Portunus segnis

Pictures

Picture	Title	Caption	Copyright
Title Adult Caption Portunus segnis; adult. Copyright ©Prof. Bella Galil/National Institute of Oceanography/Israel Oceanographic & Limnological Research Israel	Adult	Portunus segnis; adult.	©Prof. Bella Galil/National Institute of Oceanography/Israel Oceanographic & Limnological Research Israel

Identity

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

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

• 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

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

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; 1^st acutely triangular, larger than those immediately following, 2^nd to 8^th sharp, 9^th 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

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

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.

History of Introduction and Spread

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

Risk of Introduction

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

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

Biology and Ecology

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

Water Tolerances

Natural enemies

Notes on Natural Enemies

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

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

Pathway Vectors

Impact Summary

Environmental Impact

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

• 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

• Altered trophic level
• Modification of natural benthic communities
• Reduced native biodiversity
• Threat to/ loss of native species

• Competition - monopolizing resources
• Pest and disease transmission
• Predation

• Highly likely to be transported internationally accidentally
• Highly likely to be transported internationally deliberately
• Difficult to identify/detect in the field

Uses

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

Human food and beverage

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

Similarities to Other Species/Conditions

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

Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

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

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.

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Kyomo J, 1999. Distribution and abundance of crustaceans of commercial importance in Tanzania mainland coastal waters. Bulletin of Marine Science, 65(2):321-335.

Lai JCY; Ng PKL; Davie PJF, 2010. A revision of the Portunus pelagicus (Linnaeus, 1758) species complex (Crustacea: Brachyura: Portunidae), with the recognition of four species. Raffles Bulletin of Zoology, 58(2):199-237.

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Zenetos A; Gofas S; Verlaque M; Cinar ME; Garcia Raso JE; Bianchi CN; Morri C; Azzurro E; Bilecenoglu M; Froglia C; Siokou I; Violanti D; Sfriso A; San Martin G; Giangrande A; Katagan T; Ballesteros E; Ramos-Espla A; Mastrototaro F; Ocana O; Zingone A; Gambi MC; Streftaris N, 2010. 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, 11(2):381-493. http://www.medit-mar-sc.net/files/201102/23-103521437_MMS_v11n2_ZENETOS.pdf

Distribution References

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. In: Environment International, 27 (4) 321-326.

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Ghisotti F, 1966. (Il Callinectes sapidus Rathbun nel Mediterraneo (Crustacea, Decapoda)). In: Natura (Milano), 57 (3) 177-180.

Gruvel A, 1928. (Repartition geographique de quelques Crustaces comestibles sur les cotes d'Egypte et de Syrie). In: Comptes rendus de la société de biogéographie, 5 (39) 45-56.

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Khvorov SA, Al-Sinawi MN, Al-Kharusi LH, Al-Mosharfi MN, 2012. Catalogue of brachyuran crabs (Decapoda) of the Sultanate of Oman: Part I. The Arabian Sea coast (Dhofar, Al-Wusta, southern part of Sharqiya). In: Crustaceana, 85 (14) 1735-1744.

Kocatas A, Katagan T, 2003. The Decapod Crustacean fauna of the Turkish Seas. In: Zoology in the Middle East, 29 63-74.

Kyomo J, 1999. Distribution and abundance of crustaceans of commercial importance in Tanzania mainland coastal waters. Bulletin of Marine Science. 65 (2), 321-335.

Lai JCY, Ng PKL, Davie PJF, 2010. A revision of the Portunus pelagicus (Linnaeus, 1758) species complex (Crustacea: Brachyura: Portunidae), with the recognition of four species. In: Raffles Bulletin of Zoology, 58 (2) 199-237.

Naderloo R, Turkay M, 2012. Decapod crustaceans of the littoral and shallow sublittoral Iranian coast of the Persian Gulf: Faunistics, Biodiversity and Zoogeography. In: Zootaxa, 3374 1-67.

Nobili G, 1906. (Faune carcinologique de la Mer Rouge. Décapodes et Stomatopodes). In: Annales des Sciences Naturelles, Zoologie (Series 9), 4 1-347.

Ozcan T, Katagan T, Kocatas A, 2005. Brachyuran crabs from Iskenderun Bay (southeastern Turkey). In: Crustaceana, 78 (2) 237-243.

Sigana DO, Mavuti KM, Ruwa RK, 2009. Fish species composition and distribution in Kilifi Creek. In: Advances in Coastal Ecology People, processes and ecosystems in Kenya, 20 15-27.

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.

Stephenson W, 1972. An annotated check list and key to the Indo-West-Pacific swimming crabs (Crustacea: Decapoda: Portunidae). Bulletin of the Royal Society of New Zealand. 1-64.

Tirmizi NM, Kazmi QM, 1983. Carcinological studies in Pakistan, with remarks on species common to the Red Sea and the Mediterranean. In: Bulletin of the Institute of Oceanography and Fisheries, 9 347-380.

Torchio M, 1967. (Il Callinectes sapidus Rathbun nelle acque siciliane (Crustacea, Decapoda)). In: Natura, 58 (1) 81.

Yokes M B, Karhan S Ü, Okus E, Yüksek A, Aslan-Yİlmaz A, Yİlmaz I N, Demİrel N, Demİr V, Galil B S, 2007. Alien crustacean decapods from the Aegean coast of Turkey. Aquatic Invasions. 2 (3), 162-168. http://www.aquaticinvasions.ru/2007/AI_2007_2_3_Yokes_etal.pdf DOI:10.3391/ai.2007.2.3.2

Zenetos A, Gofas S, Verlaque M, Cinar ME, Garcia Raso JE, Bianchi CN, Morri C, Azzurro E, Bilecenoglu M, Froglia C, Siokou I, Violanti D, Sfriso A, San Martin G, Giangrande A, Katagan T, Ballesteros E, Ramos-Espla A, Mastrototaro F, Ocana O, Zingone A, Gambi MC, Streftaris N, 2010. 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. In: Mediterranean Marine Science, 11 (2) 381-493. http://www.medit-mar-sc.net/files/201102/23-103521437_MMS_v11n2_ZENETOS.pdf

Links to Websites

Contributors

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

Distribution Maps