INTRODUCTION
The infraorder Brachyura is regarded as the most diverse taxon within the Crustacea, with more than 6500 species (de Grave et al., Reference De Grave, Pentcheff, Ahyong, Chan, Crandall, Dworschak, Felder, Feldmann, Fransen, Goulding, Lemaitre, Low, Martin, Ng, Schweitzer, Tan, Tshudy and Wetzer2009) related to 102 families (Ahyong et al., Reference Ahyong, Lowry, Alonso, Bamber, Boxshall, Castro, Gerken, Karaman, Goy, Jones, Meland, Rogers, Svavarsson and Zhang2011).
The brachyuran fauna from the north-eastern Atlantic, and especially from European coasts, is currently one of the best known worldwide. The knowledge on diversity and distribution of decapod fauna in this region was summarized by d'Udekem d'Acoz (Reference d'Udekem d'Acoz1999). Further publications, mostly focused on the Mediterranean Sea, provided new information on bathymetry and/or biogeographic distributions of decapods (Maynou & Cartes, Reference Maynou and Cartes2000; Sánchez-Jerez et al., Reference Sánchez-Jerez, Barberá-Cebrián and Ramos-Esplá2000; Modena et al., Reference Modena, Mori and Vacchi2001; Abelló et al., Reference Abelló, Carbonell and Torres2002; Pipitone & Arculeo, Reference Pipitone and Arculeo2003; Company et al., Reference Company, Maiorano, Tselepides, Politou, Plaity, Rotllant and Sardà2004; Politou et al., Reference Politou, Maiorano, D'Onghia and Mytilineou2005; Ungaro et al., Reference Ungaro, Marano, Ceriola and Martino2005; Ateş et al., Reference Ateş, Katağan and Kocataş2006; Fanelli et al., Reference Fanelli, Colloca and Ardizzone2007; Pipitone & Vaccaro, Reference Pipitone, Vaccaro, Pessani, Tirelli and Froglia2011; El Lakhrach et al., Reference El Lakhrach, Hattour, Jarboui, Elhasni and Ramos-Esplá2012), or information related to new findings or the biology of a particular brachyuran species (Mura & Cau, Reference Mura and Cau2002; Giacobbe & Spano, Reference Giacobbe and Spano2006; Rossetti et al., Reference Rossetti, Sartor, Francesconi, Mori and Belcari2006; Guerao & Abelló, Reference Guerao and Abelló2007; Mavidis et al., Reference Mavidis, Türkay and Koukouras2008; Isajlović et al., Reference Isajlović, Vrgoč and Dulčić2009; Massi et al., Reference Massi, Micalizzi, Giusto and Pipitone2010; Neudecker et al., Reference Neudecker, Schiefenhövel, Kehlert and Becker2011; Capezzuto et al., Reference Capezzuto, Maiorano, Panza, Indennidate, Sion and D'Onghia2012; Porporato et al., Reference Porporato, de Domenico, Mangano, Rinelli and Spanò2012).
However, where West Africa is concerned, the only comprehensive monographs on brachyuran crabs are those by Monod (Reference Monod1956) and Manning & Holthuis (Reference Manning and Holthuis1981). In addition, some local studies on crustacean or decapod fauna were also undertaken along the West African coast, the most representative being those by Barnard (Reference Barnard1950, Reference Barnard1955) in South Africa; Macpherson (Reference Macpherson1983, Reference Macpherson1991) in Namibia; Henriksen (Reference Henriksen2009) in the Gulf of Guinea; Muñoz et al. (Reference Muñoz, García-Isarch, Sobrino, Burgos, Funny and González-Porto2012) in Guinea-Bissau; Fransen (Reference Fransen1991) in both the Canarian-Cape Verdean Region and the Banc d'Arguin (Mauritania); Anadón (Reference Anadon1981) in northern Mauritania and south-western Sahara; González Pérez (Reference González Pérez1995) in the Canary Islands and García-Raso (Reference García Raso1996) in Ibero-Moroccan waters. All these works include citations for brachyurans and collectively have improved the knowledge of this group. However, no other references focus particularly on Mauritania other than a succinct list of brachyurans (Monod, Reference Monod1933) and some scant records (Monod, Reference Monod1956; Anadón, Reference Anadon1981; Manning & Holthuis, Reference Manning and Holthuis1981; Fransen, Reference Fransen1991).
From 2007 to 2010, the ECOAFRIK project, led by the Spanish Institute of Oceanography (IEO), in collaboration with the University of Vigo (Spain), carried out four annual multidisciplinary surveys in Mauritanian waters (MAURIT surveys). This work presents the taxonomic results for the deep brachyuran fauna and is the first contribution associated exclusively with this group in waters off Mauritania. We also provide new data about the geographic distribution, bathymetric range and spawning periods for some of the species studied.
MATERIALS AND METHODS
The MAURIT surveys were conducted annually from 2007 to 2010 onboard the Spanish RV ‘Vizconde de Eza’ along the Mauritanian coast (16°05′49″N 20°48′13″W, see Map 1) in November–December.
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Map 1. Sampling area showing the location of the MAURIT stations with presence of brachyurans.
A total of 316 stations were sampled at depths between 79 and 1867 m by means of two different bottom trawl gears. A total of 291 stations were sampled with a commercial trawl (Lofoten type, coded MU), following a stratified random sampling procedure. Another 25 stations were sampled with a 3.5 m beam trawl (coded MUBV) along five transects perpendicular to the coastline in five bathymetric strata (150, 300, 500, 1000, 1500 m), distributed along the Mauritanian coast (see Map 1). Station data, including coordinates, depth, date and collected species, are summarized in Table 1.
Table 1. Data of the MAURIT stations: date, coordinates, bathymetric range and captured species.
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Samples were sorted and identified onboard to the lowest taxonomic level. For each species all the specimens were counted, weighed and pictured. A representative collection of each station and of each species was preserved in 70% ethanol and stored for further studies in the laboratory.
The classification adopted in the present work follows Ahyong et al. (Reference Ahyong, Lowry, Alonso, Bamber, Boxshall, Castro, Gerken, Karaman, Goy, Jones, Meland, Rogers, Svavarsson and Zhang2011) to the family level, and genera assignments follow de Grave et al. (Reference De Grave, Pentcheff, Ahyong, Chan, Crandall, Dworschak, Felder, Feldmann, Fransen, Goulding, Lemaitre, Low, Martin, Ng, Schweitzer, Tan, Tshudy and Wetzer2009). Subsections, superfamilies, families and species are usually listed in alphabetical order.
For each species we provide some relevant literature references, including those with the descriptions of the species that best fit with our specimens. We also provide the material examined (with additional material for some species), mention of the station code, the depth range and the number of specimens captured, in parentheses. As a measurement of specimens, we used the carapace length (CL), measured along the dorsal midline, from the base of rostrum to the posterior margin of the carapace. This measurement was obtained for males, females and ovigerous females, or combinations of all three, when appropriate. We also summarize the biological features and geographic distributions (mainly those published in the last 30 years), including the new data reported with this work, along with some remarks when required. In addition, we include pictures, mostly taken onboard when captured, of each species described.
In the laboratory, pictures were taken with a motorized Nikon SMZ25 stereomicroscope, using NIS-Elements Microscope Imaging Software, with an Extended Depth of Focus (EDF) patch.
The specimens examined for this work are largely deposited in the collections of the University of Vigo (Spain) (Marine Zoology Laboratory) and at the Oceanographic Centres of the Spanish Institute of Oceanography (IEO) in Cádiz (Collection of Decapod and Stomatopod Crustaceans, CCDE-IEOCD) and Málaga (The Marine Fauna Collection, CFM-IEOMA).
Abbreviations used are: ICMD: Biological Reference Collections samples code of the ICM-CSIC (Instituto de Ciencias del Mar-Consejo Superior de Investigaciones Científicas), Barcelona, Spain; USNM: United States Natural Museum, Smithsonian National Museum of Natural History, Washington, USA; IEO-CD: Instituto Español de Oceanografía, Centro Oceanográfico de Cádiz, Spain.
RESULTS
SYSTEMATICS
Order decapoda Latreille, Reference Latreille1803
Infraorder brachyura Latreille, Reference Latreille1802
Section dromiacea de Haan, Reference Haan and Siebold1833
Superfamily homoloidea de Haan, Reference Haan and Siebold1839
Family homolidae de Haan, Reference Haan and Siebold1839
Genus Homola Leach, Reference Leach1815
Homola barbata (Fabricius, Reference Fabricius1793)
(Figure 1)
Cancer barbatus Fabricius, Reference Fabricius1793: 460.
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Fig. 1. Homola barbata (Fabricius, 1793), ovigerous female CL: 21.11 mm dorsal view, MU281.
Homola barbata: Guinot & Richer de Forges, Reference Guinot and Macpherson1995: 323 figs. 1A, 7A, B, 8a, b, e, 9a, 13a (references).
MATERIAL EXAMINED
MU87, 271–305 m, (2); MU88, 94–120 m, (1); MU119, 82–80 m, (1); MU120, 109–105 m, (1); MU122, 82–80 m, (1); MU128, 218–404 m, (1); MU131, 102–104 m, (1); MU158, 80–98 m, (1); MU162, 148–149 m, (1); MU184, 213–202 m, (1); MU186, 174 m, (1); MU200, 352–334 m, (7); MU204, 155–145 m, (1); MU223, 116–117 m, (2); MU281, 100–106 m, (1); MUBV02, 318–330 m, (1); MUBV21, 107–109 m, (1).
Males: 12.78–14.51 mm, females: 12.77–23.30 mm, ovigerous females: 15.53–21.11 mm.
IDENTIFICATION
Our specimens agree well with the description provided by Manning & Holthuis (Reference Manning and Holthuis1981: 25) and González-Gurriarán & Méndez (Reference González-Gurriarán, Méndez, Castro-Sada and Coruña1986: 59).
BIOLOGY
A photophilic and sciaphilic species (Pipitone & Vaccaro, Reference Pipitone, Vaccaro, Pessani, Tirelli and Froglia2011) previously cited from bottoms of mud, mud with rocks, muddy sand, muddy shells, seagrass, rocks and sometimes in caves (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999; Pipitone & Arculeo, Reference Pipitone and Arculeo2003; Pipitone & Vaccaro, Reference Pipitone, Vaccaro, Pessani, Tirelli and Froglia2011). Bathymetric range between 2 and 637 m (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999). Often holding algae or small pieces of sponges over the carapace with the fifth pair of pereiopods, and sometimes reported below the tentacles of the anemone Telmatactis cricoides (Duchassaing, 1850) (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999).
Ovigerous females have been recorded in February (García Raso, Reference García Raso1984) and from April to September (Zariquiey Álvarez, Reference Zariquiey Álvarez1968; Manning & Holthuis, Reference Manning and Holthuis1981).
Our specimens were collected on coarse sand, coarse muddy sand, sandy mud with shell debris and muddy sand bottoms, at depths between 80–98 and 334–352 m. Ovigerous females were captured in November and December.
GEOGRAPHIC DISTRIBUTION
Eastern Atlantic, from the Bay of Biscay to Angola, including the Azores, Madeira, Cape Verde, Desertas, Canary and Gulf of Guinea Islands; the Walter Shoals (E South Africa) (Guinot & Richer de Forges, Reference Guinot and Macpherson1995); Central and Western Mediterranean Basin (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999).
Later records for this species (Monteiro et al., Reference Monteiro, Araújo, Erzini and Castro2001; Pipitone & Arculeo, Reference Pipitone and Arculeo2003; Ungaro et al., Reference Ungaro, Marano, Ceriola and Martino2005; Henriksen, Reference Henriksen2009; Pipitone & Vaccaro, Reference Pipitone, Vaccaro, Pessani, Tirelli and Froglia2011; Muñoz et al., Reference Muñoz, García-Isarch, Sobrino, Burgos, Funny and González-Porto2012) fit well within this geographic distribution.
REMARKS
The ovigerous females recorded for the first time in November and December indicate that this species spawns all year round except in winter.
Paromola Wood-Mason & Alcock, Reference Wood-Mason and Alcock1891
Paromola cuvieri (Risso, Reference Risso1816)
(Figure 2A, B)
Dorippe cuvieri Risso, Reference Risso1816: 35
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Fig. 2. Paromola cuvieri (Risso, 1816), adult: (A) dorsal view, MUBV03; (B) frontal view, MU140.
Paromola cuvieri: Guinot & Richer de Forges, Reference Guinot and Macpherson1995: 362, figures 2, 21a, b (references).
MATERIAL EXAMINED
MU14, 502–511 m, (1); MU18, 519–402 m, (4); MU23, 532–415 m, (17); MU37, 403–442 m, (2) MU121, 274–400 m, (1); MU140, 376–377 m, (1); MU150, 292–341 m, (2) MU157, 278–454 m, (1); MU175, 618–850 m, (1); MU188, 627 m, (1); MU263, 615–624 m, (2) MUBV03, 528–538 m, (1); MUBV22, 300 m, (1).
Male: 95.20 mm, female: 76.20 mm, megalopa: 7.78 mm.
IDENTIFICATION
Our specimens agree well with the descriptions and figures provided by Capart (Reference Capart1951: 25) and Zariquiey Álvarez (Reference Zariquiey Álvarez1968: 301).
BIOLOGY
Deep-water species taken on bottoms with mud or sandy mud, over a cold-water coral community (Pipitone & Arculeo, Reference Pipitone and Arculeo2003; Isajlović et al., Reference Isajlović, Vrgoč and Dulčić2009; Capezzuto et al., Reference Capezzuto, Maiorano, Panza, Indennidate, Sion and D'Onghia2012). The isolated record from the Menez Gwen hydrothermal vent must be considered with caution until confirmed (Martin & Haney, Reference Martin and Haney2005). This species is often reported holding a sponge over the carapace using the fifth pereiopods (Capezzuto et al., Reference Capezzuto, Maiorano, Panza, Indennidate, Sion and D'Onghia2012) and is also found with cirripeds, gorgonians and sea anemones on carapace and legs (González, Reference González Pérez1995; Guinot & Richer de Forges, Reference Guinot and Macpherson1995). Bathymetric range usually between 80 and 300 m (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999), although the species was once reported at 10 m (Manning & Holthuis, Reference Manning and Holthuis1981) and up to 1165 m depth (Cartes et al., Reference Cartes, Maynou, Moranta, Massuti, Lloris and Morales-Nin2004). Ovigerous females have been recorded in March, from May to July, October and November (Zariquiey Álvarez, Reference Zariquiey Álvarez1968; Manning & Holthuis, Reference Manning and Holthuis1981).
Our specimens were collected in muddy sand and sandy mud bottoms, in depths between 274–400 and 618–850 m. No ovigerous females were caught.
GEOGRAPHIC DISTRIBUTION
North and East Atlantic, from southern Iceland, the Hebrides and southern Scandinavia, south to north-western South Africa (Tripp Seamount), including the Azores, Madeira, Canary and Cape Verde Islands; Mediterranean Sea (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999; Martin & Haney, Reference Martin and Haney2005).
Further additional records (Macpherson, Reference Macpherson1991; Maynou & Cartes, Reference Maynou and Cartes2000; Abelló et al., Reference Abelló, Carbonell and Torres2002; Biagi et al., Reference Biagi, Sartor, Ardizzone, Belcari, Belluscio and Serena2002; Pipitone & Arculeo, Reference Pipitone and Arculeo2003; Sardà et al., Reference Sardà, Company and Castellón2003; Cartes et al., Reference Cartes, Maynou, Moranta, Massuti, Lloris and Morales-Nin2004; Company et al., Reference Company, Maiorano, Tselepides, Politou, Plaity, Rotllant and Sardà2004; Politou et al., Reference Politou, Maiorano, D'Onghia and Mytilineou2005; Ungaro et al., Reference Ungaro, Marano, Ceriola and Martino2005; Fanelli et al., Reference Fanelli, Colloca and Ardizzone2007; Isajlović et al., Reference Isajlović, Vrgoč and Dulčić2009; Capezzuto et al., Reference Capezzuto, Maiorano, Panza, Indennidate, Sion and D'Onghia2012; Muñoz et al., Reference Muñoz, García-Isarch, Sobrino, Burgos, Funny and González-Porto2012) fall within this geographic range.
Section cyclodorippoida Ahyong et al., Reference Ahyong, Lai, Sharkey, Colgan and Ng2007
Superfamily cyclodorippoidea Ortmann, Reference Ortmann1892
Family cymonomidae Bouvier, Reference Bouvier1897
Genus Cymonomus A. Milne-Edwards, Reference Milne-Edwards1880
Cymonomus granulatus (Norman, in Thomson, Reference Thomson1873)
(Figure 3)
Ethusa granulata Norman in Thomson, Reference Thomson1873: 176.
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Fig. 3. Cymonomus granulatus (Norman, in Thomson, 1873), adult dorsal view, MUBV10.
Cymonomus granulatus: Zariquiey Álvarez, Reference Zariquiey Álvarez1968: 813 (references).
MATERIAL EXAMINED
MUBV10, 332–344 m, (35).
Males: 4.64–5.23 mm, female: 3.52 mm, ovigerous females: 3.26–3.73 mm.
IDENTIFICATION
Our specimens agree with those described in Milne-Edwards & Bouvier (Reference Milne-Edwards, Bouvier and Milne-Edwards1900: 34) (see Remarks).
BIOLOGY
Mainly found on shell debris and muddy bottoms (Mura & Cau, Reference Mura and Cau2002), between 155 m (García Raso, Reference García Raso1996) and 2425 m (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999). Ovigerous females reported from May to July (García Raso, Reference García Raso1996; Mura & Cau, Reference Mura and Cau2002).
Our specimens were collected at 332–344 m and ovigerous females were captured in November.
GEOGRAPHIC DISTRIBUTION
Eastern Atlantic from SW Scotland to Mauritania; West and Central Mediterranean Sea (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999). In the Mediterranean Sea, this species has been reported in the Alborán Sea (Abelló et al., Reference Abelló, Carbonell and Torres2002) and in the Sardinian Channel (Mura & Cau, Reference Mura and Cau2002). In the NE Atlantic, Cartes et al. (Reference Cartes, Serrano, Velasco, Parra and Sánchez2007) reported the species on Le Danois Bank (Cantabrian Sea).
REMARKS
In our specimens, the mesial margin of the ocular peduncles are more tuberculate than spinulose as described by Milne-Edwards & Bouvier (Reference Milne-Edwards, Bouvier and Milne-Edwards1900: 34). The rest of the characters agree well with the referred description and, consequently, our specimens remain within this species.
This is the first time that ovigerous females are reported at the end of the year, which suggests a biannual spawning strategy for this species.
Section eubrachyura de Saint Laurent, Reference Saint Laurent1980
Subsection heterotremata Guinot, Reference Guinot1977
Superfamily calappoidea de Haan, 1833
Family calappidae de Haan, 1833
Genus Acanthocarpus Stimpson, Reference Stimpson1871
Acanthocarpus brevispinis Monod, Reference Monod1946
(Figure 4)
Acanthocarpus bispinosus Milne-Edwards, Reference Milne-Edwards1880 var. brevispinis, Monod, Reference Monod1946: 7, figures 1–4, pl. figures 1–2.
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Fig. 4. Acanthocarpus brevispinis Monod, 1946, adult dorsal view, MU51.
Acanthocarpus brevispinis: Manning & Holthuis, Reference Manning and Holthuis1981: 50 (references).
MATERIAL EXAMINED
MU51, 468–466 m, (1); MU99, 569–598 m, (1); MU121, 274–400 m, (1); MU134, 311–436 m, (1); MU143, 322 m, (4); MUBV02, 318–330 m, (1); MUBV13, 493–517 m, (3); MUBV14, 300–281 m, (9); MUBV19, 306 m, (12); MUBV22, 300 m, (7).
Males: 15.56–55.29 mm; female: 49.85 mm; ovigerous female: 55.29 mm.
IDENTIFICATION
Our specimens agree well with the descriptions given by Capart (Reference Capart1951: 36; as Acanthocarpus africanus) and Manning & Holthuis (Reference Manning and Holthuis1981: 50).
BIOLOGY
Previously reported on sandy and shelly mud bottoms (Manning & Holthuis, Reference Manning and Holthuis1981), at depths from 100 m (Manning & Holthuis, Reference Manning and Holthuis1981) to 517 m (Muñoz et al., Reference Muñoz, García-Isarch, Sobrino, Burgos, Funny and González-Porto2012). Ovigerous females have been recorded in March and October (Manning & Holthuis, Reference Manning and Holthuis1981).
Our specimens were collected from 274–400 to 569–598 m on sandy mud bottoms; ovigerous females were captured in December.
GEOGRAPHIC DISTRIBUTION
West Africa from Cape Juby (S Morocco) to Namibia (Manning & Holthuis, Reference Manning and Holthuis1981; Macpherson, Reference Macpherson1983, Reference Macpherson1991). Further records for this species were reported by Henriksen (Reference Henriksen2009) from Nigeria and Gabon, and by Muñoz et al. (Reference Muñoz, García-Isarch, Sobrino, Burgos, Funny and González-Porto2012) from Guinea-Bissau.
REMARKS
Material from the station MU99 at 569–598 m is the deepest known record for this species; the finding of ovigerous females in December confirms a biannual spawning strategy.
Genus Calappa Weber, Reference Weber1795
Calappa pelii Herklots, Reference Herklots1851
(Figure 5)
Calappa Pelii Herklots, Reference Herklots1851: 12.
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Fig. 5. Calappa pelii Herklots, 1851, female CL: 33.94 mm dorsal view, MU158.
Calappa pelii: Manning & Holthuis, Reference Manning and Holthuis1981: 52 (references).
MATERIAL EXAMINED
MU158, 80–98 m, (1); MUBV01, 112 m, (4); MUBV15, 148–135 m, (1); MUBV21, 107–109 m, (7).
Males: 21.11–31.91 mm; females: 30.14–40.79 mm; ovigerous female: 58.87 mm.
IDENTIFICATION
Our specimens agree well with the descriptions given by Capart (Reference Capart1951: 39, as Calappa peli) and Manning & Holthuis (Reference Manning and Holthuis1981: 52).
BIOLOGY
Bottoms of mud and broken shells, usually at depths between 8–20 and 400 m. Ovigerous females reported in March, May, August, October and December (Manning & Holthuis, Reference Manning and Holthuis1981).
Our specimens were collected in depths between 80–98 and 135–148 m on sandy bottoms, and ovigerous females were captured in November.
GEOGRAPHIC DISTRIBUTION
West Africa from Western Sahara to Namibia, including Principe Island, and Central Mediterranean Sea (Macpherson, Reference Macpherson1991; d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999). Recent records for this species are from Guinea-Bissau (Muñoz et al., Reference Muñoz, García-Isarch, Sobrino, Burgos, Funny and González-Porto2012) and from the Gulf of Guinea (Henriksen, Reference Henriksen2009).
Superfamily cancroidea Latreille, Reference Latreille1802
Family atelecyclidae Ortmann, Reference Ortmann1893
Genus Atelecyclus Leach, Reference Leach and Brewster1814
Atelecyclus rotundatus (Olivi, Reference Olivi1792)
(Figure 6)
Cancer rotundatus Olivi, Reference Olivi1792: 47, pl. 2 figure 2.
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Fig. 6. Atelecyclus rotundatus (Olivi, 1792), female CL: 27.64 mm dorsal view, MUBV10.
Atelecyclus rotundatus: Manning & Holthuis, Reference Manning and Holthuis1981: 68 (references); González Gurriarán & Méndez, Reference González-Gurriarán, Méndez, Castro-Sada and Coruña1986: 113, figure 38, photo 26.
MATERIAL EXAMINED
MUBV10, 332–344 m, (1).
Female: 27.64 mm.
IDENTIFICATION
Our specimen agrees well with the specifications provided by Forest (Reference Forest1957: 469) and the descriptions in González Gurriarán & Méndez (Reference González-Gurriarán, Méndez, Castro-Sada and Coruña1986: 113).
BIOLOGY
Recorded from sandy bottoms, often with gravel and small stones as well as muddy sand, shelly sand, gravel with shell debris and mud; also recorded on rocks, coralligenous substrates and in Posidonia meadows (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999; Pipitone & Arculeo, Reference Pipitone and Arculeo2003; Ateş et al., Reference Ateş, Katağan and Kocataş2006; Guillén et al. Reference Guillén, Gras, Soler and Triviño2011). Bathymetric distribution from intertidal to 795 m, usually between 9 and 300 m (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999). Ovigerous females were recorded in January and December (García Raso, Reference García Raso1984).
Only one female was collected in the MAURIT surveys between 332–344 m.
GEOGRAPHIC DISTRIBUTION
Eastern Atlantic from SW Faroe Islands to South Africa (beyond Cape of Good Hope, Barnard, Reference Barnard1950), including the Canary, Cape Verde and Ascension Islands; Mediterranean Sea (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999).
Later records (Abelló et al., Reference Abelló, Carbonell and Torres2002; Pipitone & Arculeo, Reference Pipitone and Arculeo2003; Ateş et al., Reference Ateş, Katağan and Kocataş2006; Serrano et al., Reference Serrano, Sánchez and García-Castrillo2006, Reference Serrano, Sánchez, Punzón, Velasco and Olaso2011; García-Muñoz et al., Reference García-Muñoz, Manjón-Cabeza and García-Raso2008; Sánchez et al., Reference Sánchez, Serrano, Parra, Ballesteros and Cartes2008; Muñoz et al., Reference Muñoz, García-Isarch, Sobrino, Burgos, Funny and González-Porto2012; Ellis et al., Reference Ellis, Martinez, Burt and Scott2013) fit well within this geographic distribution.
Superfamily dorippoidea MacLeay, 1838
Family dorippidae MacLeay, 1838
Genus Medorippe Manning & Holthuis, Reference Manning and Holthuis1981
Medorippe lanata (Linnaeus, Reference Linnaeus1767)
(Figure 7)
Cancer lanatus Linnaeus, Reference Linnaeus1767: 1044
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Fig. 7. Medorippe lanata (Linnaeus, 1767), ovigerous female CL: 23.68 mm dorsal view, MUBV01.
.
Medorippe lanata: Manning & Holthuis, Reference Manning and Holthuis1981: 31, figures 4a–h (references).
MATERIAL EXAMINED
MU210, 86–90 m, (1); MUBV01, 112 m, (2); MUBV21, 107–109 m, (19).
Males: 6.88–20.84 mm, female: 11.27 mm, ovigerous female: 23.68 mm.
IDENTIFICATION
Our specimens agree well with the description and figures in Capart (Reference Capart1951: 30; as Dorippe lanata) and in Manning & Holthuis (Reference Manning and Holthuis1981: 31).
BIOLOGY
Mostly recorded on soft heterogeneous substrates (mud, sandy mud, muddy sand and sand) at depths varying from 9 to 769 m (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999; El Lakhrach et al., Reference El Lakhrach, Hattour, Jarboui, Elhasni and Ramos-Esplá2012). On Mediterranean soft bottoms, this species has been associated with demersal assemblages currently exploited by trawling (Rosseti et al., Reference Rossetti, Sartor, Francesconi, Mori and Belcari2006). Ovigerous females have been reported from March to November (Zariquiey Álvarez, Reference Zariquiey Álvarez1968: 313 as Dorippe lanata; Manning & Holthuis, Reference Manning and Holthuis1981: 32; Modena et al., Reference Modena, Mori and Vacchi2001; Rosseti et al., Reference Rossetti, Sartor, Francesconi, Mori and Belcari2006).
Our specimens were collected from 86–90 to 112 m, on sand with biogenic debris bottoms. Ovigerous females were captured in November.
GEOGRAPHIC DISTRIBUTION
East Atlantic, from Portugal to South Africa (up to Natal) and Mozambique, including the Canary Islands and Mediterranean Sea (Barnard, Reference Barnard1950, Reference Barnard1955; Manning & Holthuis, Reference Manning and Holthuis1981; d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999).
Further records (Modena et al., Reference Modena, Mori and Vacchi2001; Abelló et al., Reference Abelló, Carbonell and Torres2002; Biagi et al., Reference Biagi, Sartor, Ardizzone, Belcari, Belluscio and Serena2002; Pipitone & Arculeo, Reference Pipitone and Arculeo2003; Ungaro et al., Reference Ungaro, Marano, Ceriola and Martino2005; Rosseti et al., Reference Rossetti, Sartor, Francesconi, Mori and Belcari2006; Fanelli et al., Reference Fanelli, Colloca and Ardizzone2007; Henriksen, Reference Henriksen2009; El Lakhrach et al., Reference El Lakhrach, Hattour, Jarboui, Elhasni and Ramos-Esplá2012; Muñoz et al., Reference Muñoz, García-Isarch, Sobrino, Burgos, Funny and González-Porto2012) fit well within its geographic distribution.
REMARKS
Manning & Holthuis (Reference Manning and Holthuis1981) erected the genera Medorippe and Phyllodorippe in order to accommodate the Atlantic species of dorippids previously included in the genus Dorippe. Medorippe can be differentiated from Phyllodorippe by the male gonopod morphology (short, stubby, straight, without distal appendages and lobulated at the outer margin of base vs long, slender, S-shaped, with two short distal appendages and lacking a lobe in the proximal part) and by the presence of a row of spines on the dorsal margin of pereiopods 2 and 3.
Family ethusidae Guinot, Reference Guinot1977
Genus Ethusa Roux, Reference Roux1830
Ethusa rosacea A. Milne-Edwards & Bouvier, Reference Milne-Edwards and Bouvier1897
(Figure 8)
Ethusa rosacea A. Milne-Edwards & Bouvier, Reference Milne-Edwards and Bouvier1897: 298; Monod, Reference Monod1956: 88 (references); Manning & Holthuis, Reference Manning and Holthuis1981: 38 (references).
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Fig. 8. Ethusa rosacea A. Milne Edwards & Bouvier, 1897, ovigerous female CL: 11.16 mm dorsal view, MUBV17.
MATERIAL EXAMINED
MUBV17, 1022–1026 m, (1).
Ovigerous female: 11.16 mm.
IDENTIFICATION
Our specimen agrees well with the original description and the figures provided later by Milne-Edwards & Bouvier (Reference Milne-Edwards, Bouvier and Milne-Edwards1900: pls III figure 5, X figures 5–8) and by Capart (Reference Capart1951: figure 5).
BIOLOGY
Bottoms of sand, muddy sand and sandy mud (Manning & Holthuis, Reference Manning and Holthuis1981; d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999), with a bathymetric range of 84 m (Henriksen, Reference Henriksen2009) to 1113 m (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999). Ovigerous females have been recorded in March, April and June (Manning & Holthuis, Reference Manning and Holthuis1981).
Only one ovigerous female was collected in December between 1022–1026 m.
GEOGRAPHIC DISTRIBUTION
The species is recorded in the Eastern Atlantic from scattered localities between Mauritania and Angola, including the Canary and Cape Verde Islands (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999). Henriksen (Reference Henriksen2009) reported this species from the Gulf of Guinea.
REMARKS
The poor development of the outer frontal teeth, the sparse setation of the carapace and the transversal broadness of the walking legs dactylus ensure the identification of our specimen as E. rosacea.
Ovigerous females are recorded for the first time at the end of the year, suggesting a biannual spawn strategy.
Ethusa rugulosa A. Milne-Edwards & Bouvier, Reference Milne-Edwards and Bouvier1897
(Figure 9)
Ethusa rugulosa A. Milne-Edwards & Bouvier, Reference Milne-Edwards and Bouvier1897: 297; Manning & Holthuis, Reference Manning and Holthuis1981: 39 (references).
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Fig. 9. Ethusa rugulosa A. Milne-Edwards & Bouvier, 1897, adult male dorsal view, MUBV21.
MATERIAL EXAMINED
MUBV01, 112 m, (1); MUBV02, 318–330 m, (1); MUBV10, 332–344 m, (26); MUBV20, 155 m, (1); MUBV21, 107–109 m, (1).
Males: 8.83–16.60 mm, females: 11.64–14.93 mm, ovigerous females: 11.09–12.64 mm.
IDENTIFICATION
Our specimen agrees well with the description and figures of the type provided by Milne-Edwards & Bouvier (Reference Milne-Edwards, Bouvier and Milne-Edwards1900: 24).
BIOLOGY
This species has been recorded from different soft bottoms, including sand and shells, muddy sand, shelly mud and also hard substrates (Manning & Holthuis, Reference Manning and Holthuis1981: 39; Henriksen, Reference Henriksen2009: 154). Bathymetric range between 55–60 and 275 m (Manning & Holthuis, Reference Manning and Holthuis1981: 39). Ovigerous females have been recorded in July (Manning & Holthuis, Reference Manning and Holthuis1981: 39).
Our specimens were collected from 107–109 to 332–344 m with ovigerous females in November.
GEOGRAPHIC DISTRIBUTION
Eastern Atlantic, recorded from scattered localities: Mauritania (present work), the Cape Verde Islands, Senegal, Sierra Leona, Liberia and Angola (Manning & Holthuis, Reference Manning and Holthuis1981) (see Remarks). This species was also reported from the Gulf of Guinea by Henriksen (Reference Henriksen2009).
REMARKS
Our finding is the first record of this species in Mauritania, extending the northern distribution up to Cape Blanc. Also, the bathymetric range is extended from 275 m reported in the literature to 332–344 m in this work.
The presence of ovigerous females for the first time at the end of the year suggests a biannual spawn.
Superfamily goneplacoidea MacLeay, 1838
Family goneplacidae MacLeay, 1838
Genus Goneplax Leach, Reference Leach and Brewster1814
Goneplax barnardi (Capart, Reference Capart1951)
(Figure 10)
Carcinoplax barnardi Capart, Reference Capart1951: 170, figures 65 a, b.
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Fig. 10. Goneplax barnardi (Capart, 1951), male CL: 20.01 mm dorsal view, MUBV09.
Goneplax barnardi: Castro, Reference Castro2007: 689, figure 27b (references).
MATERIAL EXAMINED
MU44, 606–596 m, (1); MU54, 414–451 m, (2); MU57, 430–406 m, (5); MU123, 278–362 m, (1); MU152, 381–316 m, (1); MU214, 650–588 m, (1); MU216, 422 m, (5); MUBV03, 528–538 m, (6); MUBV09, 549–555 m, (1); MUBV13, 493–517 m, (14); MUBV14, 300–281 m, (7); MUBV18, 559–574 m, (11); MUBV19, 306 m, (1); MUBV25, 499–520 m, (2).
Males: 9.73–20.01 mm, females: 12.52–15.61 mm, ovigerous females: 11.18–15.51 mm.
IDENTIFICATION
Our specimens agree well with the description of the type and with the detailed figures provided in Monod (Reference Monod1956: 351).
BIOLOGY
Previously reported from mud and sandy mud bottoms, from 200 to 590 m (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999: 241). Ovigerous females have been recorded in March and May (Manning & Holthuis, Reference Manning and Holthuis1981: 160).
Our specimens were collected between 278–362 and 588–650 m on sand and sandy mud bottoms. Ovigerous females were captured in December.
GEOGRAPHIC DISTRIBUTION
East Atlantic, from Western Sahara to Angola, including the Cape Verde Islands (Castro, Reference Castro2007) (see Remarks). Muñoz et al. (Reference Muñoz, García-Isarch, Sobrino, Burgos, Funny and González-Porto2012) later reported this species from Guinea-Bissau.
REMARKS
Castro (Reference Castro2007) removed this species from the former Carcinoplax to Goneplax because of its longer eye peduncles, dorsal margin of the ambulatory legs (P2–P5) meri armed with an acute distal tooth (although this character is also present in the Western Pacific Carcinoplax spinosissima Rathbun, 1914), and slender dactyli carinated on both sides. Obviously, Castro (Reference Castro2007: 690) meant Eastern Atlantic when he wrote ‘Western Atlantic along the west coast of Africa’.
This record constitutes the first for the species in Mauritanian waters and it is the first time that ovigerous females are reported at the end of the year. The known spawn period (March, May and December) suggests that the species is a biannual spawner.
Goneplax rhomboides (Linnaeus, Reference Linnaeus1758)
(Figure 11)
Cancer rhomboides Linnaeus, Reference Linnaeus1758: 626.
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Fig. 11. Goneplax rhomboides (Linnaeus, 1758), male CL: 13.06 mm dorsal view, MU143.
Goneplax rhomboides: Castro, Reference Castro2007: 687, figure 27A (references).
MATERIAL EXAMINED
MU143, 322 m, (1); MU207, 88–117 m, (1); MU227, 183–181 m, (3); MUBV14, 300–281 m, (2).
Males: 9.98–13.06 mm, ovigerous female: 10.05 mm.
IDENTIFICATION
Our specimens agree well with the descriptions and figures in Monod (Reference Monod1956: 354; as Goneplax angulata) and in Zariquiey Álvarez (Reference Zariquiey Álvarez1968: 414).
BIOLOGY
Burrowing species (Atkinson, Reference Atkinson1974; Neudecker et al., Reference Neudecker, Schiefenhövel, Kehlert and Becker2011), usually found in mud, sandy mud, muddy sand and sand bottoms; also on rocks and shells (Manning & Holthuis, Reference Manning and Holthuis1981; d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999; Pipitone & Arculeo, Reference Pipitone and Arculeo2003; Ateş et al., Reference Ateş, Katağan and Kocataş2006; Trenkel et al., Reference Trenkel, Le Loc'h and Rochet2007; Mutlu & Ergev, Reference Mutlu and Ergev2008; Fanelli et al., Reference Fanelli, Cartes, Badalamenti, Rumolo and Sprovieri2009; Metin et al., Reference Metin, Gökçe, Aydin and Bayramiç2009; Guillén et al., Reference Guillén, Gras, Soler and Triviño2011; Neudecker et al., Reference Neudecker, Schiefenhövel, Kehlert and Becker2011; El Lakhrach et al., Reference El Lakhrach, Hattour, Jarboui, Elhasni and Ramos-Esplá2012; Ellis et al., Reference Ellis, Martinez, Burt and Scott2013). Bathymetric range from intertidal zone (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999) to 600–999 m (Company et al., Reference Company, Maiorano, Tselepides, Politou, Plaity, Rotllant and Sardà2004; Cartes et al., Reference Cartes, Maynou, Fanelli, Papiol and Lloris2009). Ovigerous females were reported from January to March, from May to August, and in November and December (Zariquiey Álvarez, Reference Zariquiey Álvarez1968; Manning & Holthuis, Reference Manning and Holthuis1981; García Raso, Reference García Raso1984, Reference García Raso1996).
Our specimens were collected between 88–117 and 322 m on sandy mud and muddy sand bottoms; and the ovigerous female was captured in December.
GEOGRAPHIC DISTRIBUTION
East Atlantic, from the coast of SE Norway and SW Sweden, the North Sea, SW United Kingdom and Ireland, the continental European coast southwards to Western Africa, at least up to Senegal (see Remarks), including Madeira and the Canary Islands, and in the Mediterranean Sea (Castro, Reference Castro2007; Guinot & Castro, Reference Guinot and Castro2007; Berggren, Reference Berggren2008; Berggren & Stefan, Reference Berggren and Stefan2010).
Further records for this species from European waters and in the Mediterranean Sea, not included in the above-mentioned references, are Maynou & Cartes (Reference Maynou and Cartes2000), Monteiro et al. (Reference Monteiro, Araújo, Erzini and Castro2001), Biagi et al. (Reference Biagi, Sartor, Ardizzone, Belcari, Belluscio and Serena2002), Pipitone & Arculeo (Reference Pipitone and Arculeo2003), Company et al. (Reference Company, Maiorano, Tselepides, Politou, Plaity, Rotllant and Sardà2004), Ungaro et al. (Reference Ungaro, Marano, Ceriola and Martino2005); Vincent (Reference Vincent2005), Ateş et al. (Reference Ateş, Katağan and Kocataş2006), Sartor et al. (Reference Sartor, Francesconi, Rossetti and de Ranieri2006), Serrano et al. (Reference Serrano, Sánchez and García-Castrillo2006), Fanelli et al. (Reference Fanelli, Colloca and Ardizzone2007), Trenkel et al. (Reference Trenkel, Le Loc'h and Rochet2007), Mutlu & Ergev (Reference Mutlu and Ergev2008), Cartes et al. (Reference Cartes, Maynou, Fanelli, Papiol and Lloris2009), Fanelli et al. (Reference Fanelli, Cartes, Badalamenti, Rumolo and Sprovieri2009), Metin et al. (Reference Metin, Gökçe, Aydin and Bayramiç2009), Neudecker et al. (Reference Neudecker, Schiefenhövel, Kehlert and Becker2011), Serrano et al. (Reference Serrano, Sánchez, Punzón, Velasco and Olaso2011), El Lakhrach et al. (Reference El Lakhrach, Hattour, Jarboui, Elhasni and Ramos-Esplá2012) and Ellis et al. (Reference Ellis, Martinez, Burt and Scott2013).
REMARKS
Goneplax rhomboides has long been described as a species with a wide geographic distribution in East Atlantic waters, from the North Atlantic to South Africa (Barnard, Reference Barnard1950; Manning & Holthuis, Reference Manning and Holthuis1981; d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999). However, Guinot & Castro (Reference Guinot and Castro2007) described the new species, Goneplax clevai, from the South Atlantic and western limits of the Indo-West Pacific region, including in their new species some specimens reported from the Ivory Coast to South Africa and previously identified as G. rhomboides. Guinot & Castro (Reference Guinot and Castro2007: 25) accept as valid the material from Senegal mentioned by Monod (Reference Monod1956) for G. rhomboides. Consequently, the southern distribution limit of G. rhomboides will remain unclear southwards from Senegal until all the recorded specimens from the West African coasts can be properly checked.
Superfamily leucosioidea Samouelle, Reference Samouelle1819
Family leucosiidae Samouelle, Reference Samouelle1819
Genus Ebalia Leach, Reference Leach1817
Ebalia nux Norman in A. Milne-Edwards, Reference Milne-Edwards1883
(Figure 12)
Ebalia nux A. Milne-Edwards, Reference Milne-Edwards1883: pl. 5; Holthuis & Manning, Reference Manning and Holthuis1981: 61 (references)
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Fig. 12. Ebalia nux Norman in A. Milne-Edwards, 1883, male CL: 7.02 mm dorsal view, MUBV10.
.
MATERIAL EXAMINED
MU87, 271–305 m, (1); MUBV10, 332–344 m, (3).
Males: 7.02–7.20 mm.
IDENTIFICATION
Our specimens agree well with those figured by Milne-Edwards & Bouvier (Reference Milne-Edwards, Bouvier and Milne-Edwards1900: plates III, figure 7, XIII: figures 1–5), and with the descriptions provided by Zariquiey Álvarez (Reference Zariquiey Álvarez1968: 328), and by González Gurriarán & Méndez (Reference González-Gurriarán, Méndez, Castro-Sada and Coruña1986: 71).
BIOLOGY
This species has been recorded on shell, sand, sand and rocks, sand with calcareous algae, mud and sandy mud bottoms (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999; Ateş et al., Reference Ateş, Katağan and Kocataş2006); it has been also reported living on the sea pen, Pteroeides spinosum (Ellis, 1764), in the Mediterranean Sea (Porporato et al., Reference Porporato, de Domenico, Mangano, Rinelli and Spanò2012). Depths records vary between 80 and 2983 m but usually from 150 to 500 m (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999). Ovigerous females have been recorded in March (Manning & Holthuis, Reference Manning and Holthuis1981) and May–June (García Raso, Reference García Raso1996).
Our specimens, all males, were collected between 271–305 and 332–344 m.
GEOGRAPHIC DISTRIBUTION
Eastern Atlantic, from the Shetland Islands and Norway to Mauritania, including the Azores, Canary and Cape Verde Islands; and Mediterranean Sea (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999).
Later records for this species (Abelló et al., Reference Abelló, Carbonell and Torres2002; Ateş et al., Reference Ateş, Katağan and Kocataş2006; Porporato et al., Reference Porporato, de Domenico, Mangano, Rinelli and Spanò2012) are all from the Mediterranean, with the exception of a report by Cartes et al. (Reference Cartes, Serrano, Velasco, Parra and Sánchez2007) from Le Danois Bank (Cantabrian Sea).
Genus Pseudomyra Capart, Reference Capart1951
Pseudomyra mbizi Capart, Reference Capart1951
(Figure 13)
Pseudomyra mbizi Capart, Reference Capart1951: 49, figure 14, pl. II figure 24; Manning & Holthuis, Reference Manning and Holthuis1981: 66 (references).
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Fig. 13. Pseudomyra mbizi Capart, 1951, adult dorsal view, MUBV01.
MATERIAL EXAMINED
MUBV01, 112 m, (9); MUBV15, 148–135 m, (93).
Males: 16.54–20.31 mm, female: 15.96–17.45 mm, ovigerous females: 15.45–17.63 mm.
IDENTIFICATION
Our specimens agree well with the original description and figures.
BIOLOGY
The species has been collected on a variety of bottoms, such as mud, sandy mud with shells, mud with foraminifera, muddy sand, broken shells, and also on coral and rocks. Bathymetric range between 12–15 and 300 m, although usually recorded from 50 to 100 m (Manning & Holthuis, Reference Manning and Holthuis1981). Ovigerous females have been recorded from March to August, October and December (Manning & Holthuis, Reference Manning and Holthuis1981).
Our specimens were captured in depths between 112 and 135–148 m. The ovigerous females were caught in December.
GEOGRAPHIC DISTRIBUTION
West Africa from Mauritania (present work) to Angola (Manning & Holthuis, Reference Manning and Holthuis1981) (see Remarks). Henriksen (Reference Henriksen2009) reports this species from the Gulf of Guinea.
REMARKS
This is the first record of this species from Mauritanian waters, which extends its north distribution up to the Banc d'Arguin.
Superfamily majoidea Samouelle, Reference Samouelle1819
Family epialtidae MacLeay, 1838
Genus Pisa Leach, Reference Leach and Brewster1814
Pisa armata (Latreille, Reference Latreille1803)
(Figure 14)
Maja armata Latreille, Reference Latreille1803: 98
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Fig. 14. Pisa armata (Latreille, 1803), female CL: 22.12 mm dorsal view, MU139.
.
Pisa armata: Manning & Holthuis, Reference Manning and Holthuis1981: 318 (references).
MATERIAL EXAMINED
MU120, 109–105 m, (1); MU131, 102–104 m, (2); MU136, 103–112 m, (1); MU139, 96–97 m, (2); MU147, 134–139 m, (1); MU154, 92–102 m, (2); MU211, 92–109 m, (1); MU261, 111–146 m, (1); MUBV15, 148–135 m (1).
Males: 17.74–22.14 mm, females: 21.08–22.12 mm, ovigerous females: 22.27–24.67 mm.
IDENTIFICATION
Our specimens agree well with the descriptions and figures provided by Capart (Reference Capart1951: 90) and by González-Gurriarán & Méndez (Reference González-Gurriarán, Méndez, Castro-Sada and Coruña1986: 183).
BIOLOGY
This species has been recorded from rocky bottoms with gorgonians and corals, and also from sandy mud, mud, sand and shells bottoms (Manning & Holthuis, Reference Manning and Holthuis1981; d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999; García Raso & Manjón-Cabeza, Reference García Raso and Manjón-Cabeza2002; Pipitone & Arculeo, Reference Pipitone and Arculeo2003; Ateş et al., Reference Ateş, Katağan and Kocataş2006). Pipitone & Arculeo (Reference Pipitone and Arculeo2003) recorded the species on bottoms with Posidonia meadows, although they showed no correlation with the meadow structure (Sánchez-Jerez et al., Reference Sánchez-Jerez, Barberá-Cebrián and Ramos-Esplá2000). Bathymetric range extends from 3–10 m (Pipitone & Arculeo, Reference Pipitone and Arculeo2003) to 162 m (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999). Ovigerous females were recorded from March to May, from July to August and from November to December (Zariquiey Álvarez, Reference Zariquiey Álvarez1968; Manning & Holthuis, Reference Manning and Holthuis1981; García Raso, Reference García Raso1984).
Our specimens, often with sponges, ascidians and bryozoans over the carapace, were captured between 96–97 and 135–148 m on shell debris, sand and sandy mud bottoms. Ovigerous females were collected in December.
GEOGRAPHIC DISTRIBUTION
East Atlantic from Southern North Sea and SW British Isles to Angola, including the Azores, Canary and Cape Verde Islands; and Mediterranean Basin (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999).
Later records for this species (Sánchez-Jerez et al., Reference Sánchez-Jerez, Barberá-Cebrián and Ramos-Esplá2000; Abelló et al., Reference Abelló, Carbonell and Torres2002; García Raso & Manjón-Cabeza, Reference García Raso and Manjón-Cabeza2002; Pipitone & Arculeo, Reference Pipitone and Arculeo2003; Ateş et al., Reference Ateş, Katağan and Kocataş2006) fit well within its distribution.
REMARKS
Ng et al. (Reference Ng, Guinot and Davie2008) reviewed the nomenclatural and taxonomic problems within brachyurans, and rearranged the former family Pisididae as a subfamily into Epialtidae.
Family inachidae MacLeay, 1838
Inachus Weber, Reference Weber1795
Inachus aguiarii de Brito Capello, Reference de Brito Capello1876
(Figure 15A, B)
Inachus aguiarii de Brito Capello, Reference de Brito Capello1876: 265, pl. 2 figures 1–3
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Fig. 15. Inachus aguiarii Brito Capello, 1876, ovigerous female CL: 11.36 mm, MU277: (A) general dorsal view; (B) ventral view, detail of sternal callosities.
Inachus aguiarii: Manning & Holthuis, Reference Manning and Holthuis1981: 283 (references).
MATERIAL EXAMINED
MU120, 109–105 m, (4); MU129, 95–93 m, (1); MU131, 102–104 m, (1); MU226, 109–107 m, (1); MU277, 112–110 m, (1).
Males: 9.35–11.61 mm, ovigerous females: 10.35–11.36 mm.
IDENTIFICATION
Our specimens agree with those described in Zariquiey Álvarez (Reference Zariquiey Álvarez1948: 301 as Inachus thoracicus ssp. aguiarii).
BIOLOGY
Previously recorded from hard bottoms with sponges and kelp; also from soft bottoms of sandy mud often mixed with shell fragments, sand or sand with calcareous algae. Recorded at depths from 20 to 100 m (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999). Ovigerous females recorded in February, March and August (Zariquiey Álvarez, Reference Zariquiey Álvarez1968; García Raso, Reference García Raso1989).
Our specimens were collected from 93–95 to 110–112 m on sand, coarse sand, gravel and sandy mud bottoms. Ovigerous females were captured in November and December.
GEOGRAPHIC DISTRIBUTION
East Atlantic from Portugal to Guinea including the Desertas and Canary Islands (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999); in the Mediterranean Sea, the species was reported only in the Alboran and Aegean Seas (Guerao & Abelló, Reference Guerao and Abelló2007).
REMARKS
Bathymetric range is slightly increasing from that reported in the literature.
Our records of ovigerous females in November and December suggests that this species spawns all year round.
Inachus angolensis Capart, Reference Capart1951
(Figure 16)
Inachus angolensis Capart, Reference Capart1951: 72, figure 72, pl. I figure 7, pl. II figure 10; Manning & Holthuis, Reference Manning and Holthuis1981: 283 (references)
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Fig. 16. Inachus angolensis Capart, 1951, adult dorsal view, MUBV01.
.
MATERIAL EXAMINED
MU88, 94–120 m, (1); MU130, 252–362 m, (1); MU133, 87 m, (1); MU137, 81–84 m, (3); MU161, 89–92 m, (1); MU168, 87–92 m, (1); MU174, 85–84 m, (4); MU205, 89–93 m, (1); MU210, 86–90 m, (9); MU212, 163–200 m, (1); MU251, 107 m, (1); MU260, 101–120 m, (1); MU291, 106–137 m, (1); MUBV01, 112 m, (12); MUBV02, 318–330 m, (1); MUBV14, 300–281 m, (1); MUBV15, 148–135 m, (11); MUBV21, 107–109 m, (6).
Males: 10.04–15.11 mm, females: 8.36–13.67 mm, ovigerous females: 11.56–15.35 mm.
IDENTIFICATION
Our specimens agree well with the original description and also with the notes and the figures provided by Monod (Reference Monod1956: 524, figures 712 and 713).
BIOLOGY
This species has been recorded from mud, sandy mud and sandy bottoms (Manning & Holthuis, Reference Manning and Holthuis1981), at depths from 0–81 m (Henriksen, Reference Henriksen2009) up to at least 350 m (Manning & Holthuis, Reference Manning and Holthuis1981). Ovigerous females were recorded from February to June, September and October (Manning & Holthuis, Reference Manning and Holthuis1981).
Our specimens were captured between 81–84 and 252–362 m on coarse sand, sandy mud with gravel or shell fragments. Ovigerous females were collected in November and December.
GEOGRAPHIC DISTRIBUTION
West Africa from Western Sahara (Manning & Holthuis, Reference Manning and Holthuis1981) to Namibia (Macpherson, Reference Macpherson1991). Henriksen (Reference Henriksen2009) reports this species from the Gulf of Guinea.
REMARKS
Our finding of ovigerous females in November and December suggests that this species spawns throughout the year.
Inachus grallator Manning & Holthuis, Reference Manning and Holthuis1981
(Figure 17A, B)
Inachus grallator Manning & Holthuis, Reference Manning and Holthuis1981: 287, figures 73, 74.
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Fig. 17. Inachus grallator Manning & Holthuis, 1981, male CL: 6.91 mm, MUBV10, in ethanol: (A) general dorsal view; (B) dorsal carapace detail. Scale bars: (A) 1 cm; (B) 1 mm.
MATERIAL EXAMINED
MUBV10, 332–344 m, (1); MUBV14, 300–281 m, (1).
Males: 6.91 and 7.72 mm.
IDENTIFICATION
Our specimens agree well with the original description.
BIOLOGY
Reported from green mud, brown sandy mud, muddy sand and green muddy sand, at depths between 100 and 250–300 m (Manning & Holthuis, Reference Manning and Holthuis1981) and down to 325 m (Fransen, Reference Fransen1991). Ovigerous females have been recorded in January and November (Manning & Holthuis, Reference Manning and Holthuis1981).
Only two males were collected during the MAURIT surveys, at 281–300 and 332–344 m depth.
GEOGRAPHIC DISTRIBUTION
West Africa in Mauritania (Fransen, Reference Fransen1991 in part; present work) and from Nigeria to Namibia (Manning & Holthuis, Reference Manning and Holthuis1981; Macpherson, Reference Macpherson1991) (see Remarks).
REMARKS
Although the possibility of finding this species up to the Western Sahara (Manning & Holthuis, Reference Manning and Holthuis1981) or the Canary Islands (González Pérez & Quiles Lucas, Reference González Pérez, Quiles Lucas, Moro, Martín, Garrido and Izquierdo2003) has been mentioned in the literature, to date no records further north than Nigeria – excepting those in Fransen (Reference Fransen1991) – have been reported from Mauritania and the Canary Islands. The fact that Manning & Holthuis (Reference Manning and Holthuis1981) described I. grallator as a ‘deep-water species, known to occur at depths between 100 and 250–300 m’ suggests that further revision is required for Fransen's (Reference Fransen1991) records for this species at less than 100 m depth. Our record confirms the presence of this species in Mauritanian waters and slightly increases the previously reported bathymetric range.
This species closely resembles another Inachus species, I. dorsettensis. However, after analysing some specimens of I. dorsettensis from Galicia (NW Spain), we conclude that the slenderness of the pereiopods and the shape of the dactyli of the fifth pereiopods ensure the identification of the Mauritanian material as I. grallator.
Inachus leptochirus Leach, Reference Leach1817
(Figure 18A–C)
Inachus leptochirus Leach, Reference Leach1817, in Leach Reference Leach1815–1875: 1 p, pl. XXII.b figures 1, 2, 3; Manning & Holthuis, Reference Manning and Holthuis1981: 291 (references)
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Fig. 18. Inachus leptochirus Leach, 1817, MUBV10: (A) adult general view; (B) male CL: 12.42 mm, in alcohol, carapace dorsal view; (C) same ventral view. Scale bars: B, C, 1 mm.
.
MATERIAL EXAMINED
MU200, 352–334 m, (4); MU204, 155–145 m, (1); MUBV10, 332–344 m, (29).
Males: 9.28–19.21 mm, female: 13.93–19.33 mm, ovigerous females: 10.56–18.37 mm.
IDENTIFICATION
Our specimens agree well with notes and figures in Bouvier (Reference Bouvier1940: 356) and Zariquiey Álvarez (Reference Zariquiey Álvarez1968: 472).
BIOLOGY
Bottoms of gravel, sand, mud and maerl beds, at depths between 27 m (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999) and 500–650 m (Serrano et al., Reference Serrano, Sánchez, Punzón, Velasco and Olaso2011). Ovigerous females have been observed in February, May and November (Zariquiey Álvarez, Reference Zariquiey Álvarez1968).
Our specimens were captured between 145–155 and 334–352 m on coarse sand and muddy sand bottoms, mixed with shell debris. Ovigerous females were collected in November.
GEOGRAPHIC DISTRIBUTION
East Atlantic, from the Faroe Islands to Mauritania, including the Azores; and Mediterranean Sea (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999). This species was recently reported in the N Iberian Peninsula by Serrano et al. (Reference Serrano, Sánchez, Punzón, Velasco and Olaso2011).
Inachus nanus Manning & Holthuis, Reference Manning and Holthuis1981
(Figure 19)
Inachus nanus Manning & Holthuis, Reference Manning and Holthuis1981: 291, figure 75a–e.
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Fig. 19. Inachus nanus Manning & Holthuis, 1981, ovigerous female CL: 6.72 mm, MU166, in ethanol, dorsal view. Scale bar 1 mm.
MATERIAL EXAMINED
MU166, 87–85 m, (2); MU207, 88–117 m, (1); MU212, 163–200 m, (1); MU261, 111–146 m, (1).
Male: 9.50 mm, ovigerous female: 6.72 mm.
IDENTIFICATION
Our specimens agree well with the original description.
BIOLOGY
Previously reported on broken shells, bryozoans, gravel, mud with foraminifera and muddy sand bottoms. Depth records between 29 and 118 m (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999: 198). Ovigerous females have been collected in February, May to July, September and October (Manning & Holthuis, Reference Manning and Holthuis1981).
Our specimens were found between 85–87 and 163–200 m on muddy sand, coarse muddy sand and shell debris bottoms. Ovigerous females were captured in December.
GEOGRAPHIC DISTRIBUTION
West Africa from Mauritania to Cameroon, including the Canary Islands (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999).
REMARKS
This record slightly increases the bathymetric range of this species down to 163–200 m; the presence of ovigerous females in December suggests that this species spawns throughout the year.
Genus Macropodia Leach, Reference Leach and Brewster1814
Macropodia gilsoni (Capart, Reference Capart1951)
(Figure 20A, B)
Achaeopsis gilsoni Capart Reference Capart1951: 65, pl. I figure 4, 10, pl. II figure 3.
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Fig. 20. Macropodia gilsoni (Capart, 1951), ovigerous female CL: 14.35 mm, MUBV21: (A) general dorsal view; (B) dorsal carapace detail.
Macropodia gilsoni: Manning & Holthuis, Reference Manning and Holthuis1981: 297 (references).
MATERIAL EXAMINED
MU119, 82–80 m, (2); MU120, 109–105 m, (3); MU131, 102–104 m, (1); MU138, 123–130 m, (2); MU139, 96–97 m, (1); MU149, 93–146 m, (1); MU151, 110–134 m, (1); MU156, 107–102 m, (1); MU161, 89–92 m, (2) MU166, 87–85 m, (2) MU168, 87–92 m, (2) MU170, 102–92 m, (13); MU210, 86–90 m, (4); MU217, 111–113 m, (1); MU223, 116–117 m, (2); MU233, 165–189 m, (1); MU235, 123 m, (1); MUBV15, 148–135 m, (10); MUBV21, 107–109 m, (1).
Males: 4.71–9.36 mm, females: 3.21–5.09 mm, ovigerous females: 11.05–4.35 mm.
IDENTIFICATION
Our specimens agree well with the description of the type and also with the figures provided by Monod (Reference Monod1956: 555).
BIOLOGY
Species recorded on various kinds of muddy bottoms, including bottoms with shells and bryozoans or foliate foraminifera (Manning & Holthuis, Reference Manning and Holthuis1981), at depths from 37 m (Holthuis & Manning, Reference Manning and Holthuis1981) up to 264 m (Henriksen, Reference Henriksen2009, see Remarks). Ovigerous females recorded throughout the year, except in July (Manning & Holthuis, Reference Manning and Holthuis1981: 297).
Our specimens were captured between 80–82 and 165–189 m on sandy mud, sand and muddy sand bottoms usually mixed with shell debris. Some specimens carried the hydrozoan Obelia bidentata Clark, 1875, on the carapace and legs. Ovigerous females were collected in November and December.
GEOGRAPHIC DISTRIBUTION
West Africa from Mauritania (present work) to Angola (Manning & Holthuis, Reference Manning and Holthuis1981) (see Remarks). Henriksen (Reference Henriksen2009) reported this species off Nigeria, Cameroon and Congo.
REMARKS
Our record is the first report of this species in Mauritania, extending its geographic distribution northwards from Senegal to the Banc d'Arguin.
The record for this species in Henriksen (Reference Henriksen2009) from Cameroon at station 938, Long.: 9.150 Lat.: 3.900, at 0 m depth, must be considered as an error (0 m, null sampling?).
Macropodia hesperiae Manning & Holthuis, Reference Manning and Holthuis1981
(Figure 21)
Macropodia hesperiae Manning & Holthuis, Reference Manning and Holthuis1981: 298, figure 77a–e.
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Fig. 21. Macropodia hesperiae Manning & Holthuis, 1981, male CL: 6.48 mm, MU119, in ethanol, dorsal view. Scale bar 1 mm.
MATERIAL EXAMINED
MU119, 82–80 m, (1); MU129, 95–93 m, (1); MU154, 92–102 m, (2).
Male: 6.48 mm, female: 6.52 mm, ovigerous female: 5.82 mm.
IDENTIFICATION
Our specimens agree well with the original description.
BIOLOGY
Previously recorded on bottoms of mud, sand and compacted sand, at depths varying from 46–49 to 82–97 m. Ovigerous females have been reported in May (Manning & Holthuis, Reference Manning and Holthuis1981).
Our specimens were captured between 80–82 and 92–102 m on compact coarse sand bottoms.
Some specimens carried the hydrozoan species Nemertesia sp. and some colonies of the branched bryozoan species Synnotum aegyptiacum (Audouin, 1826) on their carapaces. The only ovigerous female was collected in December.
GEOGRAPHIC DISTRIBUTION
West Africa from Mauritania (present work) to Nigeria (Manning & Holthuis, Reference Manning and Holthuis1981).
REMARKS
González Pérez (Reference González Pérez1995) mentioned one specimen from the Canary Islands identified as Macropodia aff. hesperiae and collected at 821 m. However, since all the previous records of this species, including those in this study, were found in depths between 46–49 and 92–102 m, we don't consider as valid the record from Canary Islands until further confirmation of the species identification.
Our finding extends the distribution area of this species northwards, from Senegal up to Cape Timiris.
The record of ovigerous females in December indicates that this species spawns at least biannually.
Macropodia longipes (A. Milne-Edwards & Bouvier, Reference Milne-Edwards and Bouvier1899)
(Figure 22)
Stenorhynchus longipes A. Milne-Edwards & Bouvier, Reference Milne-Edwards and Bouvier1899: 48
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Fig. 22. Macropodia longipes (A. Milne-Edwards and Bouvier, 1899), male CL: 7.45 mm MU277, dorsal view. Scale bar: 1 mm.
Macropodia longipes: Manning & Holthuis, Reference Manning and Holthuis1981: 300 (references).
MATERIAL EXAMINED
MU277, 112–110 m, (1).
Male: 7.45 mm.
IDENTIFICATION
Our specimen agrees well with descriptions provided in Forest & Zariquiey Álvarez (Reference Forest and Zariquiey Álvarez1964: 226) and González-Gurriarán & Méndez (Reference González-Gurriarán, Méndez, Castro-Sada and Coruña1986: 169).
BIOLOGY
Species reported in the literature from bottoms of sand and broken shells (Anadon, Reference Anadon1981), sand, silt, gravel and coarse sands (Serrano et al., Reference Serrano, Sánchez, Punzón, Velasco and Olaso2011), and in seagrasses (Ateş et al., Reference Ateş, Katağan and Kocataş2006). Depth records from 9 m (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999) to 1249 m (Cartes et al., Reference Cartes, Maynou, Moranta, Massuti, Lloris and Morales-Nin2004). Ovigerous females recorded in February, March, August and September (Zariquiey Álvarez, Reference Zariquiey Álvarez1968; Pipitone & Tumbiolo, Reference Pipitone and Tumbiolo1993: 362).
The only specimen examined here was a male, collected between 110 and 112 m on a muddy sand bottom.
GEOGRAPHIC DISTRIBUTION
East Atlantic, from the Gulf of Gascogne to Mauritania, including the Cape Verde Islands; and Mediterranean Sea (Forest, Reference Forest1978 as Macropodia tenuirostris longipes).
Further records for this species all refer to the Mediterranean Sea (Modena et al., Reference Modena, Mori and Vacchi2001; Abelló et al., Reference Abelló, Carbonell and Torres2002; Biagi et al., Reference Biagi, Sartor, Ardizzone, Belcari, Belluscio and Serena2002; Cartes et al., Reference Cartes, Maynou, Moranta, Massuti, Lloris and Morales-Nin2004; Colloca et al., Reference Colloca, Carpentieri, Balestri and Ardizzone2004; Politou et al., Reference Politou, Maiorano, D'Onghia and Mytilineou2005; Ungaro et al., Reference Ungaro, Marano, Ceriola and Martino2005; Ateş et al., Reference Ateş, Katağan and Kocataş2006; Fanelli et al., Reference Fanelli, Colloca and Ardizzone2007; García-Muñoz et al., Reference García-Muñoz, Manjón-Cabeza and García-Raso2008; Serrano et al., Reference Serrano, Sánchez, Punzón, Velasco and Olaso2011).
REMARKS
Since Forest (Reference Forest1978) declared Macropodia tenuirostris longipes as the deepest form of M. tenuirostris tenuirostris, some authors subsequently synonymized both subspecies under the name Macropodia tenuirostris (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999; Pipitone & Arculeo, Reference Pipitone and Arculeo2003). However, we follow Ng et al. (Reference Ng, Guinot and Davie2008), who kept them as two different species, only including here references that refer to M. longipes as a separate species or subspecies.
Although the rostrum of our specimen is a little damaged, the slightly curved (not arcuated) P4, P5 dactylus, the basal article of antenna with strong spines ventrally, the presence of a nuchal spine and the absence of spines on the supraorbital margin ensure its identification as M. longipes.
Macropodia macrocheles (A. Milne-Edwards & Bouvier, Reference Milne-Edwards and Bouvier1898)
(Figure 23A–C)
Stenorhynchus macrocheles A. Milne-Edwards & Bouvier, Reference Milne-Edwards and Bouvier1898: 153
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Fig. 23. Macropodia macrocheles (A. Milne-Edwards & Bouvier, 1898): (A) male CL: 8.86 mm dorsal view, MU184; (B) ovigerous female CL: 5.17 mm, MU141, in ethanol, dorsal view; (C) same, epistome detail. Scale bars B, C: 1 mm.
Macropodia macrocheles: Manning & Holthuis, Reference Manning and Holthuis1981: 301 (references)
MATERIAL EXAMINED
MU100, 236–238 m, (3); MU105, 343–346 m, (1); MU141, 280–277 m, (11); MU149, 93–146 m, (1); MU155, 210–257 m, (3); MU157, 278–454 m, (3); MU159, 224–229 m, (7); MU173, 314–540 m, (1); MU179, 303–304 m, (1); MU184, 213–202 m, (2); MU233, 165–189 m, (1); MU280, 230–239 m, (2); MUBV22, 300 m, (1).
Males: 4.41–9.75 mm, ovigerous females: 5.17–6.45 mm.
IDENTIFICATION
Our specimens agree well with the descriptions and figures in Milne-Edwards & Bouvier (Reference Milne-Edwards, Bouvier and Milne-Edwards1900: 159) and Capart (Reference Capart1951: 77).
BIOLOGY
Collected on mud, sand and muddy sand bottoms, at depths between 96 and 300 m (Manning & Holthuis, Reference Manning and Holthuis1981). Ovigerous females have been recorded in January, February, May, June and December (Manning & Holthuis, Reference Manning and Holthuis1981).
Our specimens were captured in depths between 93–143 and 314–540 m on muddy sand and sandy mud bottoms. Some specimens carried hydroid colonies of Clytia gracilis (Sars, 1850), Clytia paulensis (Vanhöffen, 1910), Antennella secundaria (Gmelin, 1791) and a branched unidentified bryozoan species of the genus Bugula Oken, 1815, on their carapaces. Ovigerous females were collected in December.
GEOGRAPHIC DISTRIBUTION
West Africa, from Mauritania (Manning & Holthuis, Reference Manning and Holthuis1981) to Namibia (Macpherson, Reference Macpherson1991).
REMARKS
Our findings slightly extend the bathymetric range previously reported in the literature.
Family majidae Samouelle, Reference Samouelle1819
Genus Eurynome Leach, Reference Leach and Brewster1814
Eurynome aspera (Pennant, Reference Pennant1777)
(Figure 24)
Cancer Asper Pennant, Reference Pennant1777: 7, pl IX.A, figure 20
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Fig. 24. Eurynome aspera (Pennant, 1777), male CL: 10.35 mm dorsal view, MU209.
.
Eurynome aspera: Griffin, Reference Griffin1974; Manning & Holthuis, Reference Manning and Holthuis1981: 311 (references).
MATERIAL EXAMINED
MU183, 138–177 m, (1), MU209, 115–150 m, (1).
Males: 10.35–11.87 mm.
IDENTIFICATION
Our specimens agree with the description and figures provided by Zariquiey Álvarez (Reference Zariquiey Álvarez1968: 462) and González-Gurriarán & Méndez (Reference González-Gurriarán, Méndez, Castro-Sada and Coruña1986: 178).
BIOLOGY
Mainly reported on firm substrates of relatively large particles, bottoms of shelly sand, calcareous algae, muddy shells and gravel, maerl beds and on rock (Manning & Holthuis, Reference Manning and Holthuis1981; Ballesteros, Reference Ballesteros2006) and in Posidonia meadows (Pipitone & Arculeo, Reference Pipitone and Arculeo2003; Ateş et al., Reference Ateş, Katağan and Kocataş2006). Bathymetric range between 10 and 1216 m (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999: 188). Ovigerous females reported in April (Zariquiey Álvarez, Reference Zariquiey Álvarez1968).
Specimens examined were two males collected at 115–150 and 138–177 m.
GEOGRAPHIC DISTRIBUTION
East Atlantic, from Norway to Angola, including the Azores, Desertas, Canary and Cape Verde Islands; Mediterranean Sea; West Indian Ocean from False Bay, South Africa, to Durban (Griffin, Reference Griffin1974; d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999).
Later records for this species (Abelló et al., Reference Abelló, Carbonell and Torres2002; García Raso & Manjón-Cabeza, Reference García Raso and Manjón-Cabeza2002; Pipitone & Arculeo, Reference Pipitone and Arculeo2003; Ateş et al., Reference Ateş, Katağan and Kocataş2006; Serrano et al., Reference Serrano, Sánchez and García-Castrillo2006; Cartes et al., Reference Cartes, Serrano, Velasco, Parra and Sánchez2007; García-Muñoz et al., Reference García-Muñoz, Manjón-Cabeza and García-Raso2008; Ellis et al., Reference Ellis, Martinez, Burt and Scott2013) fit well within its geographic distribution.
Superfamily parthenopoidea MacLeay, 1838
Family parthenopidae MacLeay, 1838
Genus Distolambrus Tan & Ng, Reference Tan and Ng2007
Distolambrus maltzami (Miers, Reference Miers1881)
(Figure 25)
Heterocrypta Maltzami Miers, Reference Miers1881: 209, pl. 13: figure 1
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Fig. 25. Distolambrus maltzami (Miers, 1881), ovigerous female CL: 8.08 mm, MUBV21, in ethanol, dorsal view. Scale bar: 1 mm.
Heterocrypta maltzami: Manning & Holthuis Reference Manning and Holthuis1981: 322 (references)
Distolambrus maltzami: Tan & Ng, Reference Tan and Ng2007: 103, figure 5; Henriksen, Reference Henriksen2009: 80, figure 40
MATERIAL EXAMINED
MUBV21, 107–109 m, (2).
Ovigerous female: 8.08 mm.
IDENTIFICATION
Our specimen agrees well with the description provided by Milne-Edwards & Bouvier (Reference Milne-Edwards, Bouvier and Milne-Edwards1900: 121, as Heterocrypta Maltzani and Heterocrypta Maltzani var. Marioni).
BIOLOGY
Previously reported from bottoms of shell debris, shelly sand, sand, muddy sand, mud, calcareous algae and rock (Manning & Holthuis, Reference Manning and Holthuis1981, as Heterocrypta maltzami; d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999, as Heterocrypta maltzami marionis). Bathymetric range oscillates between 22 and 550 m (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999, as Heterocrypta maltzami marionis), but all previous West African records are from less than 100 m depth (0–70 m) (Manning & Holthuis, Reference Manning and Holthuis1981; as Heterocrypta maltzami).
On the West African coast, ovigerous females have been found throughout the year (Zariquiey Álvarez, Reference Zariquiey Álvarez1968; as Heterocrypta maltzami).
Only one ovigerous female was collected in December during the MAURIT surveys, between 107 and 109 m.
GEOGRAPHIC DISTRIBUTION
Eastern Atlantic Ocean from the Bay of Biscay to Angola, including the Azores and Cape Verde Islands (Manning & Holthuis, Reference Manning and Holthuis1981), as well as in the Mediterranean Sea (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999).
Later records (Pipitone & Arculeo, Reference Pipitone and Arculeo2003; Serrano et al., Reference Serrano, Sánchez and García-Castrillo2006; Henriksen, Reference Henriksen2009; Massi et al., Reference Massi, Micalizzi, Giusto and Pipitone2010) fit well within its geographic distribution.
REMARKS
In their systematic revision of the subfamily Parthenopinae, Tan & Ng (Reference Tan and Ng2007) relocated the former Heterocrypta maltzami in their new genus Distolambrus. Both genera can be easily differentiated by the presence of a V-shaped ridge on the gastric region of Distolambrus (U-shaped in Heterocrypta); the branchial ridge not continuous with the gastric ridge (continuous in Heterocrypta); male with fused thoracic sternites without a transverse groove (with a broad transverse groove in Heterocrypta); third maxilliped merus subtriangular (subquadrate in Heterocrypta); and the posterior margin not produced beyond the base of the abdomen (produced in Heterocrypta).
Genus Solenolambrus Stimpson, Reference Stimpson1871
Solenolambrus noordendei (Capart, Reference Capart1951)
(Figure 26)
Heterocrypta noordendei Capart, Reference Capart1951: 108, figure 37, pl. II figure 15.
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Fig. 26. Solenolambrus noordendei (Capart, 1951), ovigerous female CL: 9.04 mm dorsal view, MU233.
Solenolambrus noordendei: Manning & Holthuis, Reference Manning and Holthuis1981: 336 (references); Tan, Reference Tan2004: 500, figures 137G, H; 140B (references).
MATERIAL EXAMINED
MU86, 91–103 m, (1); MU170, 102–92 m, (13); MU171, 105–100 m, (19); MU183, 138–177 m, (1); MU210, 86–90 m, (1); MU212, 163–200 m, (2); MU233, 165–189 m, (1); MUBV01, 112 m, (63); MUBV02, 318–330 m, (2); MUBV10, 332–344 m, (6); MUBV15, 148–135 m, (11); MUBV21, 107–109 m, (18).
Males: 8.26–13.61 mm, ovigerous females: 6.71–10.81 mm.
IDENTIFICATION
Our specimens agree well with the original description.
BIOLOGY
Species found on a variety of bottoms such as mud, muddy sand, mud with branched foraminifera, sandy mud, shelly mud, broken shells, coral or rock, at depths between 64 and 215 m. Ovigerous females have been previously recorded in February, March, May, July and November (Manning & Holthuis, Reference Manning and Holthuis1981).
Our specimens were captured in depths from 86–90 to 332–344 m, in bottoms of coarse sand with mud and shell debris, muddy sand and sandy mud. Ovigerous females were collected in November and December.
GEOGRAPHIC DISTRIBUTION
West Africa from Mauritania (present work) to Angola (Manning & Holthuis, Reference Manning and Holthuis1981).
REMARKS
Our records extend northwards the geographic distribution, from Senegal to Cape Blanc. They also broaden the bathymetric range of the species down to 322–344 m. The presence of ovigerous females also in December suggests that this species spawns throughout the year.
Genus Spinolambrus Tan & Ng, Reference Tan and Ng2007
Spinolambrus notialis (Manning & Holthuis, Reference Manning and Holthuis1981)
(Figure 27)
Parthenope notialis Manning & Holthuis, Reference Manning and Holthuis1981: 331, figures 85, 86a–b.
![](https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary-alt:20241103143914-78872-mediumThumb-S002531541600062X_fig27g.jpg?pub-status=live)
Fig. 27. Spinolambrus notialis (Manning & Holthuis, 1981), ovigerous female CL: 10.08 mm dorsal view, MU131.
Spinolambrus notialis: Tan, Reference Tan2004: 524, figures 144A–B, 146B; Tan & Low, Reference Tan and Low2014: 96, figure 2B.
MATERIAL EXAMINED
MU131, 102–104 m, (1); MU186, 174 m, (1); MUBV21, 107–109 m, (1).
Female: 7.21 mm, ovigerous female: 10.08 mm.
IDENTIFICATION
Our specimens agree well with the original description.
BIOLOGY
Species recorded on bottoms of mud, sandy mud or sand, mostly mixed with broken shells, bryozoans, branched or foliate foraminifera, calcareous algae, corals and rocks. Bathymetric range from 18 to 162 m (Manning & Holthuis Reference Manning and Holthuis1981: 335 as Parthenope notialis). Ovigerous females have been reported in February, March, May, June, July, September and November (Zariquiey Álvarez, Reference Zariquiey Álvarez1968: 441 as Parthenope macrochelos; Manning & Holthuis, Reference Manning and Holthuis1981: 335 as Parthenope notialis).
Our specimens were captured from 102–104 to 174 m, on coarse sand and muddy sand bottoms. Ovigerous females were collected in December.
GEOGRAPHIC DISTRIBUTION
West Africa from Mauritania (present work, see Remarks) to Angola (Manning & Holthuis, Reference Manning and Holthuis1981: 335 as Parthenope notialis). Muñoz et al. (Reference Muñoz, García-Isarch, Sobrino, Burgos, Funny and González-Porto2012) reported this species from Guinea-Bissau.
REMARKS
This species was relocated by Tan & Ng (Reference Tan and Ng2007) in their new genus Spinolambrus erected to include some species of the subfamily Parthenopinae previously placed within the genera Lambrus and Parthenope from both the Atlantic and the Eastern Pacific.
Manning & Holthuis (Reference Manning and Holthuis1981) established the distribution range of this species at least from Senegal to Angola, inasmuch as the authors pointed out that most of the records of S. macrochelos (as P. macrochelos) from tropical West Africa were based on S. notialis. In consequence, they described the records of Maurin (Reference Maurin1968; as Lambrus macrocheles) from the Western Sahara and Mauritanian waters as dubious, and a further taxonomic revision would assign them to S. notialis or S. macrochelos. The same occurs with the citation in Fransen (Reference Fransen1991), although in this case Fransen referred to the specimens as ‘Parthenope notialis/miersi’ [Parthenope miersii was synonymized with S. macrochelos (Tan, Reference Tan2004)].
The southernmost distribution of S. macrochelos was confirmed by Tan & Low (Reference Tan and Low2014) in Salé (Morocco) and the authors suggest that the distributions of S. macrochelos and S. notialis may overlap in North-west Africa.
Our record off Banc d'Arguin confirms the presence of this species in Mauritanian waters and slightly extends its bathymetric distribution.
The presence of ovigerous females in December confirms that this species spawns all the year round.
Superfamily portunoidea Rafinesque, 1815
Family carcinidae MacLeay, 1838
Genus Liocarcinus Stimpson, Reference Stimpson1871
Liocarcinus corrugatus (Pennant, Reference Pennant1777)
(Figure 28A–C)
Cancer corrugatus Pennat, Reference Pennant1777: 5, pl. V figure 9.
![](https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary-alt:20241103143914-77403-mediumThumb-S002531541600062X_fig28g.jpg?pub-status=live)
Fig. 28. Liocarcinus corrugatus (Pennant, 1777), male CL: 13.07 mm, MUBV08: (A) dorsal view; (B) same, in ethanol; (C) dorsal carapace, in alcohol. Scale bars: B, 1 cm; C, 1 mm.
Liocarcinus corrugatus: Manning & Holthuis, Reference Manning and Holthuis1981: 84 (references).
MATERIAL EXAMINED
MUBV08, 174–168 m, (2).
Male: 13.07 mm.
IDENTIFICATION
Our specimen agrees well with those described by Zariquiey Álvarez (Reference Zariquiey Álvarez1968) and González-Gurriarán & Méndez (Reference González-Gurriarán, Méndez, Castro-Sada and Coruña1986).
BIOLOGY
Mainly reported on coarse soft bottoms such as gravel, coarse sand, sandy mud, muddy sand and maerl beds (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999; as Polybius (Necora) corrugatus; El Lakhrach et al., Reference El Lakhrach, Hattour, Jarboui, Elhasni and Ramos-Esplá2012), but also recorded on rocks and in Posidonia oceanica (Linnaeus) Delile, 1813 meadows (Pipitone & Arculeo, Reference Pipitone and Arculeo2003; as Polybius corrugatus; Pipitone & Vaccaro, Reference Pipitone, Vaccaro, Pessani, Tirelli and Froglia2011). Depth records from 5–10 m (Pipitone & Arculeo, Reference Pipitone and Arculeo2003; as Polybius corrugatus) to 147 m (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999; as Polybius (Necora) corrugatus). Ovigerous females reported from November to January, March, May, June and July (Zariquiey Álvarez, Reference Zariquiey Álvarez1968; as Macropipus corrugatus; García Raso, Reference García Raso1984).
Only one male was captured during the MAURIT surveys, between 168 and 174 m.
GEOGRAPHIC DISTRIBUTION
East Atlantic, from the Orkney Islands to Angola, including the Azores, Madeira, Canary and Cape Verde Islands; and Mediterranean Basin (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999: 219) (see Remarks).
Further records in the literature (Abelló et al., Reference Abelló, Carbonell and Torres2002; García Raso & Manjón-Cabeza, Reference García Raso and Manjón-Cabeza2002; Pipitone & Arculeo, Reference Pipitone and Arculeo2003; as Polybius corrugatus; Ateş et al., Reference Ateş, Katağan and Kocataş2006; García-Muñoz et al., Reference García-Muñoz, Manjón-Cabeza and García-Raso2008; Pipitone & Vaccaro, Reference Pipitone, Vaccaro, Pessani, Tirelli and Froglia2011; El Lakhrach et al., Reference El Lakhrach, Hattour, Jarboui, Elhasni and Ramos-Esplá2012; Muñoz et al., Reference Muñoz, García-Isarch, Sobrino, Burgos, Funny and González-Porto2012) fit well within this geographic distribution.
REMARKS
Liocarcinus corrugatus was also reported from many localities in the Indo-Pacific, but d'Udekem d'Acoz (Reference d'Udekem d'Acoz1999) stated that all these records probably belong to one or more closely allied species. Ng et al. (Reference Ng, Wang, Ho and Shih2001) followed d'Udekem d'Acoz and excluded L. corrugatus from the checklist of brachyurans from Taiwan, but later Yaldwin & Webber (Reference Yaldwyn and Webber2011) included this species in the Decapoda checklist of New Zealand.
Our data slightly increase the bathymetric range previously reported in the literature.
Family geryonidae Colosi, Reference Colosi1923
Genus Chaceon Manning & Holthuis, Reference Manning and Holthuis1989
Chaceon maritae (Manning & Holthuis, Reference Manning and Holthuis1981)
(Figure 29A, B)
Geryon maritae Manning & Holthuis, Reference Manning and Holthuis1981: 112, figures 24a, 25, 26.
![](https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary-alt:20241103143914-80903-mediumThumb-S002531541600062X_fig29g.jpg?pub-status=live)
Fig. 29. Chaceon maritae (Manning & Holthuis, 1981): (A) adult dorsal view, MU243; (B) juvenile CL: 21.94 mm dorsal view, MU177.
MATERIAL EXAMINED
MU01, 817–820 m, (2); MU15, 670–675 m, (1); MU17, 818–861 m, (3); MU18, 519–402 m, (40); MU19, 1222–1218 m, (1); MU23, 532–415 m, (2); MU26, 744 m, (1); MU33, 741–736 m, (2); MU46, 848–847 m, (3); MU48, 1239–1218 m, (1); MU55, 1310–1218 m, (1); MU56, 1091–1159 m, (1); MU62, 1236–1244 m, (1); MU67, 1381–1390 m, (1); MU68, 1136–1146 m, (5); MU70, 755–801 m, (3); MU73, 1330–1284 m, (1); MU79, 554–576 m, (5); MU126, 668–826 m, (2); MU177, 584–580 m, (9); MU182, 726 m, (1); MU188, 627 m, (1); MU243, 827–850 m, (3); MU267, 673–670 m, (16); MUBV03, 528–538 m, (1); MUBV13, 493–517 m, (1).
Males: 23.63–68.34 mm; females: 21.94–77.95 mm.
IDENTIFICATION
Our specimens agree well with the original description.
BIOLOGY
Species inhabiting bottoms of mud, sandy mud and corals at depths from 100–300 m (Manning & Holthuis, Reference Manning and Holthuis1981; as Geryon maritae) to 1994 m (Le Roux, Reference Le Roux2001).
Although several works focus on the population dynamics of C. maritae for fishing purposes (Gaertner & Laloé, Reference Gaertner and Laloé1986; Melville Smith, Reference Melville Smith1988; Le Roux, Reference Le Roux2001), surprisingly, none of them include data about the spawning period for this species.
Our specimens were collected between 402–519 and 1381–1390 m on muddy sand and sandy mud bottoms.
GEOGRAPHIC DISTRIBUTION
West Africa from the Western Sahara to Namibia (Manning & Holthuis, Reference Manning and Holthuis1981; d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999) (see Remarks). Le Roux (Reference Le Roux2001) reported this species off Namibia and Muñoz et al. (Reference Muñoz, García-Isarch, Sobrino, Burgos, Funny and González-Porto2012) from Guinea-Bissau.
REMARKS
Species occurrence in the Canary Islands (González et al., Reference González, Santana and Fernández-Vergaz1996) needs confirmation (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999).
Family macropipidae Stephenson & Campbell, Reference Stephenson and Campbell1960
Genus Bathynectes Stimpson, Reference Stimpson1871
Bathynectes piperitus Manning & Holthuis, Reference Manning and Holthuis1981
(Figure 30)
Bathynectes piperitus Manning & Holthuis, Reference Manning and Holthuis1981: 77, figures 16, 17.
![](https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary-alt:20241103143914-36892-mediumThumb-S002531541600062X_fig30g.jpg?pub-status=live)
Fig. 30. Bathynectes piperitus Manning & Holthuis, 1981, male CL: 13.60 mm dorsal view, MU123.
MATERIAL EXAMINED
MU02, 616–626 m, (1); MU14, 502–511 m, (6); MU15, 670–675 m, (1); MU18, 519–402 m, (31); MU23, 532–415 m (2); MU37, 403–442 m (5); MU44, 606–596 m, (14); MU45, 420–427 m, (2); MU51, 464–468 m, (2); MU52, 774–792 m, (2); MU54, 414–451 m, (1); MU57, 430–406 m, (1); MU63, 848–798 m, (59); MU64, 452–468 m, (11); MU70, 755–801 m, (1); MU71, 812–837 m, (6); MU79, 554–576 m, (2); MU123, 278–362 m, (1); MU126, 668–826 m, (3); MU127, 260–353 m, (1); MU134, 311–436 m, (1); MU143, 322 m, (2); MU175, 618–850 m, (2); MU177, 584–580 m, (2); MU179, 303–304 m, (1); MU214, 650–588 m, (1); MU222, 729–723 m, (2); MU267, 673–670 m, (1); MU276, 637–562 m, (1); MUBV02, 318–330 m, (6); MUBV03, 528–538 m, (4); MUBV09, 549–555 m, (10); MUBV10, 332–344 m, (4); MUBV13, 493–517 m (30); MUBV14, 300–281 m, (2); MUBV18, 559–574 m, (24); MUBV19, 306 m, (30); MUBV22, 300 m, (16); MUBV25, 499–520 m, (3).
Males: 13.60–55.06 mm; females: 13.13–41.98 mm; ovigerous females: 35.52–40.72 mm.
IDENTIFICATION
Our specimens agree well with the original description.
BIOLOGY
Collected on mud, muddy sand, sand and gravel bottoms and on corals, in depths from 200 to 628 m. Ovigerous females recorded from February to June (Manning & Holthuis, Reference Manning and Holthuis1981).
Our specimens were captured at depths from 260–353 to 618–850 m on sand and sandy mud bottoms. Ovigerous females were collected in November and December.
GEOGRAPHIC DISTRIBUTION
Known in West Africa, from the Cape Verde Islands (Manning & Holthuis, Reference Manning and Holthuis1981) and Mauritania (present work) to Namibia (Macpherson, Reference Macpherson1991), the species is probably present up to Western Saharan waters (see Remarks). Henriksen (Reference Henriksen2009) reports this species off Gabon.
REMARKS
Specimens of Bathynectes piperitus have been largely identified under the name of the European and Mediterranean B. maravigna (Prestandrea, 1839). The differences between both species were summarized by Manning & Holthuis (Reference Manning and Holthuis1981). Our Mauritanian record of B. piperitus supports the opinion expressed by Manning & Holthuis (Reference Manning and Holthuis1981) that the records of B. superbus (=B. maravigna) from Cape Bojador (Western Sahara) and Mauritania, given by Maurin (Reference Maurin1968), and those of Bathynectes, given by Filhol (Reference Filhol1885), may in fact be referable to B. piperitus.
Our records of ovigerous females determine a spawning period between November and June.
Genus Macropipus Prestandrea, 1833
Macropipus rugosus (Doflein, Reference Doflein and Chun1904)
(Figure 31)
Elliptodactylus rugosus Doflein, Reference Doflein and Chun1904: 94, pl. XXX figures 1–3, pl. XXXII figure 7.
![](https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary-alt:20241103143914-77138-mediumThumb-S002531541600062X_fig31g.jpg?pub-status=live)
Fig. 31. Macropipus rugosus (Doflein, 1904), ovigerous female CL: 23.43 mm dorsal view, MUBV01.
Macropipus rugosus Manning & Holthuis, Reference Manning and Holthuis1981: 86 (references).
MATERIAL EXAMINED
MU86, 91–103 m, (3); MU90, 110 m, (6); MU101, 104–96 m, (2304); MU120, 109–105 m, (1); MU125, 103–101 m, (132); MU135, 185–173 m, (25); MU137, 81–84 m, (1); MU138, 123–130 m, (233); MU144, 119–138 m, (8); MU147, 134–139 m, (7); MU148, 215–245 m, (106); MU149, 93–146 m, (5); MU154, 92–102 m, (4819); MU155, 210–257 m, (25); MU160, 143–147 m, (565); MU161, 89–92 m, (6); MU162, 148–149 m, (232); MU166, 87–85 m, (37); MU167, 101–108 m, (354); MU168, 87–92 m, (1); MU181, 142–148 m, (18); MU184, 213–202 m, (54); MU186, 174 m, (1); MU205, 89–93 m, (1); MU208, 96–79 m, (1); MU224, 173–177 m, (12); MU227, 183–181 m, (3); MU266, 103 m, (57); MU285, 128–132 m, (31); MU291, 106–137 m, (256); MUBV01, 112 m, (1); MUBV21, 107–109 m, (4).
Males: 7.39–32.41 mm, females: 7.43–24.95 mm, ovigerous females: 18.49–23.43 mm.
IDENTIFICATION
Our specimens agree well with the original description and with the notes provided by Guinot (Reference Guinot1961: 2).
BIOLOGY
Bottoms of mud, muddy sand, sand, broken shells, foraminifera, bryozoans and also on calcareous algae and rocks, at depths from 5 to 400 m. Ovigerous females previously recorded in June and September (Manning & Holthuis, Reference Manning and Holthuis1981: 86).
Our specimens were captured between 81–84 and 210–257 m on muddy sand, sandy mud, coarse muddy sand, sand and on coarse sand with shell debris bottoms. Ovigerous females were collected in November and December.
GEOGRAPHIC DISTRIBUTION
West Africa from Western Sahara to Angola, including São Tomé and Príncipe Islands (Manning & Holthuis, Reference Manning and Holthuis1981: 86). Further records can be found at Nigeria, Principe and Congo (Henriksen, Reference Henriksen2009) and Guinea-Bissau (Muñoz et al., Reference Muñoz, García-Isarch, Sobrino, Burgos, Funny and González-Porto2012).
REMARKS
Ovigerous females were reported for the first time at the end of the year, indicating that this species spawns in the second half of the year.
Superfamily xanthoidea MacLeay, 1838
Family xanthidae MacLeay, 1838
Genus Monodaeus Guinot, Reference Guinot1967
Monodaeus cristulatus Guinot & Macpherson, Reference Guinot, Richer de Forges and Crosnier1988
(Figures 32A–C & 33E–H)
Monodaeus cristulatus Guinot & Macpherson, Reference Guinot, Richer de Forges and Crosnier1988: 744, figures 8, 9, 14, 19, 20, 25, 26, pl. 3 figures D–G.
![](https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary-alt:20241103143914-94819-mediumThumb-S002531541600062X_fig32g.jpg?pub-status=live)
Fig. 32. Monodaeus cristulatus Guinot & Macpherson, 1988: (A) medium size specimen, CL ~ 5 mm dorsal view, MUBV10; (B) male CL: 14.08 mm dorsal view, MUBV14; (C) same in ethanol.
![](https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary-alt:20241103143914-74476-mediumThumb-S002531541600062X_fig33g.jpg?pub-status=live)
Fig. 33. Monodaeus couchii (Couch, 1851), males: (A) CL: 22.11 mm dorsal view; (B) same, right pereiopods P3, P4, P5 detail; (C) CL: 16.09 mm dorsal view; (D) same, right pereiopods P2 to P5 detail. Monodaeus cristulatus Guinot & Macpherson, 1988, males: (E) CL: 12.02 mm dorsal view; (F) same, right pereiopods P2 to P5 detail; (G) CL: 8.36 mm dorsal view; (H) same, left pereiopods P2 to P5 detail. (A, B) ICMD 291/1981(s01); (C, D) ICMD000013; (E, F) MU212, UVIGO/03087; (G, H) MUBV10, UVIGO/11948. Scale bars: A, C, E, G, 1 cm; B, D, F, H, 1 mm.
MATERIAL EXAMINED
MU120, 109–105 m, (1); MU171, 105–100 m, (5); MU212, 163–200 m, (1); MU219, 125–129 m, (1); MU290, 311 m, (1); MUBV08, 174–168 m, (2); MUBV10, 332–344 m, (12); MUBV14, 300–281 m (1).
Males: 4.36–14.08 mm, females: 3.53–4.96 mm, ovigerous female: 6.57 mm.
COMPARATIVE MATERIAL EXAMINED
Monodaeus cristulatus: Namibia, Exp. Benguela XII, station P42, 185 m, 28°56′59.99″S 14°55′47.99″E, Macpherson, E. det., 10 specimens (USNM 221965); Exp. Benguela XIV, station P53, 179–183 m, 29°19′S 15°05′E, Macpherson, E. det., three specimens: 1 male CL: 15.58 mm (ICMD000034) and two juveniles CL: 4.30 mm (ICMD000037) and 3.77 mm (ICMD000039).
Monodaeus couchii (Couch, 1851): Gulf of Cádiz, Exp. ARSA0311, station L14, 692 m, 36°12′54.0″N 7°00′32.4″W, 2 females CL: 11.31 and 9.20 mm (IEO-CD-AR11/001); station L31, 359 m, 36°02′27.6″N 6°20′09.6″W, 1 male CL: 7.31 mm (IEO-CD-AR11/017). Morocco (Atlantic coast): Exp. CCLME 2012, station 297, 239 m, 34°52′03.0″N 6°45′24.1″W, 1 female CL: 14.23 mm (IEO-CD-CCLME12/1226). NW Mediterranean: Barcelona, Exp. Pont 90, station G3, 192–430 m, 41°07′59.9″N 2°03′00.0″E, Sardá, F. det., 4 specimens: 2 males CL: 22.11 mm (ICMD 291/1981(s01)) and 22.42 mm (ICMD 291/1981(s02)), 2 ovigerous females CL: 17.03 mm (ICMD 291/1981(s03)) and 14.25 mm (ICMD 291/1981(s04)). SW Mediterranean: Málaga, Exp. Medits94 station Lance 16, 553–563 m, 36°23′48.1″N 4°28′54.5″W (off Pta. Calaburras), Abelló, P. det., 1 female CL: 12.34 mm (ICMD000005); Exp. Medits97 station Lance 6, 721–735 m, 36°18′33.1″N 4°44′34.1″W (off Marbella), Abelló, P. det., 1 male CL: 16.09 mm (ICMD000013); Exp. Medits97 station Lance 19, 524–526 m, 36°36′59.4″N 3°48′56.8″W (off Nerja), Abelló, P. det., 1 male CL: 20.60 mm (ICMD000017).
IDENTIFICATION
Our specimens agree with the original description (see Remarks).
BIOLOGY
This species was captured on coral, at depths from 160 to 300 m; ovigerous females were reported from July and August (label with paratypes ICMD000031, ICMD000032 and ICMD000033; Guinot & Macpherson, Reference Guinot, Richer de Forges and Crosnier1988: 744).
Our specimens were found on sandy bottoms and hidden in the holes of large stones hauled up in the net, at depths from 100–105 to 332–344 m. The ovigerous female was captured in December.
GEOGRAPHIC DISTRIBUTION
Western Africa in Mauritania (present work) and Namibia (Guinot & Macpherson, Reference Guinot, Richer de Forges and Crosnier1988) (see Remarks).
REMARKS
Monodaeus cristulatus has only been previously reported from Namibian coasts (28°–29°S). Following Guinot & Macpherson (Reference Guinot, Richer de Forges and Crosnier1988), the closest species is Monodaeus couchii, previously reported from various localities of NW Africa (Manning & Holthuis, Reference Manning and Holthuis1981), including Mauritanian waters (Milne-Edwards & Bouvier, Reference Milne-Edwards, Bouvier and Milne-Edwards1900; Anadon, Reference Anadon1981; Fransen, Reference Fransen1991). We have compared our material with specimens of M. cristulatus from Namibia and with samples of M. couchii from the Western Mediterranean, Gulf of Cádiz and Moroccan Atlantic coasts (see comparative material examined). After these comparisons, we included our specimens in M. cristulatus, due to the presence of a dorsal crest on P2–P5 meri and a proximal protuberance on P2–P4 dorsal carpi (Figure 33E–H), which are absent in M. couchii (Figure 33A–D). In juveniles of M. cristulatus the specific characters are not clearly developed (Guinot & Macpherson, Reference Guinot, Richer de Forges and Crosnier1988), which hinders species separation when studying isolated specimens. However, checking a series of different features on specimens of various sizes allows for accurate identification.
The possible misidentification of the two species and our findings of Monodaeus cristulatus in Mauritanian waters point to a need for a revision of the records of M. couchii south of Cape Blanc. Our records suggest that M. cristulatus can be found along the West African coast between Banc d'Arguin, Mauritania and Namibia.
Subsection thoracotremata Guinot, Reference Guinot1977
Superfamily grapsoidea MacLeay, 1838
Family plagusiidae Dana, Reference Dana1851
Genus Euchirograpsus H. Milne-Edwards, Reference Milne-Edwards1853
Euchirograpsus liguricus H. Milne-Edwards, Reference Milne-Edwards1853
(Figure 34)
Euchirograpsus liguricus H. Milne-Edwards, Reference Milne-Edwards1853: 175; Türkay, Reference Türkay1975: 105, figures 1–3, 17, 23 (references)
![](https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary-alt:20241103143914-78951-mediumThumb-S002531541600062X_fig34g.jpg?pub-status=live)
Fig. 34. Euchirograpsus liguricus H. Milne-Edwards, 1853, male CL: 20.68 mm dorsal view, MUBV14.
.
MATERIAL EXAMINED
MU290, 311 m, (3); MUBV14, 300–281 m, (3).
Males: 6.61–20.68 mm; females: 6.30–11.39 mm.
IDENTIFICATION
Our specimens agree well with the description provided by Türkay (Reference Türkay1975: 105) and with the comments and pictures after Giacobbe & Spano (Reference Giacobbe and Spano2006).
BIOLOGY
Collected on gravel, shell, sand, sandy mud and muddy bottoms, the species is also recorded on bottoms of rock with gorgonians, sponges, calcareous algae and kelp, and coral reefs (Manning & Holthuis, Reference Manning and Holthuis1981: 248; d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999: 250, Giacobbe & Spano, Reference Giacobbe and Spano2006). Depth records range between 10 m (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999: 250) and 620–621 m (Abelló et al., Reference Abelló, Carbonell and Torres2002). Ovigerous females were collected only in March (Manning & Holthuis, Reference Manning and Holthuis1981).
Our material was collected at 281–300 and 311 m. The species was caught in a ghost net hauled with the Lofoten trawl and in the holes of a large stone captured with the beam trawl.
GEOGRAPHIC DISTRIBUTION
East Atlantic from West Portugal to Namibia including the Azores, Madeira, Savage, Canary and Cape Verde Islands; Western and Central Mediterranean Sea (d'Udekem d'Acoz, Reference d'Udekem d'Acoz1999; Giacobbe & Spano, Reference Giacobbe and Spano2006). Abelló et al. (Reference Abelló, Carbonell and Torres2002) report this species from the Eastern Alborán Sea.
DISCUSSION
A total of 10,514 brachyuran specimens were studied from waters off Mauritania at depths from 79 to 1867 m, comprising 33 species belonging to 17 families and 24 genera. The brachyuran diversity found in recent works carried out with a similar methodology on West African coasts is slightly lower than the values of our study. García Raso (Reference García Raso1996) reported 31 brachyuran species from the Ibero-Moroccan Bay and adjacent waters at depths from 137 to 2142 m; Muñoz et al. (Reference Muñoz, García-Isarch, Sobrino, Burgos, Funny and González-Porto2012) listed 44 brachyuran species off Guinea-Bissau at depths between 20 and 1000 m, of which only 24 were found deeper than 80 m; and Macpherson (Reference Macpherson1991) included 21 brachyuran species in Namibian waters from 100 to 1000 m.
The most abundant species so far was Macropipus rugosus (89%), mainly due to the high number of specimens captured at two stations off Nouakchott (MU101, 2304 specimens; MU154, 4819 specimens). However, the remaining stations showed similar abundances to those reported for this species in Guinea-Bissau (Muñoz et al., Reference Muñoz, García-Isarch, Sobrino, Burgos, Funny and González-Porto2012), Guinea (Le Loeuf, Reference Le Lœuff1993) and the Gulf of Guinea (Henriksen, Reference Henriksen2009). Conversely, the following three species, Atelecyclus rotundatus, Ethusa rosacea and Macropodia longipes, were represented by only one specimen each. The low abundance of A. rotundatus is probably explained by its burrowing behaviour (González-Gurriarán & Méndez, Reference González-Gurriarán, Méndez, Castro-Sada and Coruña1986; S. de Matos-Pita, unpublished data), which hinders or prevents the capture of the species by the trawl. Macropodia longipes, although reported from the Gulf of Gascogne to the Cape Verde Islands and the Mediterranean Sea, has only been recorded in Mauritanian waters on the Banc d'Arguin (Forest, Reference Forest1978, as Macropodia tenuirostris longipes). Ethusa rosacea and M. longipes are both considered as rare species because they were seldom reported and always in very low abundance (Monod, Reference Monod1956; Manning & Holthuis, Reference Manning and Holthuis1981; Henriksen, Reference Henriksen2009).
Different patterns are observed for the latitudinal distribution of the species in the area studied. Eleven species, Homola barbata, Paromola cuvieri, Acanthocarpus brevispinis, Goneplax barnardi, Inachus angolensis, Macropodia gilsoni, M. macrocheles, Chaceon maritae, Bathynectes piperitus, Macropipus rugosus and Monodaeus cristulatus, are evenly distributed along most of the Mauritanian coast. Three species, Medorippe lanata, Eurynome aspera and Distolambrus maltzami, were sampled only north of Cape Timiris, and another six, Ethusa rosacea, Pisa armata, Inachus aguiarii, Macropodia hesperiae, M. longipes and Euchirograpsus liguricus were only captured between Cape Timiris and the Senegalese border. Calappa pelii, Ethusa rugulosa, Goneplax rhomboides, Pseudomyra mbizi, Inachus grallator, I. nanus, Solenolambrus noordendei and Spinolambrus notialis were scarcely sampled north and south of Cape Timiris. Finally, another five species, Cymonomus granulatus, Atelecyclus rotundatus, Ebalia nux, Inachus leptochirus and Liocarcinus corrugatus, were sampled only off Cape Blanc.
Analysis of the geographic distribution reveals three different contingents within the Mauritanian brachyuran fauna. The main zoogeographic group (18 species, 54.5%) is composed of species distributed along the West African coast, mainly in the tropical region. Within this group, 14 species, A. brevispinis, C. pelii, E. rosacea, E. rugulosa, G. barnardi, P. mbizi, I. angolensis, M. gilsoni, M. macrocheles, S. noordendei, S. notialis, C. maritae, B. piperitus and M. rugosus were mostly evenly reported from the Western Sahara and Mauritania to Angola and Namibia. Two species, I. nanus and M. hesperiae, were reported between Mauritania and the Gulf of Guinea. Inachus grallator, previously reported between the Gulf of Guinea and Namibia, and M. cristulatus, only known from the type locality (Namibia), have a discontinuous latitudinal distribution and are now both recorded in Mauritanian waters. Species with a restricted and/or discontinuous distribution in tropical West Africa have been reported in recent decades (Manning & Holthuis, Reference Manning and Holthuis1981; Macpherson, Reference Macpherson1991), but a more complete sampling effort in West African waters will certainly extend their known distribution.
The second zoogeographic group (10 species, 30.3%) is represented by species widely distributed in the Lusitanian province, including the Mediterranean Sea. The southern limit of the Lusitanian province was recently established at Cape Juby (south Morocco) by Briggs & Bowen (Reference Briggs and Bowen2012), but set up at Cap Vert (Senegal) by Briggs (Reference Briggs1995). Macropodia longipes is the only species with a strictly Lusitanian distribution, while the distributional limits of the remaining species extend further northwards or southwards. Five species range southwards to Guinea (I. aguiarii), Angola (D. maltzami), Namibia (E. liguricus), South Africa (H. barbata) and even into the Indian Ocean up to Mozambican waters (M. lanata). Another four species, C. granulatus, G. rhomboides, E. nux and I. leptochirus, extend their distribution area to northern European seas.
Finally, the third zoogeographic group includes five species (15%) widely distributed in the E Atlantic and Mediterranean, with records from northern European seas southwards to Angola (P. armata and L. corrugatus), South Africa (P. cuvieri and A. rotundatus) and even to Durban in the West Indian Ocean (E. aspera).
In this region, the affinity with Atlantic-Mediterranean fauna was previously reported for different taxa: Porifera (van Soest, Reference Van Soest1993), Hydrozoa (Ansín Agís et al., Reference Ansín Agís, Ramil and Vervoort2001), Bryozoa (Aristegui & Cruz, Reference Aristegui and Cruz1986) and Echinodermata (Hernández et al., Reference Hernández, Clemente, Tuya, Pérez-Ruzafa, Sangil, Moro-Abad, Bacallado-Aránega, Alvarado and Solís-Marín2013; Calero et al., Reference Calero, Ramil, Ramos, Ramos, Sanz and Ramilin preparation). For all these taxa, the Canary Current was highlighted as the main means of larval dispersion southwards, also enabling the presence of typical boreal species, such as C. granulatus, G. rhomboides, E. nux and I. leptochirus, in NW African waters. Conversely, the presence of tropical species in Mauritania (C. pelii, E. rosacea, E. rugulosa, G. barnardi, P. mbizi, I. angolensis, I. grallator, I. nanus, M. gilsoni, M. hesperiae, M. macrocheles, S. noordendei, S. notialis, C. maritae, B. piperitus and M. rugosus) is more difficult to explain because of dominant current systems precluding their dispersion northwards (Aristegui & Cruz, Reference Aristegui and Cruz1986; Hernández et al., Reference Hernández, Clemente, Tuya, Pérez-Ruzafa, Sangil, Moro-Abad, Bacallado-Aránega, Alvarado and Solís-Marín2013). However, a thin surface coastal current flowing northwards along the Gabonese and Congolese coasts (Stramma & Schott, Reference Stramma and Schott1999), and the so-called ‘Mauritania Current’ (Stramma & Schott, Reference Stramma and Schott1999) flowing northwards in summer and autumn between Cap Vert and Cape Blanc (Pastor et al., Reference Pastor, Pelegrí, Hernández-Guerra, Font, Salat and Emelianov2008), probably facilitate the northwards dispersion of tropical species along the West African coastline. The importance of small currents in the dispersion of species was highlighted by Briggs (Reference Briggs1995) as the main way of successfully transporting species in an opposite direction to that of the main flow across the Tropical Pacific and the Atlantic Oceans.
There are three bathymetrical groups of brachyurans in Mauritanian waters. The first group includes 13 species collected exclusively on the shelf (80–200 m; C. pelii, M. lanata, P. mbizi, P. armata, I. aguiarii, I. nanus, M. gilsoni, M. hesperiae, M. longipes, E. aspera, D. maltzami, S. notialis and L. corrugatus); the second is represented by nine species collected both on the shelf and upper slope (80–400 m; H. barbata, E. rugulosa, G. rombhoides, I. angolensis, I. leptochirus, M. macrocheles, S. noordendei, M. rugosus and M. cristulatus); and the third accounts for nine species exclusively recorded along the upper slope (300–700 m; P. cuvieri, C. granulatus, A. brevispinis, A. rotundatus, G. barnardi, E. nux, I. grallator, B. piperitus and E. liguricus) (Figure 35). In addition, C. maritae was collected on the upper and middle slope (460–1385 m) and E. rosacea only at 1024 m.
![](https://static.cambridge.org/binary/version/id/urn:cambridge.org:id:binary:20170825081516642-0825:S002531541600062X:S002531541600062X_fig35g.gif?pub-status=live)
Fig. 35. Depth records of brachyuran species. Translucent continuous line from MAURIT surveys, dotted line from the literature.
Despite the increase in decapod diversity with depth described in several regions, peaking between 1000 and 2000 m (see Muñoz et al., Reference Muñoz, García-Isarch, Sobrino, Burgos, Funny and González-Porto2012 for references), brachyuran crabs do not seem to follow this general trend in Mauritanian waters.
Our results emphasize the highest specific richness of the deep shelf (22 species, 66.66%) and its drastic reduction beyond 700 m (two species, 6.06%). The importance of crustaceans and at least of some brachyuran families (Inachidae, Portunidae, Parthenopidae and Dorippiidae) on the Mauritanian deep shelf was previously pointed out by Duineveld et al. (Reference Duineveld, Lavalaye, Van Noort, Wolff, Van Der Land, Nieuhuis and de Wilde1993). A similar bathymetric pattern was described for the brachyurans off Guinea-Bissau by Muñoz et al. (Reference Muñoz, García-Isarch, Sobrino, Burgos, Funny and González-Porto2012), clearly dominated by shelf species and only three typical deep species, which also include C. maritae. Soto (Reference Soto1991) also reported a strong reduction in species diversity beyond 500 and 700 m in the Straits of Florida. This reduction was related to the high trophic level of brachyurans and the more limited food availability in deep waters (Escobar-Briones et al., Reference Escobar-Briones, Gaytan-Caballero and Legendre2008).
Analysis of bathymetric ranges shows that in Mauritanian waters all but three of the studied species are stenobathic, with depth ranges narrower than 300 m (Menzies et al., Reference Menzies, George and Rowe1973; Pielou, Reference Pielou1992). Only P. cuvieri, C. maritae and B. piperitus (depth ranges of 322, 397 and 925 m, respectively) can be considered eurybathic species. However, considering data from the literature, only 13 of the studied species are stenobathic (E. rugulosa, P. mbizi, P. armata, I. aguiarii, I. grallator, I. nanus, M. gilsoni, M. hesperiae, M. macrocheles, S. noordendei, S. notialis, L. corrugatus and M. cristulatus). In consequence, the Mauritanian brachyuran fauna is dominated by eurybathic species, a feature also shared with the brachyurans of the Straits of Florida (Soto, Reference Soto1991).
ACKNOWLEDGEMENTS
The authors wish to thank all those who made possible the study of the material captured in Maurit surveys, especially Dr Ana Ramos and the Spanish Institute of Oceanography (IEO). We thank Drs Félix Pérez and Pere Abelló from the Instituto de Ciencias Marinas (ICM-CSIC) for their invaluable help and patience in locating some of the comparison material checked here. We would also like to thank Luis Miguel Agudo for providing the map.
FINANCIAL SUPPORT
This work was undertaken within the framework of the ECOAFRIK project and was partially funded by the Galician government (Spain, Project Ref. 10MMA602020PR) and by the ‘MAVA Fondation pour la nature’ (MAVA contract 12/87 AO C4/2012). This is ECOAFRIK publication number 5.