Brachyuran crabs (Crustacea: Decapoda) from the Canary Islands (eastern Atlantic): checklist, zoogeographic considerations and conservation

Brachyuran crabs (Crustacea: Decapoda) from the Canary Islands (eastern Atlantic): checklist, zoogeographic considerations and conservation José Antonio González Ecología Marina Aplicada y Pesquerías, i-UNAT, Universidad de Las Palmas de Gran Canaria, Campus Universitario de Tafira, 35017 Las Palmas de Gran Canaria, Spain. E-mail: [email protected] Summary: Just 20 years have passed since González (1995) finished one of his seminal works on decapod crustaceans of the Canary Islands, thanks to the help of the reputed carcinologists L.B. Holthuis and C.H.J.M. Fransen. This publication allowed d’Udekem d’Acoz (1999) to include the Canarian decapods in his inventory of the NE Atlantic. No checklists of decapod fauna specifically covering this area have been published since then, and an update is needed. The current list of Canarian brachyuran crabs comprises 132 species. Additional species have been recorded thanks to intensified research into deep water, natural range expansions from nearby areas, introduction by anthropogenic activities and description of new taxa; several of these changes are detailed in this review. Although the description of new brachyuran species is not expected to occur at a significant rate, an increase in the number of species from the Canaries is expected to result from trawling and dredging sampling, as well as from introduction of non-native species. For the first time, some zoogeographic comments on the Canarian brachyuran carcinofauna are made. Finally, crab species of commercial interest are listed, their current threats are identified and some updated conservation measures are proposed. Keywords: checklist; zoogeography; conservation; Brachyura; Decapoda; Crustacea; Canary Islands; eastern Atlantic. Cangrejos braquiuros (Crustacea: Decapoda) de las Islas Canarias (Atlántico oriental): lista comentada, consideraciones zoogeográficas y conservación Resumen: Se cumplen 20 años desde que González (1995) publicó uno de sus trabajos seminales sobre crustáceos decápodos de Canarias, gracias a la ayuda de los reputados carcinólogos L.B. Holthuis y C.H.J.M. Fransen. Dicha publicación permitió a d’Udekem d’Acoz (1999) incluir los decápodos canarios en su inventario del Atlántico NE. Desde entonces no ha sido publicada ninguna lista completa de decápodos que cubra específicamente esta área y es necesaria una actualización. La lista actual de braquiuros canarios consta de 132 especies. Otras especies han sido citadas debido a la intensificación de las investigaciones dirigidas a aguas profundas, expansiones naturales de su rango de distribución desde áreas cercanas, introducciones causadas por actividades antropogénicas y a descripción de nuevas especies; algunos de estos cambios se detallan en esta revisión. Aunque no se espera que la descripción de nuevos braquiuros se produzca a un ritmo significativo, es esperable un incremento en el número de especies en Canarias como resultado de muestreos con arrastre bentónico y dragas y de la introducción de especies exóticas. Por primera vez, se realizan consideraciones zoogeográficas sobre la carcinofauna de braquiuros de Canarias. Por último, las especies de interés comercial son listadas, sus amenazas actuales identificadas y se proponen algunas medidas de conservación. Palabras clave: lista; zoogeografía; conservación; Brachyura; Decapoda; Crustacea; Canarias; Atlántico oriental. Citation/Como citar este artículo: González J.A. 2016. Brachyuran crabs (Crustacea: Decapoda) from the Canary Islands (eastern Atlantic): checklist, zoogeographic considerations and conservation. Sci. Mar. 80(1): 89-102. doi: http://dx.doi. org/10.3989/scimar.04350.10A Editor: E. Macpherson. Received: September 28, 2015. Accepted: November 11, 2015. Published: January 20, 2016. Copyright: © 2016 CSIC. This is an open-access article distributed under the Creative Commons Attribution-Non Commercial Lisence (by-nc) Spain 3.0. Scientia Marina 80(1) March 2016, 89-102, Barcelona (Spain) ISSN-L: 0214-8358 doi: http://dx.doi.org/10.3989/scimar.04350.10A INTRODUCTION Just 20 years have passed since González (1995) finished one of his seminal works on decapod crustaceans of the Canary Islands, thanks to the invaluable help of the reputed carcinologists L.B. Holthuis (see Holthuis 1995) and C.H.J.M. Fransen. This publication allowed d’Udekem d’Acoz (1999) to include the 90 • J.A. González SCI. MAR., 80(1), March 2016, 89-102. ISSN-L 0214-8358 doi: http://dx.doi.org/10.3989/scimar.04350.10A Canarian decapods in his inventory and distribution of decapods in northeastern Atlantic north of 25°N. The Canary archipelago is an overseas Spanish territory and an outermost European Region placed in the eastern-central Atlantic. This archipelago is situated in front of the northwestern coast of Africa, fairly close to the continents of Europe and Africa (104 km from Cape Juby, Morocco) but separated from them by great depths (Fig. 1). The age of the islands varies from east to west between 19 my (Lanzarote) and 0.7 my (El Hierro). The volcanic characteristics of the Canary Islands are shown by the absence of wide insular shelves, with a bottom depth of 180-200 m near the coast. The archipelago has nearly 1300 km of coastline and is washed by the oligotrophic ocean (Braun and Molina 1984). The Canaries are under the influence of the subtropical gyre of the eastern-central Atlantic, which facilitates the transport of planktonic larvae and rafting organisms to the archipelago from the American, European and Northwest African coast. The average seawater temperature around the Canary Islands is 18.5°C in February, rising abruptly to 24°C in August–September (e.g. Barton et al. 1998). A mesoscale distribution of larval communities has recently been described in filaments of the upwelling system from the African coast reaching the southeast of the archipelago (e.g. Landeira et al. 2009, 2010). Also, the Canaries are geographically located on a very important maritime route, and both ships and oil platforms have been recognized as major vectors for the introduction of non-native species (e.g. González et al. 2012a, Triay-Portella et al. 2015). Geomorphological, geographical and oceanographic particularities of the Canaries could explain the great diversity in the biogeographic patterns of the biota inhabiting this area. These physical and biodiversity characteristics, together with the climatic condition of the Canary Islands—a temperate-subtropical area—compared with the surrounding region highlight the uniqueness of the Canary Islands and their oceanographic connectivity to the adjacent waters (e.g. González et al. 2012a). The sustainable use of marine resources and their conservation measures is a major goal on national and international policy agendas (e.g. Spalding et al. 2007). According to the comprehensive biogeographic system for classifying the oceans (for both coastal and shelf areas) proposed by Spalding et al. (2007), the Canary Islands are included in the Azores-Madeira-Canaries ecoregion (i.e. the Macaronesian archipelagos), within the Lusitanian province of the Temperate Northern Atlantic realm. After the work of González (1995) and the compilation by d’Udekem d’Acoz (1999), Fransen and Wirtz (1997) published on Canarian decapod crustaceans, and González et al. (1996, 2000) and Quiles et al. (2002) listed crab families in the Canary Islands. Moro et al. (2014) published a list of decapods from the Canary Islands based on both material examined and in situ sightings, illustrated with colour photographs. Moreover, several authors have published updated lists of brachyuran decapod fauna at different geographical scales (e.g. Türkay 2001, Ng et al. 2008, Marco-Herrero et al. 2015). However, none of these has specifically covered the diversity found around the Canary Islands; and an update is needed for this area. The systematic research landscape on decapod crustaceans has changed drastically in the last few decades. A great number of changes concerning the brachyuran crustacean species found around the Canary Islands have also taken place. These changes are due to systematic modifications, non-confirmed presence or newly recorded species for the area. Today’s most widely used classifications have all appeared after the work of Zariquiey Álvarez (1968), and there is a concerted effort by carcinologists worldwide to check the validity of taxa using multiple tools such as ecology, larvae and genetics (Marco-Herrero et al. 2015). The present work summarizes all changes in Canarian brachyuran carcinofauna since González (1995), provides scientists with an updated classification list, and makes for the first time a zoogeographic analysis of this important component of the marine biota of the Canary Islands. Moreover, crab species of commercial interest are listed, their current threats are identified, and some updated conservation measures are proposed. MATERIALS AND METHODS For the compilation of this list, all publications since 1995 about the distribution of brachyuran crabs were checked, including previous lists for the Canary Islands region (González 1995, González and Quiles 2003), data from Internet databases such as WoRMS (http://www.marinespecies.org/) and GBIF (http:// www.gbif.org/species), systematic data, new records, and unpublished or in-preparation data. Several contributions need to be highlighted here, particularly the impressive studies on European decapods (d’Udekem d’Acoz 1999, Türkay 2001), Iberian decapods (gathering many records from the Canaries, Zariquiey Álvarez 1968), and West African brachyuran crabs (Manning and Holthuis 1981), but also several specific works on brachyurans (Neumann 1996, Fransen and Wirtz 1997, Moro et al. 2014), particularly the accounts on Canarian species of several crab families (González et al. 1996, 2000, Quiles et al. 2002). Several studies on feeding habits of fishes in waters of the Canaries (Fanlo et al. 1993, 1996, Tuset et al. 1996, Dürr and González 2001, Moreno-López et al. 2002) have also been used as a source of brachyurans occurring in the area. Some projects (DGXIV/C/1 94/034, CAMARON, PESCPROF 1-3, REDECA, AQUACRU, MARPROF) have provided us with deep-water crabs from the Canaries, collected with a panoply of fishing gear. Finally, many vouchers from the ‘Museo de Ciencias Naturales de Tenerife (TFMC)’, ‘Museu d’Historia Natural do Funchal (MMF)’, and the study collection of the ‘Instituto Canario de Ciencias Marinas’ (ICCM, now transferred to the ‘Universidad de Las Palmas de Gran Canaria’) were checked. This checklist covers all marine brachyuran species present in the Canary Islands from the intertidal to deep water. As in previous regional studies on decapod crus- Brachyuran crabs from the Canaries • 91 SCI. MAR., 80(1), March 2016, 89-102. ISSN-L 0214-8358 doi: http://dx.doi.org/10.3989/scimar.04350.10A taceans (González 1995) or fishes (Brito et al. 2002), the study area is bounded by the 30°N and 27°N parallels, the 19°W meridian and, in the Canaries-Africa channel, the 13°W meridian. The depth is generally less than 1500 m. This area occupies a band of about 600 km from east to west and about 330 km from north to south. In the north and west the depth is greater than 4000 m and on the southern edge it is greater than 3500 m (Fig. 1). This updated systematic classification follows Ng et al. (2008), but also takes into account the latest changes in particular taxa (e.g. Marco-Herrero et al. (2013) for Majoidea; Schubart and Reuschel (2009) and Spiridonov et al. (2014) for Cancroidea and Portunoidea). Superfamilies are listed by systematic order following the Sections and Subsections as currently accepted, and by alphabetical order within them. Families, subfamilies, genera, species and subspecies are also listed by alphabetical order within their respective superfamilies. All changes with respect to the catalogue by González (1995) and the subsequent checklist by González and Quiles (2003) are explained, including systematic modifications, misidentifications, synonyms, species that reach Canarian waters by increasing their distribution range, invasive and introduced species in the Canary Islands, and new records from the Canaries. Each crab species was classified as pelagic or benthic according to the main spatial distribution of their adults. Moreover, according to depth boundaries found around the Canary Islands (closely linked to regional oceanographic conditions determined by the water masses present) (Pajuelo et al. 2015), each benthic species was then assigned to a depth boundary where the species mainly occur in waters of the Canary Islands: shelf and transition area between shelf and slope (ST, 0-300 m), upper slope (US, 301-800 m), middle slope (MS, 801-2000 m) and lower slope (LS, >2000 m). Each crab species was also assigned to a biogeographic pattern category adapted from Brito et al. (2002) and subsequent studies (e.g. Espino et al. 2014). Recent biogeographic studies that clearly separate the coastal and shelf biota occurring in the Macaronesian ecoregion (i.e. the Azores, Madeira and the Canaries) from that occurring in the Cape Verde Islands (officially Cabo Verde) (Brito et al. 2007, Spalding et al. 2007, Brito 2010, Wirtz et al. 2013) were also taken into account. Canarian brachyuran crabs were classified into the following 11 groups according to their distribution range and the above-mentioned considerations: 1, cosmopolitan or worldwide species (COSM); 2, pantropical or circumtropical species (PANT); 3, amphi-Atlantic species of wide distribution (AAWD); 4, amphi-Atlantic species of warm affinity (AAWA); 5, eastern Atlantic species of wide distribution (EAWD); 6, eastern Atlantic cold-temperate species (EACT); 7, eastern Atlantic warm-temperate species (EAWT); 8, Atlanto-Mediterranean species (ATLM); 9, Guinean species (restricted to tropical and subtropical eastern Atlantic) (TSEA); 10, eastern-central Atlantic island species (from the Azores to Cape Verde Islands, and southwards even to St. Helena) (ECAI); and 11, Macaronesian species (around the Azores, Madeira, Savage, and/or the Canary Islands) (MAC). Two different zoogeographic approaches (consisting in describing the different components of the brachyuran fauna) were performed. A first description included all brachyuran species. For the second approach, both pelagic and deep-water benthic species (those living deeper than the transition area between shelf and slope, >300 m depth) were excluded from the analysis. It is widely accepted that the set of littoral and upper-bathyal species (in the Canaries, those living shallower than the upper slope, 0-300 m) better characterize the brachyuran carcinofauna of the Canary Islands and their zoogeographic affinities. The species’ occurrence around the neighbouring archipelago of Madeira (to the north) and the Cape Verde Islands (to the south) has also been recorded. Apart from publications containing well-documented first records, the main references used to check the occurrence of the brachyuran species were Manning Fig. 1. – Map of the Canary Islands (northeastern Atlantic) showing their geographic situation and bathymetric characteristics. 92 • J.A. González SCI. MAR., 80(1), March 2016, 89-102. ISSN-L 0214-8358 doi: http://dx.doi.org/10.3989/scimar.04350.10A and Holthuis (1981), Türkay (1982), Fransen (1991), d’Udekem d’Acoz (1999), Wirtz and d’Udekem d’Acoz (2001), Araújo and Calado (2003), González et al. (2004), and Araújo et al. (2009, 2014) among others. RESULTS A total of 132 marine crab species are reported herein around the Canary Islands (including three species of doubtful presence and another three of probable occurrence). Their spatial distribution, depth boundary, bathymetric range and biogeographic pattern are presented in Table 1. This represents a number of species close to the 140 brachyuran species reported around the Iberian Peninsula (Marco-Herrero et al. 2015), and nearly half of the 284 brachyuran species known in European waters (including the Azores, Madeira and the Canaries), with 40 freshwater crab species (d’Udekem d’Acoz 1999). When compared with the recent brachyuran catalogue from the Iberian Peninsula (MarcoHerrero et al. 2015), it is also noteworthy that both the Iberian and the Canarian lists comprise 20 currently accepted brachyuran superfamilies (Ng et al. 2008, Spiridonov et al. 2014). The Iberian superfamilies Cyclodorippoidea, Homolodromoidea and Corystoidea do not currently occur around the Canary archipelago. The superfamily Ocypodoidea has been reported from the Canaries based on two single findings of one species. The Macaronesian superfamily Pseudozioidea, as well as the African subtropical-tropical superfamilies Trapezioidea and Cryptochiroidea are represented in Canary Island waters, but not around the Iberian Peninsula to date (Marco-Herrero et al. 2015). No freshwater crab species occur in the Canaries. REMARKS Systematic and nominal changes since 1995 Systematic changes have affected the taxonomical arrangement of Brachyura by Zariquiey Álvarez (1968) and Manning and Holthuis (1981), which were followed in the Canarian catalogue by González (1995). The main changes in the systematics of brachyuran crabs after those impressive catalogues were recently explained by Marco-Herrero et al. (2015). So, instead of the 20 crab families and 67 genera considered in González (1995), a total of 39 brachyuran families and 77 genera are presented here. Several currently accepted families were included within other families in González (1995), as follows: Ethusidae within the Dorippidae; Eriphiidae, Oziidae, Pilumnidae, Pseudoziidae, Domeciidae, and Panopeidae within the Xanthidae; Euryplacidae within the Goneplacidae; Progeryonidae within the Geryonidae; Epialtidae and Inachidae within the Majidae; Carcinidae and Polybiidae within the Portunidae; Percnidae, Plagusiidae, and Varunidae as subfamilies within the Grapsidae. The family Cryptochiridae was reported as Hapalocarcinidae in González (1995). Ocypodidae (Castro 2012) and Varunidae (Moro et al. 2014) were reported based on very few findings of few individuals. The western Atlantic family Mathildellidae (Fransen and González in prep.) and the Thiidae (Monterroso and González in prep.) are newly reported from the Canaries here. The current account of brachyuran crabs of the Canary Islands adds another 22 to the 108 valid species in González (1995). According to d’Udekem d’Acoz (1999), Homologenus rostratus (A. Milne-Edwards, 1880) (Bouvier 1922) is a misidentification with Homologenus boucheti Guinot and Richer de Forges, 1995. Maja crispata Risso, 1827 (Herrera et al. 1993, as M. crisperata) should not be considered here because it is a misidentification with Pisa tetraodon. Macropodia aegyptia (H. Milne-Edwards, 1834) should be considered a synonym of Macropodia deflexa Forest, 1978. Portunus sp. in González (1995) should be assigned to Laleonectes vocans (A. Milne-Edwards, 1878). According to Reuschel and Schubart (2006), Xantho incisus Leach, 1814 is considered here a synonym of X. hydrophilus (Herbst, 1790), and the erroneous citation of the Indo-Pacific crab Lophozozymus incisus (H. Milne-Edwards, 1834) by Moro et al. (2014) is based on a secondary homonym of Xantho incisus, the true record. According to Neumann (1996), records for Maja squinado (Herbst, 1788) from the Canaries, the type locality, should be reassigned to Maja brachydactyla Balss, 1922. Pinnotheres pinnotheres (Linnaeus, 1758) is now considered as a new combination, Nepinnotheres pinnotheres (Linnaeus, 1758), made by Manning (1993) when he described a new genus (Nepinnotheres) with several previously Pinnotheres species. After González (1995), Chaceon inglei Manning and Holthuis, 1989 was reported from the Canaries by Araújo et al. (2009) as a result of deep sea surveys with selective crab traps. Ebalia tuberculata Miers, 1881 was reported from both littoral (Quiles et al. 2002) and bathyal bottoms (Moro et al. 2014). Ebalia tumefacta (Montagu, 1808) was first recorded from the Canaries by Quiles et al. (2002). Some littoral crab species have recently been found by expert scuba divers and reported by Moro et al. (2014): Xaiva mcleayi (Barnard, 1947), Liocarcinus pusillus (Leach, 1816), and Pinnotheres pisum (Linnaeus, 1767). Other littoral species were reported based on dredging sampling: Macropodia linaresi Forest and Zariquiey Álvarez, 1964, Liocarcinus navigator (Herbst, 1794), and Thia scutellata (Fabricius, 1793) (Monterroso and González in prep.). Some additional species are now present in Canarian waters, probably due to natural range expansions from nearby eastern Atlantic or Mediterranean areas: Calappa galloides Stimpson, 1859 (González et al. 2000), Ebalia tuberculata Miers, 1881 (Quiles et al. 2002), Merocryptus boletifer A. Milne-Edwards and Bouvier, 1894, and Macropipus tuberculatus (Roux, 1830) (Moro et al. 2014). However, the finding of one specimen of Neopilumnoplax sp. in Canarian waters (Fransen and González in prep.) may represent a recent colonization from the subtropical western Atlantic, or reflect that the deep-water crabs Neopilumnoplax americana (Rathbun, 1898) or Neopilumnoplax gervaini Tavares and Guinot, 1996 are amphi-Atlantic species. Regarding the specimens of Brachynotus Brachyuran crabs from the Canaries • 93 SCI. MAR., 80(1), March 2016, 89-102. ISSN-L 0214-8358 doi: http://dx.doi.org/10.3989/scimar.04350.10A Table 1. – List of the brachyuran species occurring in waters of the Canary Islands. Spatial distribution, benthic habitat, bathymetric range (depth interval in m known at the Canaries and elsewhere), and biogeographic pattern are provided for each species. Their occurrence (X) or absence (-) around Madeira and/or the Cape Verde Islands is also provided. Species: * doubtful presence, ** probable occurrence. Depth boundaries: ST shelf and transition area between shelf and slope (0-300 m), US upper slope (301-800 m), MS middle slope (801-2000 m), LS lower slope (>2000 m). Species’ biogeographic pattern codes: COSM cosmopolitan or worldwide, PANT pantropical or circumtropical, AAWD amphi-Atlantic of wide distribution, AAWA amphi-Atlantic of warm affinity, EAWD eastern Atlantic of wide distribution, EACT eastern Atlantic cold-temperate, EAWT eastern Atlantic warm-temperate, ATLM Atlanto-Mediterranean, TSEA tropical and subtropical eastern Atlantic, ECAI eastern-central Atlantic island (from Azores to Cape Verde Is., and even to St. Helena, MAC Macaronesian (Azores, Madeira, Savage, and/or the Canaries). Taxa / Species Spatial Depth Bathymetric range Biogeographic Canarian species also in distribution boundary Canaries Elsewhere pattern Madeira Cape Verde BRACHYURA Linnaeus, 1758 PODOTREMATA Guinot, 1977 DROMIOIDEA de Haan, 1833 Dromiidae de Haan, 1833 Dromia marmorea Forest, 1974 Benthic ST 0-96 0-76 TSEA X X Dromia personata (Linnaeus, 1758) Benthic ST 2-201 0-100 EACT likely likely HOMOLOIDEA de Haan, 1839 Homolidae de Haan, 1839 Homola barbata (Fabricius, 1793) Benthic ST 40-324 2-637 AAWD X X Homologenus boucheti Guinot and Richer de Forges, 1995 Benthic MS 733-1575 738-2195 EACT X - Paromola cuvieri (Risso, 1816) Benthic US 120-860 10-1212 EAWD X X Latreilliidae Stimpson, 1858 Latreillia elegans Roux, 1830 Benthic ST 180-330 35-475 ATLM likely X EUBRACHYURA de Saint Laurent, 1980 HETEROTREMATA Guinot, 1977 CALAPPOIDEA de Haan, 1833 Calappidae de Haan, 1833 Calappa galloides Stimpson, 1859 Benthic ST 15-80 15-218 AAWA - X Calappa granulata (Linnaeus, 1758) Benthic ST 15-300 10-400 ATLM X X Cryptosoma cristatum Brullé, 1837 Benthic ST 2-89 2-75 ECAI X X CANCROIDEA Latreille, 1802 Atelecyclidae Ortmann, 1893 Atelecyclus rotundatus (Olivi, 1792) Benthic ST 65-278 0-795 EAWD - X Atelecyclus undecimdentatus (Herbst, 1783) Benthic ST 0-51 0-51 ATLM likely likely Cancridae Latreille, 1802 Cancer bellianus Johnson, 1861 Benthic US 120-871 37-700 EACT X - Cancer pagurus Linnaeus, 1758 Benthic ST 150-400 0-520 EACT - - DORIPPOIDEA MacLeay, 1838 Dorippidae MacLeay, 1838 Medorippe lanata (Linnaeus, 1767) Benthic ST 10-100 9-952 EAWT - - Ethusidae Guinot, 1977 Ethusa mascarone (Herbst, 1785) Benthic ST 3-80 5-100 ATLM - - Ethusa rosacea A. Milne-Edwards and Bouvier, 1897 Benthic ST 125-132 100-1013 TSEA - X ** Ethusina alba Filhol, 1884 Benthic LS ? 2800-4265 EACT likely X Ethusina talismani A. Milne-Edwards and Bouvier, 1897 Benthic LS 2050-20831892-2400 EACT likely - ERIPHIOIDEA MacLeay, 1838 Eriphiidae MacLeay, 1838 Eriphia verrucosa (Forskål, 1775) Benthic ST 0-6 0-6 ATLM X - Oziidae Dana, 1851 Eupilumnus africanus (A. Milne-Edwards, 1867) Benthic ST 0-20 0-35 TSEA X X GONEPLACOIDEA MacLeay, 1838 Euryplacidae Stimpson, 1871 * Machaerus atlanticus (Miers, 1881) Benthic ST ? 10-90 TSEA - - Goneplacidae MacLeay, 1838 Goneplax barnardi (Capart, 1951) Benthic US 500-570 200-590 TSEA - X Goneplax rhomboides (Linnaeus, 1758) Benthic ST 15-570 0-580 EAWT X X Mathildellidae Karasawa and Kato, 2003 Neopilumnoplax sp. Benthic ST 279-279 - AAWA - - Progeryonidae Števčić, 2005 Paragalene longicrura (Nardo, 1869) Benthic ST 130-160 20-30 ATLM X - LEUCOSIOIDEA Samouelle, 1819 Leucosiidae Samouelle, 1819 ** Ebalia affinis Miers, 1881 Benthic ST ? 4-140 TSEA X X Ebalia deshayesi Lucas, 1846 Benthic ST 2-80 5-100 ATLM X - Ebalia edwardsii Costa, 1838 Benthic ST 0-190 0-100 ATLM X - Ebalia fragifera Miers, 1881 Benthic ST ? ? MAC X - Ebalia nux A. Milne-Edwards, 1883 Benthic US 540-2983 80-2983 EACT likely X Ebalia tuberculata Miers, 1881 Benthic ST 40-300 12-110 TSEA - - Ebalia tuberosa (Pennant, 1777) Benthic ST 30-180 0-199 EACT X - Ebalia tumefacta (Montagu, 1808) Benthic ST 150-200 0-199 EACT - - Ilia nucleus (Linnaeus, 1758) Benthic ST 162-162 0.5-80 ATLM - - Ilia spinosa Miers, 1881 Benthic ST 0-107 5-132 TSEA - - Merocryptus boletifer A. Milne-Edwards and Bouvier, 1894 Benthic ST 100-150 40-629 ATLM X - MAJOIDEA Samouelle, 1819 Epialtidae MacLeay, 1838 Epialtinae MacLeay, 1838 Acanthonyx brevifrons A. Milne-Edwards, 1869 Benthic ST 0-7 0-110 ECAI X X Acanthonyx lunulatus (Risso, 1816) Benthic ST 0-15 0-90 EAWT X X 94 • J.A. González SCI. MAR., 80(1), March 2016, 89-102. ISSN-L 0214-8358 doi: http://dx.doi.org/10.3989/scimar.04350.10A Taxa / Species Spatial Depth Bathymetric range Biogeographic Canarian species also in distribution boundary Canaries Elsewhere pattern Madeira Cape Verde Pisinae Dana, 1851 Anamathia rissoana (Roux, 1828) Benthic US 100-500 100-730 ATLM X - Herbstia condyliata (Fabricius, 1787) Benthic ST 0-54 0-80 ATLM X - Herbstia rubra A. Milne-Edwards, 1869 Benthic ST 0-20 0-75 TSEA - X Micropisa ovata Stimpson, 1858 Benthic ST 0-3 0-110 TSEA - X Pisa armata (Latreille, 1803) Benthic ST 41-82 18-162 EAWT likely X Pisa carinimana Miers, 1879 Benthic ST 4-120 4-100 EAWT X - Pisa nodipes (Leach, 1815) Benthic ST 3-70 0-100 ATLM X X Pisa tetraodon (Pennant, 1777) Benthic ST 2-50 0-50 ATLM - - Rochinia carpenteri (Thomson, 1873) Benthic MS 500-1059 400-1340 EACT X - Inachidae MacLeay, 1838 Achaeus cranchii Leach, 1817 Benthic ST 0 - >20 0-70 ATLM X - Dorhynchus thomsoni Thomson, 1873 Benthic MS 570-1163 106-2080 COSM likely X Inachus aguiarii de Brito Capello, 1876 Benthic ST 55-110 20-100 ATLM X - Inachus dorsettensis (Pennant, 1777) Benthic US 540-540 0-749 EAWD - - Inachus grallator Manning and Holthuis, 1981 Benthic ST 60-125 36-325 TSEA - - Inachus nanus Manning and Holthuis, 1981 Benthic ST 45-80 29-118 TSEA - - Inachus phalangium (Fabricius, 1775) Benthic ST 0.5-35 0.5-160 EACT X - Inachus thoracicus Roux, 1830 Benthic ST 30-90 10-200 ATLM - - Macropodia deflexa Forest, 1978 Benthic ST 23-23 0-90 EACT - - Macropodia linaresi Forest and Zariquiey Álvarez, 1964 Benthic ST 5-50 3.5-140 ATLM - - Macropodia aff. hesperiae Manning and Holthuis, 1981 Benthic MS 821-821 - MAC - - Macropodia longirostris (Fabricius, 1775) Benthic ST 30-60 4-130 ATLM - - Macropodia aff. longirostris (Fabricius, 1775) Benthic ST 50-60 - MAC - - Macropodia aff. parva van Noort and Adema, 1985 Benthic ST 25-82 20-90 TSEA - - Macropodia rostrata (Linnaeus, 1761) Benthic ST 0-100 0-193 EAWD likely - Macropodia spec. 2 Fransen, 1991 Benthic ST 86-200 140-170 MAC - - Stenorhynchus lanceolatus (Brullé, 1837) Benthic ST 2-273 5-96 TSEA X X Majidae Samouelle, 1819 Eurynome aspera (Pennant, 1777) Benthic ST 10-200 10-1216 EAWD X X Maja brachydactyla Balss, 1922 Benthic ST 0-72 0-91 EAWT X - Maja goltziana d’Oliveira, 1888 Benthic ST 50-287 27-250 ATLM - - Oregoniidae Garth, 1958 Ergasticus clouei A. Milne-Edwards, 1882 Benthic US 420-570 70-1000 ATLM X X PALICOIDEA Bouvier, 1898 Palicidae Bouvier, 1898 Palicus caronii (Roux, 1828) Benthic ST 20-220 18-220 ATLM X X PARTHENOPOIDEA MacLeay, 1838 Parthenopidae MacLeay, 1838 Distolambrus maltzami (Miers, 1881) Benthic ST 45-125 22-550 EAWT likely X Parthenopoides massena (Roux, 1830) Benthic ST 25-90 3-141 ATLM X X Spinolambrus macrochelos (Herbst, 1790) Benthic ST 100-475 5-1478 ATLM likely X Velolambrus expansus (Miers, 1879) Benthic ST 2-125 30-170 ATLM X X PILUMNOIDEA Samouelle, 1819 Pilumnidae Samouelle, 1819 Pilumnus hirtellus (Linnaeus, 1761) Benthic ST 0-25 0-90 EACT X X Pilumnus inermis A. Milne-Edwards and Bouvier, 1894 Benthic ST 200-250 5-400 ATLM X X Pilumnus spinifer H. Milne-Edwards, 1834 Benthic ST 0-20 1-179 ATLM X X Pilumnus villosissimus (Rafinesque, 1814) Benthic ST 0-20 0-20 ATLM X - PORTUNOIDEA Rafinesque, 1815 Carcinidae MacLeay, 1838 * Carcinus aestuarii Nardo, 1847 Benthic ST 10-10 1-26 COSM - - * Carcinus maenas (Linnaeus, 1758) Benthic ST ? 0-60 EACT - - Portumnus latipes (Pennant, 1777) Benthic ST 0-3 0-30 ATLM likely - Xaiva mcleayi (Barnard, 1947) Benthic ST 4-4 8-73 TSEA - - Geryonidae Colosi, 1923 Chaceon affinis (A. Milne-Edwards and Bouvier, 1894) Benthic US 411-1350 130-2047 EACT X X Chaceon inglei Manning and Holthuis, 1989 Benthic LS 2156-21561640-2500 EACT X - Chaceon maritae (Manning and Holthuis, 1981) Benthic US ? 100-1000 TSEA - X Geryon trispinosus (Herbst, 1803) Benthic US 639-833 32-2220 EACT - - Pirimelidae Alcock, 1899 Pirimela denticulata (Montagu, 1808) Benthic ST 0-15 0-250 EACT X X Polybiidae Ortmann, 1893 Bathynectes longipes (Risso, 1816) Benthic ST 30-100 15-226 ATLM X - Bathynectes maravigna (Prestandrea, 1839) Benthic US 366-846 60-1410 EACT - - Liocarcinus corrugatus (Pennant, 1777) Benthic ST 1-225 1-147 EAWT X X Liocarcinus depurator (Linnaeus, 1758) Benthic ST 45-90 1-871 EACT - - Liocarcinus holsatus (Fabricius, 1798) Benthic ST <100 1-400 EACT likely - Liocarcinus navigator (Herbst, 1794) Benthic ST 5-50 0-108 EACT - - Liocarcinus pusillus (Leach, 1816) Benthic ST 20-30 0-455 EACT likely - Liocarcinus vernalis (Risso, 1816) Benthic ST 35-100 0-150 ATLM - - Liocarcinus zariquieyi Gordon, 1968 Benthic ST 23-80 5-80 ATLM - - Macropipus tuberculatus (Roux, 1830) Benthic ST 100-150 48-748 EACT likely - Polybius henslowii Leach, 1820 Pelagic - 0-5 2-1245 EACT likely - Portunidae Rafinesque, 1815 Portuninae Rafinesque, 1815 Laleonectes vocans (A. Milne-Edwards, 1878) Benthic ST 5-10 6-37 AAWA X - Brachyuran crabs from the Canaries • 95 SCI. MAR., 80(1), March 2016, 89-102. ISSN-L 0214-8358 doi: http://dx.doi.org/10.3989/scimar.04350.10A sexdentatus (Risso, 1827) found in a brackish littoral pond with mobile sandy substrate (Charca de Maspalomas, S of Gran Canaria), under stones at 1.5 m of depth (Moro et al. 2014), given that this species has been introduced in other zones (for instance, the U.K.) and that even the habitat observed is not the more appropriate for this species, the most plausible reason for that finding is a human-mediated activity (through ballast water, fouling or oil platform, or a release from aquarium or pet trade). The finding in 2011 of one single male of Uca (Afruca) tangeri (Eydoux, 1835) on a sandstone platform on a crowded urban beach (Las Canteras, Gran Canaria) could be a consequence of larvae drifting transport between the African coast and the Canary archipelago, and the subsequent process of settlement and growing of an isolated individual (Castro 2012). Another plausible explanation is a human-mediated introduction (release from aquarium or pet trade). Acanthonyx brevifrons A. Milne-Edwards, 1869 is first recorded from the Canary Islands herein, based on one male (ICCM408, 3.8 mm carapace length) collected by hand at the Bay of Melenara, Gran Canaria, 27°59’N 15°12’W, Dec. 1997, on red algae Gelidium, 0-7 m. An eastern Atlantic species, appearing to be endemic from the NE Atlantic archipelagos of Azores Taxa / Species Spatial Depth Bathymetric range Biogeographic Canarian species also in distribution boundary Canaries Elsewhere pattern Madeira Cape Verde Portunus (Portunus) hastatus (Linnaeus, 1767) Benthic ST 2-60 2-40 TSEA X X Portunus (Portunus) inaequalis (Miers, 1881) Benthic ST ? 4-73 TSEA X X Thalamitinae Paul’son, 1875 Thalamita poissonii (Audouin, 1826) Benthic ST 0.5-120 0.5-20 PANT - X Thiidae Dana, 1852 Thia scutellata (Fabricius, 1793) Benthic ST 5-50 0-110 EACT - - PSEUDOZIOIDEA MacLeay, 1838 Pseudoziidae MacLeay, 1838 Euryozius bouvieri (A. Milne-Edwards, 1869) Benthic ST 0-23 6-30 ECAI X X TRAPEZIOIDEA Miers, 1886 Domeciidae Ortmann, 1893 Domecia acanthophora africana Guinot, 1964 Benthic ST 0-0 0-35 ECAI - X XANTHOIDEA MacLeay, 1838 Panopeidae Ortmann, 1893 Panopeus africanus A. Milne-Edwards, 1867 Benthic ST 0-8 0-140 TSEA - X Xanthidae MacLeay, 1838 Actaeinae Alcock, 1898 Paractaea monodi Guinot, 1969 Benthic ST 5-82 0-200 ATLM X X Paractaea rufopunctata (H. Milne-Edwards, 1834) Benthic ST 0-45 0-91 PANT - - Euxanthinae Alcock, 1898 Glyptoxanthus cavernosus (A. Milne-Edwards, 1878) Benthic ST 0-0 0-17 ECAI - X Monodaeus couchii (Couch, 1851) Benthic ST 20-500 0-1415 EAWT likely X ** Monodaeus rouxi (Capart, 1951) Benthic ST ? 11-510 TSEA X - Xanthinae MacLeay, 1838 Coralliope parvula (A. Milne-Edwards, 1869) Benthic ST 0-33 0-355 TSEA - X Microcassiope minor (Dana, 1852) Benthic ST 0-20 0-220 AAWA X X Nanocassiope melanodactyla (A. Milne-Edwards, 1867) Benthic ST 5-110 5-225 TSEA X X Xantho hydrophilus (Herbst, 1790) Benthic ST 0-25 0-37 ATLM X X Xantho pilipes A. Milne-Edwards, 1867 Benthic ST 0-36 0-133 EAWD X - Xantho poressa (Olivi, 1792) Benthic ST 0-15 0-20 ATLM - - Xantho sexdentatus (Miers, 1881) Benthic ST 0-15 0-35 TSEA likely X Xantho sp. Fransen, 1991 Benthic ST 0-7 0-15 TSEA X X Zosiminae Alcock, 1898 Platypodiella picta (A. Milne-Edwards, 1869) Benthic ST 0-30 0-30 TSEA X - THORACOTREMATA Guinot, 1977 CRYPTOCHIROIDEA Paul’son, 1875 Cryptochiridae Paul’son, 1875 Detocarcinus balssi (Monod, 1956) Benthic ST 20-25 3-62 TSEA - - GRAPSOIDEA MacLeay, 1838 Grapsidae MacLeay, 1838 Grapsus adscensionis (Osbeck, 1765) Benthic ST 0-7 0-4 TSEA X X Pachygrapsus marmoratus (Fabricius, 1787) Benthic ST 0-6 0-20 ATLM X - Pachygrapsus maurus (Lucas, 1846) Benthic ST 0-6 0-6 ATLM X X Pachygrapsus transversus (Gibbes, 1850) Benthic ST 0-7 0-7 PANT X X Planes minutus (Linnaeus, 1758) Pelagic - 0-31 0-31 PANT X X Percnidae Števčić, 2005 Percnon gibbesi (H. Milne Edwards, 1853) Benthic ST 0-29 5-29 PANT X X Plagusiidae Dana, 1851 Euchirograpsus liguricus H. Milne-Edwards, 1853 Benthic ST 150-250 10-359 EAWT X X Plagusia depressa (Fabricius, 1775) Benthic ST 0-5 0-5 AAWA X X Varunidae H. Milne-Edwards, 1853 Brachynotus sexdentatus (Risso, 1827) Benthic ST 1.5-1.5 0-2 ATLM - - OCYPODOIDEA Rafinesque, 1815 Ocypodidae Rafinesque, 1815 Uca (Afruca) tangeri (Eydoux, 1835) Benthic ST 0-2 0-2 TSEA - X PINNOTHEROIDEA de Haan, 1833 Pinnotheridae de Haan, 1833 Nepinnotheres pinnotheres (Linnaeus, 1758) Benthic ST 2-25 6-250 ATLM - - Pinnotheres pisum (Linnaeus, 1767) Benthic ST 38-38 0-150 EACT - - 96 • J.A. González SCI. MAR., 80(1), March 2016, 89-102. ISSN-L 0214-8358 doi: http://dx.doi.org/10.3989/scimar.04350.10A (Milne-Edwards and Bouvier 1894), Madeira (Chapman and Santler 1955, as A. lunulatus var. brevifrons) and the Cape Verde Islands (Milne-Edwards 1869, type locality), associated with Cystoseira meadows or rocks with algae, from the intertidal to up more than 110 m of depth (d’Udekem d’Acoz 1999). Although Forest and Zariquiey Álvarez (1964) reviewed the Mediterranean species of Macropodia Leach, 1814 and their nomenclature, specific identification of the eastern-central Atlantic Macropodia has remained particularly difficult (Manning and Holthuis 1981, d’Udekem d’Acoz 1999). Eight Macropodia species not fully identified at specific level from the Canary-Cape Verde region were included in the Fransen (1991) account. Of them, four species were compiled in the d’Udekem d’Acoz (1999) inventory, with three of them (M. aff. hesperiae, M. aff. longirostris, and Macropodia spec. 2) potentially representing endemic forms from the Macaronesia. Species of doubtful presence in the Canary Islands Machaerus atlanticus (Miers, 1881) has only been recorded from the Canary Islands by Barquín-Diez and Carrillo (1988) (as Pilumnoplax atlantica, with no collecting data). In the Eastern Atlantic, it is known from Senegal to Congo-Brazzaville, on mixed bottoms dominated by mud, 10-90 m (d’Udekem d’Acoz 1999). From the time when Santaella (1973) conducted his research, confusion has long existed over the validity of records for the species of Carcinus from the Canaries. Unfortunately, neither González (1995) nor this work have shed too much light on this issue. Carcinus maenas (Linnaeus, 1758) was first recorded from the Canaries by Heller (1863) (with no location), and then compiled by García Cabrera (1971) (“Canaries”). Santaella (1973) did not give validity to these citations based on the absence of a well-documented finding, and also the fact that the European species of Carcinus had not yet been separated in the 19th century. Although live specimens of this species were recorded as commercialized bait (Núñez et al. 2011), they were imported. The species’ supposed natural range of distribution covered the eastern Atlantic, from SE Iceland to Atlantic Morocco, the Western Sahara and Mauritania, and it is also known from the western Mediterranean. However, it has been introduced in the NE Pacific, South Africa, S of Australia, and perhaps in the NW Atlantic and has been accidentally observed in many locations around the world (d’Udekem d’Acoz 1999). It is found in intertidal and shallow subtidal zones, 0-60 m, mainly at 0-6 m, rarely to 200 m (?), under rocks and algae, tide pools, marshes and seagrass beds, and in low to full salinity areas (d’Udekem d’Acoz 1999, Fransen 2015a in WoRMS). Carcinus aestuarii Nardo, 1847 was probably first recorded from the Canaries by Heller (1863) (as C. maenas) and Almaça (1960), and then compiled by Zariquiey Álvarez (1968) (as C. mediterraneus), García Cabrera (1971) (as C. maenas), and Manning and Holthuis (1981) among others. The presence of this species in waters of the Canaries was corroborated by Barquín-Diez et al. (1982-1983) (one spec., Tenerife, muddy sand, 10 m), and collected again on polluted bottoms of the Port of Santa Cruz de Tenerife (González 1995, based on a pers. comm.). Were both specimens well identified? If correctly identified, were both specimens introduced? It has been recorded in scattered localities worldwide: the West Pacific (Japan, introduced, d’Udekem d’Acoz 1999), the Indo-West Pacific, the Indian Ocean (Suez Canal, ?Red Sea), the western Atlantic (USA). It has been found in the eastern Atlantic, the Mediterranean and Black Seas, in Morocco, Mauritania, and South Africa (Fransen 2015b in WoRMS), on muddy sand, under rocks, and seagrass meadows, at 1-26 m (Abelló et al. 1988, d’Udekem d’Acoz 1999). If C. aestuarii is the right identification for the species occurring in the Canaries, it should be considered as introduced. If C. maenas occurs in the Canaries, this would represents a range extension of this west-African species. Species probably occurring in the Canary Islands Ethusina alba Filhol, 1884: an eastern Atlantic species, found between France and the Azores, off Azores and off Cape Verde Islands, on mud bottoms often with pumice stone, 2800-4265 m (Manning and Holthuis 1981, Fransen 1991). Ebalia affinis Miers, 1881: an eastern Atlantic species, known from Seine Bank, N of Madeira (Doflein 1904), the Cape Verde Islands (Monod 1956) and from Senegal to Angola, including São Tomé and Principe Islands (Manning and Holthuis, 1981), on calcareous algae and mud with shells, 4-140 m (d’Udekem d’Acoz 1999). Portunus (Portunus) inaequalis (Miers, 1881): an eastern Atlantic species, known from Madeira (Türkay 1976), the Cape Verde Islands and Guinea-Bissau southward to Angola, including the islands of the Gulf of Guinea, on various bottom types, 4-73 m, mainly at 4-40 m (Manning and Holthuis 1981, d’Udekem d’Acoz 1999). To date no records for these three crab species exist from the Canaries. However, as they have been recorded from northern and southern adjacent areas, their presence in the Canary Islands is very probable. Very recently, photographic evidence was provided to me proving the occurrence of large specimens of P. (P.) inaequalis in waters of both Tenerife and Gran Canaria islands. Monodaeus rouxi (Capart, 1951): an eastern Atlantic species, found in Madeira (Fransen 1991), Senegal, Gambia and southward to Angola (d’Udekem d’Acoz 1999), mainly on mud, also on sand and clay (Forest and Guinot 1966, Fransen 1991), from 11 m (Manning and Holthuis 1981) to 510 m depth (Fransen 1991). No record exists for this species from the Canaries. However, it has been recorded from Madeira and the neighbouring African coast, so its presence in the Canary Islands is very probable. Cancer pagurus Linnaeus, 1758: an eastern Atlantic species, known from Norway (70°N) to S Portugal, including the Mediterranean (very rare), rocks and sand, from the intertidal to 520 m of depth, rarely down to 100 m (d’Udekem d’Acoz 1999). Some doubtful findings from Canada, USA and the Pacific Ocean Brachyuran crabs from the Canaries • 97 SCI. MAR., 80(1), March 2016, 89-102. ISSN-L 0214-8358 doi: http://dx.doi.org/10.3989/scimar.04350.10A have been reported in global databases. Macaronesia: Azores (Borges et al. 2010). Up to three fishermen have reported to us on sporadic catches of ox crab (= ‘buey liso’ in Canarian Spanish), few individuals per trap, off Fuerteventura. Some of these catches were sold in local supermarkets, and a dried carapace used as an ornamental object was examined by us. So the northeastern sector of the Canaries seems to be the southern distribution limit for the species, coinciding with the coldest waters around the archipelago. Spatial and vertical distribution Of the 132 brachyuran species occurring around the Canary Islands, 130 are benthic forms (Table 1). Polybius henslowii has been reported as epito bathypelagic, with benthic stages (e.g. González-Gurriarán 1987). Planes minutus is epipelagic, living on loggerhead sea turtles and inanimate flotsam (e.g. Dellinger et al. 1997), but also has benthic stages when flotsam reaches the beaches. Dorhynchus thomsoni is known to be benthic with a pelagic initial stage (e.g. Hartnoll et al. 1987). Portunus (Portunus) hastatus has juveniles with pelagic stages, but adults are clearly benthic (Table 1). Of the 130 brachyuran benthic species occurring in the Canaries, 112 are forms of the shelf and transition area between shelf and slope (ST, 86.2%, mainly inhabiting at a depth interval of 0-300 m), 11 are upper slope species (US, 8.5%, mainly at a depth interval of 301-800 m), 4 are middle-slope species (MS, 3.1%, mainly at a depth interval of 801-2000 m), and 3 are lower slope species (LS, 2.3%, mainly at a depth greater than 2000 m) (Table 1). Seven brachyuran benthic species seem to occur in waters of the Canary Islands at the shallowest depth ever recorded elsewhere: Homologenus boucheti, Ilia spinosa, Velolambrus expansus, Xaiva mcleayi, Laleonectes vocans, Euryozius bouvieri, and Percnon gibbesi. Seventeen brachyuran benthic species seem to occur in waters of the Canary Islands at the greatest depth ever recorded elsewhere: Dromia marmorea, D. personata, Cryptosoma cristatum, Cancer bellianus, Paragalene longicrura, Ebalia edwardsii, E. tuberculata, E. tumefacta, Ilia nucleus, Pisa carinimana, Inachus aguiarii, Stenorhynchus lanceolatus, Maja goltziana, Liocarcinus corrugatus, L. zariquieyi, Portunus (Portunus) hastatus and Thalamita poissonii. Zoogeographic considerations To describe for the first time the different components of the brachyuran fauna, all the Canarian brachyuran crabs listed herein (132 species) are grouped by biogeographic pattern in Table 2. A first preliminary description showed three main groups: 39 ATLM species (29.5%), 28 TSEA species (21.2%), and 27 EACT species (20.5%). The remaining eight biogeographic groups varied from six EAWD species (4.5%) to one AAWD species (0.8%) (Table 2). The presence of one MAC endemic species, Macropodia aff. hesperiae Manning and Holthuis, 1981, is noteworthy. The only record for this middle-slope species from the Canary Islands was by González (1995), based on one specimen caught off Playa de Santiago, S of La Gomera, 821 m, which was identified by C.H.J.M. Fransen, and then validated by d’Udekem d’Acoz (1999). Since all Macropodia species known to date were discarded during the identification, the Canarian specimen may represent a new species, even being an endemism from the Canary Islands or the Macaronesia. The true Macropodia hesperiae Manning and Holthuis, 1981 inhabits littoral waters at depths between 46 and 97 m from Senegal to Nigeria (Manning and Holthuis 1981). The Canarian brachyuran crabs inhabiting the shelf and transition area between shelf and slope (123 species), i.e. those living shallower than the upper slope, at 0-300 m, are grouped by biogeographic pattern in Table 2. This description of the benthic littoral and/or upperbathyal species showed the same three main groups: 39 ATLM species (31.7%), 28 TSEA species (22.8%), and 21 EACT species (17.1%). The remaining eight biogeographic groups varied from ten EAWT species (8.1%) to one AAWD species (0.8%) (Table 2). The presence of four ECAI species is noteworthy. 1. Cryptosoma cristatum Brullé, 1837: known from the Azores (Wirtz and Martins 1993), Porto Santo, Madeira, Desertas Islands (Türkay 1976, as Crytosoma cristatum, erroneous spelling), the Canaries (Brullé 1837-1839, as C. cristata, type locality), the Cape Verde Islands (Monod 1956), and St. Helena Island; as an introduced species, found in the southwestern Mediterranean, with one record from the Alboran Sea (García-Raso 1993). 2. Acanthonyx brevifrons A. Milne-Edwards, 1869: from the Azores (Milne-Edwards and Bouvier 1894), Madeira (Chapman and Santler 1955, as A. lunulatus var. Table 2. – Zoogeographic composition of the Canarian brachyuran fauna showing the number of species (N) grouped by biogeographic pattern. Species’ biogeographic pattern All Canarian brachyuran (132 species, depth 0 - >2000 m) Littoral and upper bathyal benthic (123 species, depth 0-300 m) N%N% Atlanto-Mediterranean (ATLM) 39 29.5 39 31.7 Tropical and subtropical Eastern Atlantic (TSEA) 28 21.2 28 22.8 Eastern Atlantic cold-temperate (EACT) 27 20.5 21 17.1 Eastern Atlantic warm-temperate (EAWT) 10 7.6 10 8.1 Eastern Atlantic of wide distribution (EAWD) 6 4.5 6 4.9 Amphi-Atlantic of warm affinity (AAWA) 5 3.8 5 4.1 Eastern-central Atlantic island (ECAI) 5 3.8 5 4.1 Pantropical or circumtropical (PANT) 5 3.8 4 3.3 Macaronesian (MAC) 4 3.0 3 2.4 Cosmopolitan or worldwide (COSM) 2 1.5 1 0.8 Amphi-Atlantic of wide distribution (AAWD) 1 0.8 1 0.8