Review Paper Diversity, Biology and Exploitation of Brackish Water Crabs in West Africa

Available online at http://www.ifgdg.org

Int. J. Biol. Chem. Sci. 12(5): 2355-2370, October 2018

ISSN 1997-342X (Online), ISSN 1991-8631 (Print)

Review Paper http://ajol.info/index.php/ijbcs http://indexmedicus.afro.who.int

Diversity, biology and exploitation of brackish water crabs in West Africa: A review

Comlan Eugène DESSOUASSI1,2*, Antoine CHIKOU1, Djiman LEDEROUN1, Delphine ADANDEDJAN1, Luc GANGBE2 and Philippe LALEYE1

1Laboratory of Hydrobiology and Aquaculture, Faculty of Agronomic Science, University of Abomey-Calavi. 01 BP 526 Tri Postal Cotonou, Benin. 2 Benin National Institute for Agricultural Research. 04BP626 Cotonou, Benin. *Corresponding author; E-mail: [email protected]; Tel: +229 97580352/+229 94735052

ABSTRACT

Crabs are an important biological component of the West African lagoon fauna and are exploited by fishing communities. This paper presents a synthesis of the current understanding pertaining to the diversity, biology and exploitation of these crabs in order to identify promising research paths for this zoological group. A literature review was done. Thirty-two species belonging to thirteen families and twenty-three genera have been indexed by previous investigators and we plan on making use of this information in defining future research direction. The biology of crabs has been the subject of much research focused on Callinectes amnicola (Rochebrune, 1883) species. C. amnicola is an omnivorous species whose males are known to mature between 63 and 105 mm shell width, while females mature between 83.5 and 116.2 mm shell width. Fertility varies from 0.47 to 4.8 million eggs per female. However, exploitation parameters for this species in various lagoons are not available. A research direction geared towards establishing a new and thorough inventory along with currently nonexistent but relevant exploitation parameters will go a long way in defining sustainable management measures for these crab species. © 2018 International Formulae Group. All rights reserved.

Keywords: Lagoon crabs, diversity, biology and exploitation, West Africa.

INTRODUCTION intensely exploited by a human population of Lagoons and estuaries are considered a variety of backgrounds (Lalèyè et al., 2007). throughout the world as the most productive This exploitation focuses mostly on fish and aquatic ecosystems (Lalèyè et al., 2007). In crustaceans (Penaeid shrimps and Portunid the West African tropical region, a succession crabs). Crabs, on the other hand, are a major of lagoons, lakes and estuaries run along the component of lagoon fauna (Le Loeuf and coast (Hardman-Mountford and McGlade, Intès, 1968; Fischer et al., 1981), represent an 2002; Lalèyè et al., 2007; Carpenter and De important food resource for humans, and are Angelis, 2014). Along the Gulf of Guinea, the subject of a remarkable economic activity between Côte d’Ivoire and Nigeria, these (Thiam and Diallo, 2010). Worldwide, total ecosystems cover about 3,000 Km2, with crab catches were estimated at 1.2 million highly diversified living resources that are tons in 2003 (Thiam and Diallo, 2010).

However, it should be emphasized that since and exploitation in order to identify relevant crabs are coastal creatures that are quite easy research prospects for this zoological group. to catch, this figure is exclusive of individual To attend this objective, publications and small-scale catching; especially in touching on lagoon crabs in West Africa were developing countries where they constitute are identified using Google Scholar, Scopus, and a cheap source of protein (Thiam and Diallo, Agris databases as well as UAC (University of 2010). Compared with fish and shrimps, Abomey-Calavi) and the Benin Ministry of however, scientific investigation on crabs as Agriculture, Livestock, and Fisheries libraries. opposed to other fisheries groups such as fish Publications search was conducted using the and shrimp is lacking (Udoh and Nlewadim, keywords crabe lagunaire, Afrique de l’Ouest 2011; Sara et al., 2010). In fact, of the (French) or lagoon crab, West Africa identified publications covering research on (English) (Assogbadjo et al., 2011; Wiegleb, crabs in West Africa, the first ones date back 2016; Gangbè et al., 2016; Pochet, 2017). to the 1960s, although the last one was Database of crustacean catalogs of the France published in 2017. Of the earlier ones, the National Museum for Natural History, of the very first two are Monod (1956) and Manning US National Museum of Natural History, and & Holtuis (1981) who established the first of the Leiden Natural Museum were also inventory of the entire marine, lagoon, and used. A list of crab species was established land crabs on the West African coast. using species nomenclature as defined by Ng Literature review on the exploitation and et al. (2008) as well as the recent systematic production of Callinectes sp. and Cardisoma revised versions (Shahdadi and Schubart, sp. in West Africa (d’Almeida and Fiogbé, 2017). 2008) indicates an insufficient body of The present synthesis of current knowledge about these two species despite scientific knowledge on brackish water crabs their socio-economic relevance. The authors in West Africa focuses on species encountered have suggested that knowledge be improved in Benin, Ghana, Côte d’Ivoire, Nigeria, and relative to the estimation and exploitation of Togo (Figure 1). This review is made as a stocks as well as to the biological prelude to establishing an inventory and understanding necessary for a successful framing future research aimed at a promising breeding of Callinectes amnicola exploitation of these species in Benin. (Rochebrune, 1883) and Cardisoma armatum (Herklots, 1851). Considering the full breadth LAGOON CRABS DIVERSITY IN THE of diversity in crab species in West African GULF OF GUINEA lagoons (Monod, 1956; Manning and Thirty-two crab species are reported in Holthius, 1981) and in view of the scientific lagoons and associated mangrove habitats knowledge produced on this zoological group from Côte d’Ivoire to Nigeria. These species so far (Akin-Oriola et al., 2005; d’Almeida et are divided into twenty-three genera and al., 2006; Arimoro and Idoro, 2007; thirteen families (Table 1). d’Almeida et al., 2007; d’Almeida et al., According to available literature, the 2008; d’Almeida et al., 2009; George et al., highest species richness of lagoon crabs is 2009; George and Abowei, 2009; d’Almeida observed in Nigeria (twenty-three species) and et al., 2009; d’Almeida et al., 2010; Udoh and the lowest is observed in Togo (two species). Nlewadim, 2011; Omuvwie and Atobatele, Country size as well as the lagoons selected 2013; Olakolu and Fakayode, 2014; Sankare for developing the inventory justifies the et al., 2014 a; Onyekachi and Edah, 2014; observed gap. Regional surveys conducted d’Almeida et al., 2014), it appears necessary during colonial times and around the period of to have a review in order to establish what the the independence movement in West Africa current understanding on this group of have not included all the lagoons in such crustacean is. We review known lagoon crab countries as Benin or Côte d’Ivoire. In Benin, species in the Gulf of Guinea, their biology, no crab species have been reported in the 2356 C. E. DESSOUASSI et al. / Int. J. Biol. Chem. Sci. 12(5): 2355-2370, 2018 coastal lagoon, in the Porto-Novo Lagoon, nor (Sankaré, 2007; George et al., 2009; in Ahémé lake. This situation equally applies Santhanam, 2018). The diet consists of to the Grand Lahou and Aby lagoons in Côte oligochaetes, polychaetes, pelecypod d’Ivoire, as well. Subsequent surveys carried molluscs, crustaceans, and aquatic insects out in Benin, Nigeria, and Côte d’Ivoire (Sankaré, 2007) in Côte d’Ivoire waters. revealed that twelve species were not reported Specimens smaller than 5 cm in size tend to in the first regional surveys, namely: have a scavenger-type diet, while those whose Callinectes sapidus, Callinectes danae, size is between 5 and 10 cm are carnivorous Callinectes marginatus, Geryon maritae, (Sankaré, 2007). In the Lagos lagoon Gecarcinus weileri, Portunus validus, (Nigeria), diet is composed of lobsters, fish, Menippe nodifrons, Pilumnopeus caparti, mollusks, and crabs (Onyekachi and Edah, Ocypoda cursor, Grapsus grapus, 2014). Maintaining the focus on Nigeria, diet Pachygrapsus transversus, Cyclograpsus in the Okpoka River is composed of integrer. Unlike regional surveys, it is not crustaceans, fish, mollusks, annelids, algae, possible to have access to the collections of sand particles, and crab parts (George and national surveys because the authors have not Abowei, 2009). specified the places where reported specimens Reproduction in Portunidae crabs are kept. New surveys on lagoon crabs are occurs through mating. Crabs capable of needed. Possibilities of molecular tests could swimming have a sexual dimorphism that is be envisaged. Indeed, recent taxonomic already detectable in post-larval stages overhauling has affected species of the (Bourgeois-Lebel, 1980; Charles Dominique Perisesarma genus in the West African et Hem, 1981; Sankaré, 2007; Arimoro and carcinological fauna (Shadadi and Schubart, Idoro, 2007; Jivoff et al., 2007). C. amnicola 2017). Assessing biodiversity according to reproduction in Ebrié Lagoon (Côte d’Ivoire) UICN criteria would also be useful in the face is characterized by female migration from of anthropogenic pressures on natural desalinated zones to the meso-euhaline areas resources in general and on fisheries resources near Vridi channel. This migration is followed particularly. Edible species, which are by gonad maturation. Males are much more economically important, are more the focus of sedentary and are found in oligohaline waters, biological and ecological investigations. sometimes in fresh waters (Charles Dominique and Hem, 1981). Studies LAGOON CRABS BIOLOGY IN THE describing differentiation in C. amnicola GULF OF GUINEA reproduction organs, with scales of sexual Available information on the biological maturity, have been carried out (Charles aspects of lagoon crabs covers more than 75% Dominique and Hem, 1981; d’ Almeida et al., of the crab population encountered in 2006; d’ Almeida et al., 2007; Sankaré, 2007; Nigerian lagoons (Qua Iboe, Okpoka, Ojo, d’ Almeida et al., 2008; d’ Almeida et al., and Lagos lagoons), in Côte d’Ivoire lagoons 2009; d’ Almeida et al., 2010; d’ Almeida et (Ebrié, Aby, Grand Lahou and Fresco al., 2014). These sexual maturity scales lagoons), and in Ghanaian waters (Mukwe and display 7 stages. Table 2 shows size at first Sakumo). This information focuses mainly on sexual maturity for the identified species in Callinectes amnicola (Rochebrune, 1883), an some of the lagoons in the study area. The abundant and economically-important species. biological cycle of the species described by Other scarcely-studied species include Lhomme (1994) is presented on Figure 2. Callinectes pallidus (Rochebrune, 1883), Cardisoma armatum (Herklots, 1851) and Callinectes pallidus (Rochebrune, 1883) Guinearma huzardi (Desmarest, 1825). Callinectes pallidus biological cycle Callinectes amnicola (Rochebrune, 1883) occurs in both seawater and lagoon (Lhomme, Callinectes amnicola has an 1994). Indeed, C. pallidus is a coastal marine omnivorous diet that varies with crab size species that penetrates mesohaline brackish 2357 C. E. DESSOUASSI et al. / Int. J. Biol. Chem. Sci. 12(5): 2355-2370, 2018 waters (Monod, 1956; Williams, 1974; live its life on earth. This explanation supports Manning and Holthuis, 1981). Its diet was the occasional arrival of C. armatum into studied in the Ojo Estuary in Badagri, Nigeria. lagoon waters (Akin-Oriola et al., 2005). It, It consists of fish, molluscs, crustaceans, however, has to be noted that the C. armatum macroscopic plants, algae, sand particles, and sexual maturity scale, its breeding season, and unidentified masses (Jimoh et al., 2014; its first maturity size in various environments Santhaman, 2018). Males consume more fish in the sub-region are still unknown and are while females consume more plants. No part of the questions future research should reproduction parameters of this species are address. available. Guinearma huzardi (Desmarest, 1825) Cardisoma armatum Herklots, 1851 The diet of this species consists of Studies on Cardisoma armatum have mangrove leaves, fish, and other crabs focused on its ecoethological parameters, diet, (Lawal-Are and Nwankwo, 2011). The same and reproduction. This species colonizes more authors have observed a sex ratio in favor of clay soils, has a mainly phytophagous diet, males. Fertility ranges from 1.5 to 3.5 million and feeds mainly on Paspalum vaginatum eggs with egg diameter ranging from 0.21 to leaves, palm nuts (Elaeis guineensis), and 0.33 mm. coconut (Cocos nucifera) (Hounga, 1999). It Two species have mainly been has an active phase on the land surface at the focus of biology-related investigations: night and an apparent resting phase in burrows Callinectes amnicola and Cardisoma in the daytime (Mensah and Gbeto, 2001). armatum. Knowledge gained from the biology Akin-Oriola et al. (2005) have investigated the and ecology of Callinectes amnicola is an morphometric, meristic, and growth essential basis for the establishment of parameters in C. armatum compared with sustainable management measures based on those of C. pallidus. Morphometric and biological indicators. Future diet-related meristic parameters and weight-length investigations focusing on daily dietary habits relationship in C. armatum were also and an estimation of dietary requirements will compared with those in the C. amnicola be useful to feeding efficiency within the swimming blue crab in the Nokoué framework of farming this species. As for Lake/Porto-Novo Lagoon complex of Benin reproduction parameters, knowledge of the (Goussanou et al., 2017a). Embryonic size at first maturity as well as the breeding development in C. armatum has also been season and preferred breeding grounds on described (d’Almeida et al., 2014). It appears each lagoon are essential for the sustainable that its embryonic development involves a management of crabs. transformation process of a telolecith egg into Regarding Callinectes pallidus, the larva. Fertilized eggs exude from the cavity of total lack of information on the reproduction gravid or grainy females through gonopores of the species will have to be filled in order to and attach to the bristles of biramous understand its life cycle and to assess its pleopods. These females become ovigerous in exploitation rate in the lagoons of the Gulf of the abdomen where embryogenesis takes Guinea. place and involves the stages of segmentation, As far is Cardisoma armatum is gastrulation, and organogenesis. concerned, existing work makes it possible to Macroscopically, fertilized eggs and germs understand that its life cycle occurs on the shape a cluster whose color changes from earth with an occasional introduction in orange to yellow, to gray, and finally to black. lagoon waters, just as swimming crabs do At the end of this process, the embryos are (Akin-Oriola et al., 2005). The scale of released into the aquatic environment to maturity applicable to the species, its breeding undergo actual larval development which, season, and the size at first maturity in the after metamorphosis, will give a juvenile crab various West African environments are yet to that comes out of the lagoon environment to be established. Available information on its 2358 C. E. DESSOUASSI et al. / Int. J. Biol. Chem. Sci. 12(5): 2355-2370, 2018 biological life does not, as yet, allow making a Beninese lagoons between 2000 and 2008. statement on the extent of its exploitation. This increase is also observed in Ivorian lagoons with the exception of the Aby Lagoon EXPLOITATION OF LAGOON CRABS where catches have decreased (Sankaré et al., IN THE GULF OF GUINEA 2014a). The decrease in catches in the Aby Available knowledge relating to the Lagoon is related to the increase in the exploitation of lagoon crabs covers primarily number of fishermen, fishing gear, the the Callinectes amnicola species in a few catching of females, and the closure of the countries. As well, exploitation statistics do lagoon's natural channel (Sankaré et al., not cover that much of a continuous timespan 2014b). From Côte d’Ivoire to Nigeria, and are derived from short-term exploitation parameters have not been investigations. Compiled data is shown in evaluated. However, this information is Table 3. available on other economically-important In Côte d’Ivoire lagoons, most Portunid crabs such as Portunus pelagicus Callinectes amnicola fishing is done by and Portunus sanguinolentus elsewhere in the gillnets. This gear has a potential danger for world (Muller et al., 2006; Dash et al., 2013; the maintenance of the fertility of the stock, Chutapa et al., 2014). Dash et al. (2013) report since catches are generally made before an exploitation rate of E = 0.61 for Portunus breeding, and fishing pressure seems very sanguinolentus in India between 2009 and high (Charles Dominique and Hem, 1981, 2010. In Thailand, Chutapa et al. (2014) have Lhomme, 1994). On the Ebrié lagoon, Charles reported an exploitation rate of E = 0.71 for Dominique and Hem (1981) have reported the Portunus pelagicus between 2008 and 2009. following select crab fishing gear: gillnets; Analyzes are made by these authors, which crab trap, and multiply-hooked lines. On the gave recommendations for sustainable other hand, the poorly selective gear which management of crab fisheries. The FiSAT II captures almost all size classes include software (Sparre & Venema, 1998) which gillnets, seines, and purse seines. uses the size frequencies to generate the From 2006 to 2009, average catch per population and exploitation parameters is unit effort was 33 kg/landing for the Fresco used. Considering the strong fishing pressures Lagoon, 16.5 to 32.25 kg/landing for the reported on the Callinectes amnicola species Grand-Lahou Lagoon, from 25 to 38.5 in West African lagoon environments (Charles kg/landing for the Ebrié Lagoon and from 5 to Dominique and Hem, 1981; Lhomme, 1994; 88.25 kg/landing for the Aby Lagoon Gnimadi et al., 2008; d’Almeida and Fiogbé, (Sankaré et al., 2014a). 2008; Sankaré, 2007; Sankaré et al., 2014a; In the Lagos Lagoon of Nigeria, Sankaré et al., 2014b), it is highly circular lift net and wire basket traps are recommended to evaluate the exploitation fishing gears inventoried (Babatunde, 2008). parameters of the Callinectes amnicola The catch per unit effort varies from 2.5 to 15 species in the various lagoon ecosystems in crabs for circular lift net (very selective gear) the Golf of Guinea. The socio-economic as opposed to 5.6 to 17 crabs for wire basket importance of crab fisheries also deserves to trap. In Benin, crab fishermen are estimated to be studied. As for the Cardisoma armatum number roughly 2030, including 588 women land crab, apart from catching by traps known (Gnimadi et al., 2008). in West African countries, catch statistics are The socio-economic importance of not available. Callinectes amnicola in West African lagoons (Le Loeuf and Intes, 1968, Gnimadi et al., FARMING LAGOON CRABS IN THE 2008, d’Almeida and Fiogbé, 2008) justifies GULF OF GUINEA the fact that the available exploitation Callinectes amnicola farming trials information is relevant only to this species. have been tested and show that it can be done Crab catches have generally doubled in in drainable ponds (Sohou et al., 2016; 2359 C. E. DESSOUASSI et al. / Int. J. Biol. Chem. Sci. 12(5): 2355-2370, 2018

Goussanou et al., 2017b). Effective command substrate with fresh water) is still unsuccessful of the control of the Scylla serrata brackish (Mensah and Gbeto, 2001; Edea et al., 2015). water crab farming, which belongs to the The artificial reproduction of the species and same family as Callinectes amnicola (Mirera, larval development are described by Cuesta 2014; Davis, 2004), can serve as a basis for and Anger (2005) in an aquarium. A salinity further domestication studies. With regard to of 25‰ allows the best survival of eggs and Cardiosoma armatum, the conditions of its larvae through metamorphosis. Subsequent rearing in pens have also been studied (Edéa trials should take into account this optimum et al., 2015). The rearing in pens (sandy salinity value for crab farming purposes.

Figure 1: Geographical area covered by the review.

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Table 1: List of crab species identified in the lagoons of five countries on the Gulf of Guinea.

Family/Species Country Museums/ References Register Number Dromiidae Sternodromia monodi (Forest & Guinot, Nigeria RMNH.CRUS.D.27151 Manning and Holthuis (1981) 1966) RMNH.CRUS.D.27152 Dorippidae Phyllodorippe armata (Miers, 1881). Nigeria RMNH.CRUS.D.23325 Manning and Holthuis (1981) Menippidae Menippe nodifrons Stimpson, 1859 Côte N/A Sankaré (2005) d’Ivoire Benin N/A Hountogan (2012) Euryplacidae Machaerus oxyacanthus (Monod, 1956) Nigeria RMNH.CRUS.D.44442 Manning and Holthuis (1981) Parthenopidae RMNH.CRUS.D.24025 Manning and Holthuis (1981) Parthenopoides massena (Roux, 1830) Nigeria RMNH.CRUS.D.24026 Pilumnidae Nigeria RMNH.CRUS.D.23664 Manning and Holthuis (1981) Nigeria USMN 170334 https://collection.nmnh.si.edu/ Pilumnopeus africanus (De Man, 1902) Côte USNM Manning and Holthuis (1981) d’Ivoire Benin RMNH.CRUS.D.19944- Manning and Holthuis (1981) RMNH.CRUS.D.19946 Côte N/A Sankaré (2005) Pilumnopeus caparti (Monod, 1956) d’Ivoire Portunidae Ghana RMNH.CRUS.D.58 Manning and Holthuis (1981) Ghana USNM 170116-170118 https://collection.nmnh.si.edu/ Togo USNM 143985 Manning and Holthuis (1981) https://collection.nmnh.si.edu/ Nigeria USNM 120943 Manning and Holthuis (1981) https://collection.nmnh.si.edu/ Callinectes amnicola (Rochebrune, Côte N/A Sankaré (2005) 1883) d’Ivoire Murai et al. (2003); Adandédjan (2012); Benin N/A Hountogan (2012); Dessouassi (2014), Mensah (2014) Manning and Holthuis (1981); Nigeria RMNH.CRUS. D.15533 http://bioportal.naturalis.nl

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http://bioportal.naturalis.nl/ Nigeria RMNH.CRUS.D.15532 (Manning and Holthuis (1981); Amadi(1990); Akin- Callinectes pallidus (Rochebrune, Oriola et al.(2005) 1883) Ghana USNM 14879 Manning and Holthuis (1981) https://collection.nmnh.si.edu/ Côte N/A Sankaré (2005) d’Ivoire Benin N/A Aka et al. (1999) Nigeria N/A Amadi (1990) ; Akin-Oriola Callinectes marginatus (A. Milne- et al. (2005) Edwards, 1861) Côte N/A d’Ivoire Sankaré (2005) Callinectes sapidus Rathbun, 1896 Benin N/A Adandédjan (2012) Callinectes danae Smith, 1869 Benin N/A Adandédjan (2012) Sanquerus validus (Herklots, 1851) Benin N/A Murai et al. (2003) RMNH.CRUS.D.24575- Monod (1956) ; Manning and Portunus inaequalis (Miers, 1881) Nigeria 24577 Holthuis (1981) RMNH.CRUS.D.25287 Panopeidae Nigeria RMNH.CRUS.D.15548 http://bioportal.naturalis.nl/ ; Monod (1956) Panopeus africanus A. Milne-Edwards, Côte 1867 N/A Sankaré (2005) d’Ivoire Benin RMNH.CRUS.D.18735 Manning and Holthuis (1981) Xanthidae Nanocassiope melanodactylus (A. Nigeria USNM Manning and Holthuis (1981) Milne- Edwards, 1867) Gecarcinidae Côte d’Ivoire N/A Monod (1956) Ghana RMNH.CRUS.D.58 Manning and Holthuis (1981). Ghana MNHN 2403 Monod (1956) Cardisoma armatum Herklots, 1851 Togo N/A Monod (1956) Manning and Holthuis (1981) Benin RMNH.CRUS.D.17890 ; Adandédjan (2012) Nigeria RMNH.CRUS.D.15472 Manning and Holthuis (1981) Johngarthia weileri (Slendler, 1912) Nigeria N/A Amadi (1990) ; Akin-Oriola et al. (2005) Grapsidae Ghana RMNH.CRUS.D.68 Manning and Holthuis (1981 Manning and Holthuis (1981); Goniopsis pelii (Herklots, 1851) Nigeria RMNH.CRUS. D.15518 http://bioportal.naturalis.nl Benin N/A Hountogan (2012)

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Côte Monod (1956) ; Manning and N/A d’Ivoire Holthuis (1981) Manning and Holthuis (1981); Nigeria RMNH.CRUS. D.15553 http://bioportal.naturalis.nl Ghana USNM 170340 Manning and Holthuis (1981) Pachygrapsus gracilis (Saussure, 1858) https://collection.nmnh.si.edu/ Benin RMNH.CRUS.D.19935 Manning and Holthuis (1981) Côte Sankaré (2005) N/A d’Ivoire Pachygrapsus transversus (Gibbes, Côte Sankaré (2005) N/A 1850) d’Ivoire Grapsus grapsus (Linneaus, 1758) Côte N/A Sankaré (2005) d’Ivoire Cyclograpus integer H. Milne Edwards, Côte N/A Sankaré (2005) 1837 d’Ivoire Sesarmidae Metagrapsus curvatus (H. Milne RMNH.CRUS.D.31013 Manning and Holthuis (1981); Edwards, 1837) Nigeria RMNH.CRUS.D.30887 Shahdadi and Schubart (2017) RMNH.CRUS. D.15537 http://bioportal.naturalis.nl Côte N/A Sankaré (2005) d’Ivoire Benin N/A Adandédjan (2012) Ghana RMNH.CRUS. D.182 Manning and Holthuis (1981). Nigeria RMNH.CRUS. D.15538 http://bioportal.naturalis.nl Ghana MNHN 2398 Monod (1956) Chiromantes(Sesarma)angolense De Benin Brito Capello, 1864 N/A Hountogan (2012) Côte N/A Monod (1956) d’Ivoire Nigeria RMNH.CRUS.D.30988 Manning and Holthuis (1981) Chiromantes (Sesarma) buettikoferi De Côte RMNH.CRUS.D.19862 Manning and Holthuis (1981) Man, 1883 d’Ivoire RMNH.CRUS. D.150- Ghana Monod (1956) 151 Côte N/A Sankaré (2005) Armases elegans (Herklots, 1851) d’Ivoire Benin N/A Adandédjan (2012) Manning and Holthuis (1981) Nigeria RMNH.CRUS. D.15567 ;http://bioportal.naturalis.nl Manning and Holthuis (1981) Nigeria RMNH.CRUS. D.15536 ;http://bioportal.naturalis.nl Guinearma alberti (Rathbun, 1921) Côte Monod (1956) N/A d’Ivoire Côte Manning and Holthuis (1981) RMNH.CRUS.D.19849 d’Ivoire Guinearma huzardi (Desmarest, 1825) Manning and Holthuis (1981) Benin RMNH.CRUS.D.18729 ; Adandédjan (2012)

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Ghana RMNH.CRUS.D.132 Manning and Holthuis (1981).

Ghana MNHN 2398 Monod (1956) Ghana USMN 170346 https://collection.nmnh.si.edu/ Manning and Holthuis (1981) Nigeria RMNH.CRUS.D.15539 http://bioportal.naturalis.nl/ Nigeria MNHN 2498 Monod (1956) Ocypodidae Ghana RMNH.CRUS.D.27227 Manning and Holthuis (1981) Manning and Holthuis (1981) Benin RMNH.CRUS.D.18736 ; Adandédjan (2012) Ocypode africana De Man, 1881 Côte d’Ivoire N/A Sankaré (2005) Manning and Holthuis (1981); Nigeria RMNH.CRUS. D.15517 http://bioportal.naturalis.nl Ocypode cursor (Linneaus, 1758) Côte Sankaré (2005) d’Ivoire N/A Ghana RMNH Manning and Holthuis (1981) RMNH.CRUSD. Manning and Holthuis (1981 Ghana 262 Afruca tangeri (Eydoux, 1835) Côte Sankaré (2005) N/A d’Ivoire Benin N/A Adandédjan (2012) Ghana RMNH.CRUS.D.15535 Manning and Holthuis (1981 MNHN: French National Museum for Natural History (Paris), USMN: US National Museum of Natural History; RMNH: Leiden Naturalis Museum; NA: museum/register number not available.

Table 2: Size at first sexual maturity in Callinectes amnicola in the lagoons of the Gulf of Guinea.

First maturity size Reproductive Females fertility Area References No (cm) period (million eggs per crab)

Male Female

Côte d’Ivoire: Sankaré March-April 0.5 - 2 Aby-Tendo-Ehy (2007) 1 8 9 and August lagoon complex

Ghana: Mukwe Kwei (1978) 1.902 ± 0.24 to 2.82 ± 2 6.3 8.35 N/A and Sakumo 0.32 Benin : Dessouassi 3 9.16 11.62 N/A N/A Ahémé lake (2014)

Nigeria : Lagos Santhanam 4 10.5 11.0 N/A 0.47 - 4.48 lagoon (2018) N/A: Not available.

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Figure 2: Biological cycle of Callinectes amnicola in Ebrié lagoon, Côte d’Ivoire (Lhomme, 1994).

Table 3 : Catch data of Callinectes amnicola crabs in a few lagoons in the Gulf of Guinea.

Reference Annual Country Water body References years production (tons)

1990 - 2000 2678.605 D/Pêches (2001) Nokoué Lake DPH(2016); DPH(2017a); DPH 2016 828 (2017b); DPH(2017c) 1990 - 2000 342.28 D/Pêches (2001) Porto-Novo Lagoon DPH(2016); DPH(2017a); DPH 2016 74.11 (2017b); DPH(2017c) Nokoué Lake-Porto- Gnimadi et al. (2008) Benin 2008 5846.464 Novo Lagoon 1990 - 2000 541.985 D/Pêches (2001) Ahémé Lake DPH(2016); DPH(2017a); DPH 2016 79.85 (2017b); DPH(2017c) Coastal Lagoon 1990 - 2000 173.6238 D/Pêches (2001) Ahémé Lake-Coastal Gnimadi et al. (2008) 2008 1379.016 Lagoon Aby Lagoon 2001 3747 Sankaré (2007) 2006-2009 2720 Sankaré et al. (2014a) Ebrié Lagoon 1975 190 Charles Dominique and Hem (1981) Côte 1979 825.2 Charles Dominique and Hem (1981) d’Ivoire 2006-2009 2248 Sankaré et al. (2014a) Grand-Lahou Lagoon 2006 - 2009 823 Sankaré et al. (2014a) Fresco Lagoon 2006 - 2009 14.75 Sankaré et al. (2014a)

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Conclusion ACKNOWLEDGEMENTS The knowledge based on lagoon crabs We would like to thank Professeur is variable in Gulf of Guinea countries, Danièle Guinot (MNHN) for Monod (1956) namely from Côte d’Ivoire to Nigeria. West and advices provided; Karen Dorp Van African surveys conducted in the 1960s (RMNH) for assistance in catalogue database provided information on diversity. This search. information has been supplemented in some countries (Côte d’Ivoire, Benin, and Nigeria) REFERENCES by national surveys. Adandédjan D. 2012. Diversité et Thirty-two species belonging to déterminisme des peuplements de thirteen families and twenty genera are known macroinvertébrés benthiques de deux in West African lagoon environments. The lagunes du Sud-Bénin : la Lagune de highest species richness was reported in Porto-Novo et la Lagune Côtière. Thèse Nigeria (23 species) and in Côte d’Ivoire (22 de Doctorat, Université d’Abomey- species). Seventeen species have been Calavi, Abomey-Calavi, p. 239. identified in Benin, 11 in Ghana, and 02 Aka NAD, Allabi ACE, Dreyfuss G, Kinde- species in Togo. Callinectes amnicola Gazard D, Tawo L, Rondelaud D, breeding biology, diet, and growth have been Bouteille B, Avode G, Anagonou SY, studied in Côte d’Ivoire and Nigeria lagoons. Gninafon M, Massougbodji A, Dumas However, the Callinectes amnicola breeding M. 1999. Observations épidémiologiques season, dietary habits, and growth have been sur le premier cas de paragonimose studied in Ivorian and Nigerian lagoons. humaine et les hôtes intermédiaires Breeding areas of this species are not yet potentiels de Paragonimus sp. au Bénin. known in most of West African lagoons. Manuscrit n° 2017. Bull. Soc. Pathol. However, its size at first maturity, breeding Exot., 92:191-194. season and best breeding grounds are not yet Akin-Oriola G, Anetekhai M, Olowonirejuaro known in most of West African lagoons. K. 2005. Morphometric and Meristic The estimation of exploitation Studies in Two Crabs: Cardiosoma parameters of this species which is intensely armatum and Callinectes pallidus. Turk caught in different lagoons seems important J. Fish Aquat. Sc., 5: 85-89. for management measures. Amadi AA. 1990. A comparative ecology of It is also important to initiate research estuaries in Nigeria. Hydrobiologia, on the socio-economic importance of 208(1): 27-38. DOI: 10.1007 Callinectes amnicola caught in lagoons. /BF00008440 Cardisoma armatum species has also been Arimoro FO, Idoro BO. 2007. Ecological investigated in some biological surveys. The Studies and Biology of Callinectes other thirty species are potential research amnicola (Family: Portunidae) in the subjects. Lower of Reaches of Warri River, Delta State, Nigeria. WJZ., 2(2): 57-66. COMPETING INTERESTS Assogbadjo AE, Aihou K, Youssao IAK, The authors declare that they have no Fovet-rabot C, Mensah GA. 2011. competing interests. L’écriture scientifique au Bénin Guide contextualize de formation. Dépôt légal AUTHORS CONTRIBUTIONS n° 5372 du 26 septembre 2011, 3ème CED collected the diversity data and Trimestre 2011. Coco New Tech: wrote the manuscript. AC and LG collected Cotonou. the biology data. DA collected the Babatunde EE. 2008. The fisheries and the exploitation data. DL and PL read and Bionomics of swimming crab, corrected the manuscript. Callinectes amnicola (De Rochebrune, 1883) from a tropical lagoon and its 2366 C. E. DESSOUASSI et al. / Int. J. Biol. Chem. Sci. 12(5): 2355-2370, 2018

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