2006.], unpaginated.
Kocatas A, 1981. Preliminary list and distribution of the decapod Crustacea in Turkish waters. (Liste preliminaire et repartition des Crustaces Decapodes des eaux Turques.). Rapp. Coram, Int. Mer Medit. 27 (2), 161-162.
Lemaitre R, 1995. Charybdis hellerii (Milne Edwards, 1867), a non-indigenous 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.
Lenz H, 1905. East African decapods and stomatopods collected by Prof. Dr A. Voeltzkow. Scientific results of visits to Madagascar and East Africa in the years 1889-1895. Vol. 3. (Ostafrikanische Dekapoden und Stomatopoden, gesammelt von Herrn Prof. Dr. A. Voeltzkow. A Voeltzkow, Wissenschaftliche Ergebnisse der Reisen in Madagaskar und Ostafrika in den Jahren 1889-1895. Vol. 3.). Abhandlungen der Senckenbergischen naturforschenden Gesellschaft. 27 (4), 341-392.
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Mantelatto F L M, Dias L L, 1999. Extension of the known distribution of Charybdis hellerii (Milne Edwards, 1867) (Decapoda: Portunidae) along the Western Tropical South Atlantic. Crustaceana. 72 (6), 617-620.
Milne Edwards A, 1867. Descriptions of some new brachyuran Decapoda. (Descriptions de quelques espèces nouvelles de Crustacés Brachyures.). Annales de la Société entomologique de France, (ser. 4). 7 (4), 263-288.
Nobili M G, 1906. Crab fauna of the Red Sea. Decapoda and Stomatopoda. (Faune carcinologique de la Mer Rouge. Décapodes et Stomatopodes.). Annales des Sciences naturelles, Zoologie. 9 (4(1-3)), 1-347.
Rüppell E S, 1830. Descriptions and illustrations of 24 species of short-tailed crabs, as a contribution to the natural history of the Red Sea. (Beschreibungen und Abbildungen von 24 Arten kurzschwänzigen krabben, als Beiträge zur Naturgeschichte des Rothen Meeres.). In: Beschreibungen und Abbildungen von 24 Arten kurzschwänzigen krabben, als Beiträge zur Naturgeschichte des Rothen Meeres, Frankfurt a. M, Germany: HL Brönner. 1-28.
Seebens H, Blackburn T M, Dyer E E, Genovesi P, Hulme P E, Jeschke J M, Pagad S, Pyšek P, Winter M, Arianoutsou M, Bacher S, Blasius B, Brundu G, Capinha C, Celesti-Grapow L, Dawson W, Dullinger S, Fuentes N, Jäger H, Kartesz J, Kenis M, Kreft H, Kühn I, Lenzner B, Liebhold A, Mosena A (et al), 2017. No saturation in the accumulation of alien species worldwide. Nature Communications. 8 (2), 14435. http://www.nature.com/articles/ncomms14435
Steinitz W, 1929. Immigration of Indo-Pacific species into the Mediterranean Sea since the start of the Quaternary. (Die Wanderung indopazifischer Arten ins Mittelmeer seit Beginn der Quartärperiode.). Internationale Revue der gesammten Hydrobiologie und Hydrographie. 1-90.
Tavares M, Amouroux J M, 2003. First record of the non-indigenous crab, Charybdis hellerii (A. Milne- Edwards, 1867) from French Guyana (Decapoda, Brachyura, Portunidae). Crustaceana. 76 (5), 625-630.
Wee D P C, Ng P 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. 1-128.
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06/01/10 Original text by:
Baki Yokes, Halic University, Turkey
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