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Portunus segnis

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Datasheet

Portunus segnis

Summary

  • Last modified
  • 19 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Host Animal
  • Preferred Scientific Name
  • Portunus segnis
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Crustacea
  •         Class: Malacostraca
  • Summary of Invasiveness
  • P. segnis is a marine nocturnal crab, native to the Western Indian Ocean, from Pakistan westwards to the Arabian Gulf, the east coast of South Africa, Mozambique, Madagascar and Mauritius. It is one of the earl...

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Pictures

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PictureTitleCaptionCopyright
Portunus segnis; adult.
TitleAdult
CaptionPortunus segnis; adult.
Copyright©Prof. Bella Galil/National Institute of Oceanography/Israel Oceanographic & Limnological Research Israel
Portunus segnis; adult.
AdultPortunus segnis; adult.©Prof. Bella Galil/National Institute of Oceanography/Israel Oceanographic & Limnological Research Israel

Identity

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Preferred Scientific Name

  • Portunus segnis (Forskål, 1775)

Other Scientific Names

  • Portunus mauritianus Ward, 1942
  • Portunus pelagicus (Linnaeus, 1758)
  • Portunus trituberculatus

International Common Names

  • Spanish: jaiba azul
  • French: etrille bleue

Local Common Names

  • : saratan sabih
  • Indian Ocean, Western: blue manna crab; blue swimmer crab; blue swimming crab; flower crab; sand crab; swimming red crab
  • Kenya: kaa kiukizi; mswete
  • Pakistan: googoo tanga; kekra

Summary of Invasiveness

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P. segnis is a marine nocturnal crab, native to the Western Indian Ocean, from Pakistan westwards to the Arabian Gulf, the east coast of South Africa, Mozambique, Madagascar and Mauritius. It is one of the earliest introductions through the Suez Canal, having been recorded in Port Said, Egypt, in 1898. During the 1920s it was widely recorded in the Levant (Israel, Lebanon, Syria, Turkey), and has recently spread as far west as the Tyrrhenian coast of Italy and the Gulf of Gabes, Tunisia.

The introduction of Erythraean biota into the Mediterranean Sea led to displacement, extirpation (local extinction), and changes to habitat structure, although little is known about the mechanisms of the inter-relationships. The impact of P. segnis on native biota is undetermined but given that it is an omnivorous predator much larger than any of the sea’s native portunid crabs and that as adults they lack any predators, it can be assumed that its impact may be negative and that it has the potential to outcompete local taxa. Global warming is expected to favour the spread of this tropical species. It is commercially important in its native range as well as in the Levant. 

Taxonomic Tree

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

Notes on Taxonomy and Nomenclature

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Portunus segnis (Forskål, 1775) was described from material collected near Jeddah, on the Red Sea coast of Saudi Arabia, by Petrus Forskål. Forskål died in Arabia and his notes were published posthumously. It is uncertain whether the type specimens reached Denmark, and they are presumed lost. The specific name “segnis” has not been used since Forskål’s description, and was largely subsumed in P. (Portunus) pelagicus (Linnaeus, 1758). P. pelagicus was regarded as widespread throughout the Indo-West Pacific region and was “generally considered a species without any nomenclatural troubles” (Holthuis, 2004). A recent revision of the genus Portunus Weber, 1795, has provided ample morphological, biogeographical and molecular evidence to resurrect P. segnis (Lai et al., 2010). All early records, with the exception of Ward (1942) and Stephenson and Rees (1967) refer to it as Lupa pelagica, Neptunus pelagicus or Portunus pelagicus.

Description

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Carapace broad (CW/CL c. 2.2-2.3), surface evenly granular, frequently with a short pubescence between granules. Sinuous mesogastric and arched epibranchial ridges as rows of tubercles and a pair of granular elevations in cardiac region present; no other obvious ridges. Nine anterolateral teeth; 1st acutely triangular, larger than those immediately following, 2nd to 8th sharp, 9th very long, projecting laterally. Front may have four teeth except for inner supraorbital teeth; median frontal teeth usually low and obtuse or even confluent and indistinct, leaving a wide gap between spiniform lateral median teeths. Posterolateral junction of carapace rounded. Merus of third maxilliped with anterolateral corner rounded, not expanded laterally. Chelipeds relatively slender and elongate, smooth or minutely granular; merus usually with three spines on anterior border and a single terminal spine on posterodistal corner; manus with proximal and two distal spines on upper face, upper and outer face with five well-developed costae, under surface smooth, inner surface with median low and smooth costa. Ambulatory legs with merus subquadrate, posterodistal border smooth; propodus elongate, with smooth posterior border; natatorial paddle elongate oval, obtusely angled distally. Penultimate segment of male abdomen longer than broad with evenly converging lateral borders. G1 very long and slender, base with slight basal spur, curved with finely tapering tip and spinules in distal part. Female genital opening located in median part of sternite, elongate with long axis directed anteromesially; thickened cuticle along antero- and posterolateral borders (Apel and Spiridonov, 1998).

Largest specimen recorded is an ovigerous female from Rhodes, Greece (187.8 × 84.3 mm) (Corsini Foka et al., 2004).

Lai et al. (2010) describes the differences between the male and female colouring: "males with dark olive green blue carapace with many pale white spots on surface particularly posteriorly and anterolaterally; spots do not tend to merge to form reticulating bands, however, such banding if present is typically thinner than in P. pelagicus. Females similar in pattern to male except that tips of chelipeds are red tinged with a brownish red instead of blue tinged with deep rust red”. Corsini Foka et al. (2004) described a freshly deceased female specimen: “carapace and legs show yellow-whitish spots and lines on a reddish-brown background, the dactyls of chelipeds are reddish-brown, the fingers in the second, third and fourth pereiopods are reddish at the edge and light blue on the surface”.

Distribution

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P. segnis is native to the Western Indian Ocean, from Pakistan westwards to the Arabian Gulf, the east coast of Africa, Madagascar and Mauritius and Red Sea (Lai et al. 2010).

It is one of the earliest introductions through the Suez Canal, having been recorded in Port Said, Egypt, already in 1898 (Fox, 1924, as Neptunus pelagicus). During the 1920s it was widely recorded in the Levant (Israel (Fox, 1924), Lebanon (Steinitz, 1929), Syria (Gruvel, 1930), Turkey (Gruvel, 1928)), and has spread as far west as Italy (Ghisotti,1966; Crocetta, 2006) and the Gulf of Gabes, Tunisia (Rabaoui et al., 2015).  

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

Sea Areas

Indian Ocean, WesternWidespread2010Native Not invasive Lai et al., 2010Western Indian Ocean from Pakistan to South Africa
Mediterranean and Black SeaPresentIntroducedafter 1869 Invasive Lai et al., 2010; Zenetos et al., 2010; Brockerhoff and McLay, 2011Eastern Mediterranean

Asia

BahrainPresentNativeApel and Spiridonov, 1998; Lai et al., 2010
IranPresentNative1842-1843Hosseini, 2009; Naderloo and Turkay, 2012First record collected by Theodor Kotschy off Kharg
IsraelWidespreadIntroduced1924 Invasive Fox, 1924; Galil, 2007; Lai et al., 2010Off Ashjod, off Haifa
KuwaitPresentbefore 1978NativeAl-Mohanna and Subrahmanyam, 2001; Lai et al., 2010Khiran, Kuwait coast
LebanonWidespreadIntroduced1929 Invasive Steinitz, 1929
OmanPresentNativeLai et al., 2010; Khvorov et al., 2012
PakistanPresent2005NativeTirmizi and Kazmi, 1983; Lai et al., 2010Karachi, Korung Creek, Indus Delta, Sindh
Saudi ArabiaWidespreadNative1775Apel and Spiridonov, 1998; Lai et al., 2010Numerous locations
SyriaWidespreadIntroduced1930 Invasive Gruvel, 1930; Lai et al., 2010Fish market, Latakia
TurkeyPresentIntroduced1928 Invasive Gruvel, 1928; Holthuis, 1961; Kocatas and Katagan, 2003; Ozcan et al., 2005; Yokes et al., 2007Aegean coast: Palamut Buku and Gokova
United Arab EmiratesPresentNative1901Nobili, 1906; Apel and Spiridonov, 1998; Lai et al., 2010Abu Dhabi and off Arzana

Africa

EgyptPresentFox, 1924; Lai et al., 2010Introduced and invasive on Mediterranean, native in Red Sea
KenyaWidespread2004Native Not invasive Sigana et al., 2009; Lai et al., 2010Studied in Kilifi Creek, 55 km north of Mombasa city
MadagascarPresent2004NativeCrosnier, 1962; Lai et al., 2010Belaza Tulear, Nose Be
MauritiusPresentNativeLai et al., 2010
MozambiquePresent2004NativeLai et al., 2010Inyack Bay, Inhaca, Maputo Bay
SomaliaPresent1976NativeLai et al., 2010Gresira, Sar Vaule
South AfricaPresent2006NativeLai et al., 2010Durban Bay
SudanPresent1991NativeLai et al., 2010Near Port sudan
TanzaniaPresent1995NativeStephenson, 1972; Kyomo, 1999

Europe

CyprusPresentIntroduced1961Demetropoulos and Neocleous, 1969
GreeceLocalisedIntroduced1991Corsini et al., 2004Rhodes Island, collected 1991-2000
ItalyLocalisedIntroduced1966Ghisotti, 1966; Torchio, 1967; Ariani and Serra, 1969; Crocetta, 2006Livorno harbour, Tyrrhenian Sea

History of Introduction and Spread

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It is one of the earliest introductions through the Suez Canal, having been recorded in Port Said, Egypt, already in 1898 (Fox, 1924, as Neptunus pelagicus). According to Fox (1927) it was first seen in numbers in the Canal between 1889 and 1893, although Krukenberg records one specimen from the Bitter Lakes in 1886. In 1898 observations were made at Port Said, and four years later it was common in the port. During the 1920s it was widely recorded in the Levant (Israel (Fox, 1924), Lebanon (Steinitz, 1929), Syria (Gruvel, 1930) and Turkey (Gruvel, 1928)). It has spread as far west as Italy (Ghisotti, 1966; Crocetta, 2006) and the Gulf of Gabes, Tunisia (Rabaoui et al., 2015). It is established in the Mediterranean Sea. 

Introductions

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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
Mediterranean and Black Sea Indian Ocean, Western >1869 Aquaculture (pathway cause) ,
Fisheries (pathway cause) ,
Interconnected waterways (pathway cause)
Yes Brockerhoff and McLay (2011); Galil (2011) Exact means of dispersal are unknown

Risk of Introduction

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P. segnis is an Erythraean invasive already widely spread in the eastern and central Mediterranean Sea. The warming of the Mediterranean waters may facilitate the establishment of populations in the northern and western reaches of the sea. It may be secondarily introduced in ballast tanks to the rapidly warming Lusitanian province and to the western Atlantic (as did the Erythraean invasive portunid Charybdis hellerii (Milne Edwards, 1867) (Lemaitre, 1995).

Habitat

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In its native range the species is found in coastal and brackish waters, over mud and sand, at 0-40 m (Arabian Gulf; Carpenter et al., 1997), it enters estuaries and lagoons seasonably (Anam and Mostarda, 2012; Naderloo and Türkay, 2012). In the Mediterranean, the species is found under rocks and in rock pools, on sandy or muddy substrate, intertidal to 55 m, occasionally in estuaries (Holthuis and Gottlieb, 1958; Galil et al., 2002).

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Littoral
Coastal areas Principal habitat Harmful (pest or invasive)
Coastal areas Principal habitat Natural
Coastal dunes Principal habitat Harmful (pest or invasive)
Coastal dunes Principal habitat Natural
Mangroves Principal habitat Harmful (pest or invasive)
Mangroves Principal habitat Natural
Mud flats Principal habitat Harmful (pest or invasive)
Mud flats Principal habitat Natural
Intertidal zone Principal habitat Harmful (pest or invasive)
Intertidal zone Principal habitat Natural
Brackish
Estuaries Present, no further details

Biology and Ecology

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Genetics

Lai et al. (2010) investigated the genetics of 45 specimens of P. segnis. P. segnis revealed two co-dominant haplotypes separated from each other by two mutational steps. Haplotype 102 (n=16) was obtained from individuals collected from Mozambique and Madagascar, whereas Haplotype 99 (n=16) is restricted to the Red Sea and Mediterranean Sea.

Inter-specific divergence has been found between P. segnis and the other three species of the P. pelagicus group. Based on total sequence divergence at the COI locus, P. armatus and P. reticulatus form a sister group to P. pelagicus and P. segnis, with an average genetic distance of ~2% between P. armatus and P. reticulatus,3.14% between P. pelagicus and P. segnis, and ~7% divergence between P. armatus/P. reticulatus and P. pelagicus/P. segnis.

Reproductive Biology

In the Iranian Gulf and Gulf of Oman ovigerous females occur throughout the year, with the highest proportion in the fall; spawning occurs year round with a peak in winter (Kamrani et al. 2010; Safaie et al., 2013a). Interestingly, two regional studies provide different data for fecundity; 277,421 -1,114348 eggs, with average fecundity of 662,978 eggs (Kamrani et al., 2010) and 521,027 - 6,656599 eggs, with an average fecundity of 2,397967 (Safaie et al., 2013b). In the Mediterranean the mean number of eggs (fecundity) of 12 ovigerous females (whose mean CW 143.3 ± 6.2 mm) was 777,642 ± 80684 (Rabaoui et al., 2015).

Activity Patterns

It is an active nocturnal predator, buried in daytime, with only its eyes, antennae and gill openings protruding.

Chatterji et al. (1994) noted lunar periodicity in the abundance of P. pelagicus s.l. (possibly P. segnis) in trawl catches along the Goa coast, India, with higher catches during the full moon and the new moon.

Nutrition

P. segnis is an omniverous predator. According to studies of stomach contents, juvenile crabs (<90 mm CW) prefer crustaceans (48.6%) to molluscs (21.5%) and fish (17.5 %), adults (CW 111-150 mm) shift their diet to a higher proportion of fish (26.7%), though crustaceans and molluscs remain principal components (40.5%, 24.5%, respectively), and the largest adults (CW 151-170 mm) consume more fish (29.4%), and reduce the proportions of crustaceans and molluscs (37.5%, 21.6%, respectively) (Hosseini et al 2014). These results are similar to those obtained by Pazooki et al. (2012).

Associations

Chelonibia patula (Ranzani 1820), a cosmopolitan epizoon, was collected from P. segnis in the Levant (Israel, Turkey) (Pasternak et al., 2002; Ozcan, 2012).

Environmental Requirements

P. segnis is tolerant of a wide ranger of temperatures from 13.5°C (winter, Livorno, Italy) to 30°C (summer, SE Levant). It is euryhaline (adapts to a wide range of salinities), moving between brackish estuaries to marine and even hypersaline waters (the Bitter Lakes, suez Canal, fide Krukenberg, 1888). 

Latitude/Altitude Ranges

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Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
43 30

Water Tolerances

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ParameterMinimum ValueMaximum ValueTypical ValueStatusLife StageNotes
Depth (m b.s.l.) 0 55 Harmful For Portunus pelagicus s.l.
Salinity (part per thousand) 10 40 Harmful For Portunus pelagicus s.l.
Water temperature (ºC temperature) 13.5 30 Harmful For Portunus pelagicus s.l.

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Octolasmis Parasite Adult not specific Faisal et al., 2010

Notes on Natural Enemies

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Alsaqabi et al. (2010) studied external and internal parasites of P. segnis (as P. pelagicus) off the Saudi coast of the Arabian Gulf and identified a poecilasmatid parasite identified as Octolasmis sp., and ‘pepper spot’ parasites – a condition brought about by hyperparasitic protozoans infecting trematod cysts, which become melanized.  

The adult crab has no natural enemies in the Mediterranean Sea.

Means of Movement and Dispersal

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Natural Dispersal

Dispersal by currents of larvae is most likely as attested by its temporally sequential records along the Suez Canal, Levantine coast line and further west.

Accidental Introduction

Secondary introductions in ballast tanks or escape/release from farming facilities are possible. 

Pathway Causes

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CauseNotesLong DistanceLocalReferences
Interbasin transfers Yes Yes Galil, 2011
Interconnected waterways Yes Yes Galil, 2011

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
WaterLarvae stages Yes Yes

Impact Summary

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CategoryImpact
Economic/livelihood Positive and negative
Environment (generally) Negative

Environmental Impact

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Impact on Biodiversity

The introduction of Erythraean biota into the Mediterranean Sea led to displacement, extirpation (local extinction), and changes to habitat structure, although little is known about the mechanisms of the inter-relationships (Galil, 2007a,b). The populations of P. segnis, an omnivorous predator much larger than any of the sea’s native portunid crabs, and lacking predators as adults, may outcompete local taxa. 

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • 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
  • Gregarious
Impact outcomes
  • Altered trophic level
  • Modification of natural benthic communities
  • Reduced native biodiversity
  • Threat to/ loss of native species
Impact mechanisms
  • Competition - monopolizing resources
  • Pest and disease transmission
  • Predation
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally
  • Highly likely to be transported internationally deliberately
  • Difficult to identify/detect in the field

Uses

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Economic Value

P. segnis is an important fishery resource in its native region (Chande and Mgaya, 2003). It is listed (as P. pelagicus) in several regional FAO field guides (Bianchi, G., 1985; Carpenter et al 1997; Anam and Mostarda, 2012). It is fished using traps, spears, set nets, baited sticks, and bottom trawls along the coast of Kenya (Anam and Mostarda, 2012), shrimp bottom trawl, bottom set gillnet, fish traps and stake-nets along the Iranian coastline (Safaie et al., 2013). Along the Arabian Peninsula “berried females fetch high market prices” (Carpenter et al., 1997). 

It was already making its way through the Suez Canal in the 1880s and was noticed by the Canal company employees who were enthusiastic amateur fishermen (Fox, 1927). In the 1920s it was fished for sale in the markets at Alexandria and at Haifa (Fox, 1927) and by 1962 (to 1983) it dominated decapod fisheries near Alexandria (Abdel-Razek, 1987). The crab production in Bardawil lagoon started at the beginning of 1986. It is one of the most valuable fishery resources in Bardawil lagoon, on the Mediterranean coast of the Sinai Peninsula, the catch increasing from 491.7 tons in 1995 to 1321.8 tons in 2005 (Abdel Razek et al., 2006). In the 1930s it was considered of high commercial value (Steinitz, 1933); in the 1940s the annual average catch was twenty-two tons (Perlmutter, 1956), and it was reported as “..very common in trawl and beach seine catches” in the 1950s off the Israeli coast (Holthuis and Gottlieb, 1958). In the 2000s it was selling in huge quantities at the fish markets of Sicily (Crocetta, 2006). The species is commercially important on the Mediterranean coast of Turkey, fished by traps, trawls, beach seines, and gill nets (Ozcan, 2012). 

Uses List

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Human food and beverage

  • Meat/fat/offal/blood/bone (whole, cut, fresh, frozen, canned, cured, processed or smoked)

Similarities to Other Species/Conditions

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P. segnis is easily distinguished from the Chesapeake blue crab, Callinectes sapidus, the only similarly sized portunid in the Mediterranean, by the prominent inner spine on the cheliped carpus and, in males, by the triangular abdomen (Galil et al., 2002).

Prevention and Control

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Prevention

P. segnis is an Erythraean invasive already widely spread in the Mediterranean Sea. Care should be taken to prevent secondary introductions in ballast tanks to the rapidly warming Lusitanian province and to the western Atlantic (i.e. the application of the International Convention for the control and management of ships’ ballast water and sediments, IMO, 2004).

Physical/Mechanical Control

Physical/mechanical control is futile as P. segnis has already established flourishing populations in the Mediterranean Sea. Reintroducing a high-salinity barrier and locks should, however, reduce further introductions of propagules through the Suez Canal.

Gaps in Knowledge/Research Needs

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Lai et al. (2010) suggested verification of P. pelagicus s.l. records in order to refine the spatial distribution of the species, with emphasis on regions of suspected sympatry. 

There is the need to study of the impacts of the introduction of P. segnis on the native Mediterranean biota through modelling its spread under different scenarios of rising sea water temperature.

References

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Abdel-Razek FA, 1987. Crab fishery of the Egyptian waters with notes on the bionomics of Portunus pelagicus (L.). Acta Adriatica, 28(1-2):143-154.

Al-Mohanna SY; Subrahmanyam MNV, 2001. Flux of heavy metal accumulation in various organs of the intertidal marine blue crab, Portunus pelagicus (L.) from the Kuwait coast after the Gulf War. Environment International, 27(4):321-326.

Alsaqabi SM; Eshky AA; Albelali AS, 2010. Parasitic Infections In The Blue Crab Swimmer Portunus pelagicus (Linneaus, 1758) Crustacea) found in the Arabian Gulf (Kingdom of Saudi Arabia). Arab Gulf Journal of Scientific Research, 28:185-196.

Anam R; Mostarda E, 2012. Field identification guide to the living marine resources of Kenya. FAO Species Identification Guide for Fishery Purposes. Rome, Italy: FAO, 357 pp.

Anirban Chakraborty; Otta SK; Joseph B; Sanath Kumar; Hossain MS; Indrani Karunasagar; Venugopal MN; Iddya Karunasagar, 2002. Prevalence of white spot syndrome virus in wild crustaceans along the coast of India. Current Science, 82(11):1392-1397.

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.

Archdale MV; Añasco CP; Nakagawa A, 2010. Liftnets compare favorably with pots as harvesting fishing gear for invasive swimming crabs. Journal of Fisheries and Aquatic Science, 5(6):510-516. http://www.academicjournalsinc.com

Ariani AP; Serra V, 1969. Sulla presenza del Portunus pelagicus (L.) in acque italiane, con osservazioni sulla morfologia delle specie (Crustacea Decapoda). (Sulla presenza del Portunus pelagicus (L.) in acque italiane, con osservazioni sulla morfologia delle specie (Crustacea Decapoda).) Archivio Botanico e Biogeografico Italiano, 14(4):187-206.

Audouin V, 1826. Explication sommaire des planches de Crustacés de l'Égypte et de la Syrie, publiées par Jules-César Savigny, Membre de l'Institut; offrant un exposé des caractères naturels des genres, avec la distinction des espèces. Explication sommaire des planches de Crustacés de l?Égypte et de la Syrie, publiées par Jules-César Savigny, Membre de l?Institut; offrant un exposé des caractères naturels des genres, avec la distinction des espèces. Description de l?Égypte, ou recueil des observations et des recherches qui ont été faites en Égypte pendant l?expédition de l?armée française, publiée par les ordres de Sa Majesté l?Empereur Napoléon le Grand. Histoire naturelle. Imprimerie impériale, Paris. Animaux invertébrés, 1(4):77-98.

Batoy CB; Sarmago JF; Pilapil BC, 1987. Breeding season, sexual maturity and fecundity of the blue crab, Portunus pelagicus (L.) in selected coastal waters in Leyte and vicinity, Philippines. Annals of Tropical Research, 9(3):157-177.

Bianchi G, 1985. FAO species identification guide for fishery purposes. Field guide to the commercial marine and brackish-water species of Pakistan. Rome, Italy: FAO, 200 pp.

Brockerhoff A; McLay C, 2011. Human-Mediated spread of Alien Crabs. In: In the wrong place-alien marine crustaceans: distribution, biology and impacts. Invading Nature - Springer Series in Invasion Ecology, 6 [ed. by Galil, B. S. \Clark, P. F. \Carlton (eds), J. T.]. Berlin, Germany: Springer, 27-106.

Bryars SR, 1997. Unpublished PhD thesis. Adelaide, Australia: Flinders University, 256 pp.

Bryars SR; Havenhand JN, 2004. Temporal and spatial distribution and abundance of blue swimmer crab (Portunus pelagicus) larvae in a temperate gulf. Marine and Freshwater Research, 55(8):809-818.

Calman WT, 1927. Report on the Phyllocarida, Cumacea and Stomatopoda (Cambridge Exp. Suez 1924). Trans. Zool. Soc. London, 22:399-401.

Carpenter KE; Krupp F; Jones DA; Zajonz U, 1997. FAO species identification field guide for fishery purposes. The living marine resources of Kuwait, Eastern Saudi Arabia, Bahrain, Qatar, and the United Arab Emirates. Rome, Italy: Food and Agriculture Organization (FAO), viii + 293 pp.

Cavaliere A; Berdar A, 1975. Presenza di Callinectes sapidus Rathbun (Decapoda Brachyura) nello Stretto di Messina. (Presenza di Callinectes sapidus Rathbun (Decapoda Brachyura) nello Stretto di Messina.) Bollettino di Pesca, Piscicoltura e Idrobiologia, 30(2):315-322.

Chande AI; Mgaya YD, 2003. The Fishery of Portunus pelagicus and Species Diversity of Portunid Crabs along the Coast of Dar es Salaam. Tanzania Western Indian Ocean J. Mar. Sci, 2(1):75-84.

Chatterji A; Ansari ZA; Ingole BS; Sreepada RA; Kanti A; Parulekar AH, 1994. Effect of lunar periodicity on the abundance of crabs from the Goa coast. Indian Journal of Marine Sciences, 23(3):180-181.

Corsini Foka M; Kondylatos; G; Economidis PS, 2004. Occurrence of the Lessepsian species Poriunus pelagicus (Crustacea) and Apogon pharaonis (Pisces) in the marine area of Rhodes Island. Mediterranean Marine Science, 5(1):83-89.

Crocetta F, 2006. First record of Portunus pelagicus (Linnaeus, 1758) (Decapoda, Brachyura, Portunidae) in the northern Tyrrhenian Sea. Crustaceana, 79(9):1145-1148.

Crosnier A, 1962. Crustaces decapodes Portunidae. Faune de Madagascar, 16:1-154.

Demetropoulos A; Neocleous D, 1969. The fishes and crustaceans of Cyprus. Fisheries Bulletin Ministry of Agriculture and Natural Resources of Cyprus, 1:3-21.

Duruer EC; Kinacigil T; Soykan O; Tosunoglu Z, 2008. Contribution to some biological and fishery aspects of commercial penaid prawns in Mersin Bay (North-eastern Mediterranean, Turkey). Crustaceana, 81(5):577-585.

Ekman S, 1967. Zoogeography of the sea. London, UK: Siddgwick and Jackson, 419 pp.

Encyclopedia of Life, 2013. Encyclopedia of Life. http://www.eol.org

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Links to Websites

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WebsiteURLComment
GISD/IASPMR: Invasive Alien Species Pathway Management Resource and DAISIE European Invasive Alien Species Gatewayhttps://doi.org/10.5061/dryad.m93f6Data source for updated system data added to species habitat list.
WoRMShttp://www.marinespecies.org/index.php

Contributors

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07/08/2015 Updated by:

Bella Galil, National Institute of Oceanography, Israel

19/09/2013 Original text by:

Ekaterina Shalaeva, Consultant, South Croydon, Greater London, UK

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