Updating changes in the Iberian decapod crustacean fauna (excluding crabs) after 50 years

Abstract

An annotated checklist of the marine decapod crustaceans (excluding crabs) of the Iberian Peninsula has been compiled 50 years after the publication of "Crustáceos decápodos ibéricos” by Zariquiey Álvarez (1968). A total of 293 species belonging to 136 genera and 48 families has been recorded. This information increases by 116 species the total number reported by Zariquiey Álvarez in his posthumous work. The families with the greatest species richness are the Paguridae (28) and Palaemonidae (18). References by geographic sectors and for all species are given. The results show that 264 species are reported in the Atlantic sectors, while 178 have been found in the Mediterranean. The species richness and the differences between and within sectors are discussed; these are mainly due to the dimension of the areas, the depth ranges and the confluence of distinct water masses with a different origin and different physicochemical features. Consequently, the greatest richness of decapod species (excluding crabs) is found in the Gulf of Cádiz, with 194 species. The total number of decapods found in and around Iberian waters, including crabs, freshwater species and some new records not yet published, reaches 448.

INTRODUCTION

The study of decapod crustaceans in the Iberian Peninsula began in the last part of the 19th century, with information and lists obtained from different coastal areas (e.g. Brito Capello 1873, Barceló-Combis 1875, Bolívar 1892, 1916, Buen 1887, 1916, Ferrer Galdiano 1920, 1940, Miranda y Rivera 1921), as well as from the results from the research expeditions that covered parts of these waters (see García Raso 1996 and references cited therein). The first checklist for all Spanish waters was compiled by Miranda y Rivera (1933) and subsequently extended by Zariquiey-Cenarro (1935a, b, 1942). Zariquiey Álvarez (1946) published a synthesis, “Crustáceos Decápodos Mediterráneos”. In Portugal, Nobre (1931, 1936) published “Crustáceos Decápodos e Stomatópodes marinhos de Portugal”. These relatively intensive works on the Iberian decapods culminated just 50 years ago with the publication of “Crustáceos Decápodos Ibéricos”, a posthumous work by Ricardo Zariquiey Álvarez completed by Drs Lipke B. Holthuis, Jacques Forest and Isabella Gordon, which has been, and still is, a basic reference work for this taxonomic group in the Mediterranean Sea and Europe.

Recently, the works of d’Udekem d’Acoz (1999a) and Türkay (2001) offer updated lists of the European decapods, but they do not provide listings of species by geographic sectors.

Today, the number of known Iberian decapod crustaceans has increased mainly due to the large number of studies undertaken along the continental shelf and slope, as well as on bathyal bottoms. Furthermore, climate change and other anthropogenic activities have increased the number of thermophilic and invasive species, reinforcing the utility of check-lists for detecting changes in fauna over time in specific regional areas (Zenetos et al 2010, 2012). Finally, changes in nomenclature and taxonomical reorganizations at different levels (families, genera and species) have taken place in these years due, in part, to the development of molecular techniques (Cuesta et al. 2012, among others) and to larval studies (Marco-Herrero et al. 2013).

These reasons led us to consider that it would be of wider interest to review the list of Iberian decapod crustacean species. We also present this review in homage to and recognition of all these authors, especially Dr. Ricardo Zariquiey Álvarez and his father Ricardo Zariquiey-Cenarro. In this regard, Marco-Herrero et al. (2015) published a first paper with the updated list of the brachyuran crabs, and now, with this present paper, which includes the remaining marine groups, we update the definitive list of all marine Iberian decapod crustaceans. In addition, a list of freshwater species is given to account for the total number of species in Iberian waters.

MATERIALS AND METHODS

In order to draw up the present updated list of Iberian decapod crustaceans (excluding Brachyura), two basic studies were taken as a departure point: Zariquiey Álvarez (1968) and d’Udekem d’Acoz (1999a), in addition to a large number of subsequent specific publications. Data from relatively recent expeditions in remote areas of the coast, especially those from deep and pelagic waters (e.g. Türkay 1975, García Raso 1996, Fariña et al. 1997, and Cartes et al. 2007, 2014) enabled us to complete the species list. Additional information was obtained from several expeditions and research projects carried out by governmental agencies, which have published information and species lists on the internet. These include the Instituto Español de Oceanografía projects ECOMARG (http://www.ecomarg.com/biodiversidad.html), DEEPER (http://www.ma.ieo.es/deeper) and INDEMARES (http://www.indemares.es), the Alcalá University CALMEN07 Expedition (http://www.juanjunoy.info/wp-content/uploads/2012/12/CALMEN07.pdf; Junoy 2008) and studies of specific areas (e.g. Calado and Narciso 2002). Reviews of specific groups and genera (e.g. Macpherson 1988, García-Gómez 1994, Galil 2000, and Macpherson and Segonzac 2005), those with descriptions of new species (e.g. García Muñoz et al. 2014), and new records of introduced species in the area (e.g. García Raso et al. 2014b) were also included.

In addition, in studies on meso- and bathypelagic species in waters some distance from the Iberian Peninsula coast, for example between Portugal and Azores, some species are mentioned but their specific location is not (e.g. Burukovsky 1996). Some of these species, which show wide geographic distribution and extensive vertical movements, have also been included, because although they have not been captured in jurisdictional waters (economic zone) their presence is highly probable, especially considering the scarcity of studies in the pelagic habitat around Iberian waters.

The geographical sectors considered are the same as in Marco-Herrero et al. (2015) (Fig. 1): the Gulf of Biscay and Galicia, western Portugal, the Gulf of Cádiz and the southwestern Iberian Peninsula, the Alboran Sea, and the Spanish western Mediterranean. For each of these sectors and for all species, some selected bibliographic references are given.

[Figure omitted. Please see PDF.]

For the classification and nomenclature, we followed that adopted in WoRMS (http://www.marinespecies.org/) and in recent reviews (Vereshchaka 2000, 2009, Baba et al. 2008, McLaughlin et al. 2010, Osawa and McLaughlin 2010, Boyko and McLaughlin 2010, Chan 2010, De Grave and Fransen 2011, Vereshchaka et al. 2014, and Ng and De Forges 2015, among others).

RESULTS

A total of 293 species of marine and estuarine decapod crustaceans, excluding crabs (Brachyura) were recorded in coastal and offshore waters around the Iberian Peninsula (Appendix 1). They belong to 136 genera and 48 families. These data increase the number of species recorded by Zariquiey Álvarez (1968) by 116 species of 54 different genera and 22 families (the last two, including reassignments).

The updated checklist by geographical sectors of all decapod crustacean species, excluding Brachyura, recorded in waters around the Iberian Peninsula is shown in Appendix 1. The distribution of some species shows sectors in white, which are absences or “gaps” interspersed among others in which the species is cited. Unless they are at the distribution limit of the species, their presence in these gaps (although not verified) is more than likely.

The families with the highest species richness are Paguridae with 28 species, followed by Palaemonidae with 18 (excluding freshwater species) (Appendix 1).

Table 1 presents the taxonomic changes in the names of genera and species referred to in Zariquiey Álvarez (1968), as well as some synonyms mentioned in recent studies.

Table 1. – Changes in the generic or authority assignation of species in comparison with Zariquiey Álvarez (1968) as well as synonymized species, listed by taxonomic hierarchy. * According Garcia Gómez (1994) and WoRMS.

[Table omitted. Please see PDF.]

Taxonomic groups

Zariquiey Álvarez (1968) considered two supersections: Natantia with three sections, Caridea, Penaeidea and Stenopodidea; and Reptantia with three sections, Macrura Reptantia, Anomura and Brachyura. However, in the current classification these taxa belong to different infraorders (included in two suborders), and the supersection and section ranges are not accepted.

The suborder Dendrobranchiata (previously Section Penaeidea within Supersection Natantia) is represented in the Iberian Peninsula by 38 species in 26 genera and 7 families.

The most important contributions to the knowledge of Dendrobranchiata in the study area are the papers of Maurin (1965), Abbes and Casanova (1973), Casanova and Judkins (1977), Sardà et al. (1982), Abelló et al. (2002), Domenech et al. (1981), Cartes et al. (2007, 2014), Serrano et al. (2011), Abelló and Torres (1998), and the classic works of Sund (1920), Stephensen (1923) and Hansen (1920, 1922).

In the new updated checklist, a new genus and species of the family Aristeidae is recorded for Iberian waters: Cerataspis monstrosus, which was captured in the Gulf of Biscay and reported as Plesiopenaeus armatus by Saint Laurent (1985). According to its general geographic distribution (Crosnier and Forest 1973, d’Udekem d’Acoz 1999a), this species must also be present in the Atlantic areas of Portugal and the Gulf of Cádiz. Morgan et al. (1985: Fig. 3) showed the presence of mysis larval stages of several species of Cerataspis around Iberian waters, while Bracken-Grissom et al. (2012) confirmed molecularly that the adult of P. armatus and the larvae of C. monstrosus belong to the same species.

In the family Benthesicymidae, two new species of the genus Benthesicymus have been added: B. brasiliensis and B. iridescens (Appendix 1).

In the family Penaeidae, three other species must be included in the updated list: Penaeopsis serrata, Funchalia villosa and the introduced species Penaeus pulchricaudatus (Abelló and Torres 1998, Abelló et al. 2002, Judkins 2014, d’Udekem d‘Acoz 1999a, Rodríguez 1989).

In the Solenoceridae family, Metapenaeus affinis, Hymenopenaeus aphoticus and Hymenopenaeus debilis have been added to the list provided by Zariquiey Álvarez (1968) (Cartes et al. 2014, García Raso 1996, Abelló et al. 2002).

The Sergestidae family has recently been revised by Vereshchaka (2000, 2009) and Vereshchaka et al. (2014). These authors proposed some new genera, although the WoRMs does not include some of them yet. The information on the general distribution of the species can be found in Vereshchaka (2009) and in papers mentioned in the following paragraph.

In Iberian waters, this family shows a high species richness. A total of 15 species in 10 genera, most of them epipelagic and some benthopelagic and bathypelagic, have been reported. Eight of these species were not reported by Zariquiey Álvarez (1968): Cornutosergestes cornutus (synonym: Sergestes cornutus Krøyer, 1855), Deosergestes henseni (syn. Sergestes henseni Ortmann, 1893), Gardinosergia splendens (syn. Sergia splendens Sund, 1920), Parasergestes armatus (syn. Sergestes armatus Krøyer, 1855), Petalidium obesum, Sergia burukovskii, Sergia grandis and Sergia tenuiremis (syn. Sergestes kroyeri Bate, 1891) (see Appendix 1) (Cartes et al. 2014, d’Udekem d‘Acoz 1999a, Torres et al. 2014, Landeira and Fransen 2012, Hansen 1920, 1922, Dos Santos and Moreira 2003).

Judkins (2014) provided geographical distribution maps of pelagic decapod shrimps in the Atlantic Ocean, which allowed us to complete the distribution of some species (such as Gennadas tinayrei and Funchalia villosa). We also used general information from the Ocean Biogeographic Information System (OBIS) (for example for F. villosa http://www.iobis.org/explore/#/taxon/444697) and from specific websites edited by recognized research centres or researchers (e.g. F. woodwardi reported in Portugal in the French Inventaire National du Patrimoine Naturel (https://inpn.mnhn.fr/accueil/recherche-de-donnees/especes).

Additionally, Deosergestes arachnipodus (Cocco, 1832, described as Acheles arachnipodus) was considered a synonym of Sergestes arcticus by Hansen (1922) and Zariquiey Álvarez (1968). Holthuis (1977a) mentioned that it is a synonym of S. corniculum Kroyer, 1885 and that the use of the name arachnipodus should be given preference since it is older than corniculus. However, Vereshchaka (2009), following Hansen (1922), considered that D. arachnipodus is a synonym of Sergestes arcticus Kroyer, 1855 (=Eusergestes arcticus, see Judkins and Kensley 2008), but this species is still accepted as valid in the WoRMS. We have followed Vereshchaka (2009) in this article, but the situation needs to be clarified.

Other species of Dendrobranchiata not yet reported in Iberian waters but with probable presence due to their biogeographical distribution range, and because they have been reported off northern Morocco, are Benthesicymus bartletti Smith, 1882 and Benthonectes filipes Smith, 1885 (García Raso 1996), as well as Allosergestes pectinatus (Sund, 1920) (Sund 1920, d’Udekem d‘Acoz 1999a).

The suborder Pleocyemata is the most diverse and rich in species. It includes ten infraorders: Achelata, Polychelida, Astacidea, Stenopodidea, Caridea, Procarididea (with no species in Iberian waters), Axiidea, Gebiidea (both previously included within the Thalassinidea infraorder), Anomura and Brachyura.

The first three infraorders (Achelata, Polychelida and Astacidea) have 18 marine species included in 14 genera. In Zariquiey Álvarez (1968) these taxa were included within the Section Macrura Reptantia in the superfamilies Astacidea and Palinuridea [together with the Polychelidae, Palinuridae and Scyllaridae (the last two belonging presently to the Achelata)].

Some relevant contributions for the study area concerning these infraorders include those of Bouvier (1917), Miranda y Rivera (1921), Silvertsen and Holthuis (1956), Türkay (1976), Domenech et al. (1981), Saint Laurent (1985), Neves (1987), García Raso (1996) and Cartes et al. (2014), as well as the review of Polychelidae by Galil (2000).

New additions to Achelata since Zariquiey Álvarez (1968) are the scyllarid Acantharctus posteli (=Scyllarus posteli Forest, 1963) (Pozuelo et al. 1976, García Raso 1982c, González-Gordillo and Rodríguez 2003) and the introduced palinurid Jasus lalandii (Guerra and Gaudêncio 1982, d’Udekem d‘Acoz 1999a).

Within the Polychelida, new additions include Cardus crucifer (=Polycheles crucifer Thomson, 1873) (d’Udekem d‘Acoz 1999a, Türkay 1976), Pentacheles snyderi (Galil 2000), and Willemoesia leptodactyla (Saint Laurent 1985). Pentacheles laevis was quoted in Zariquiey Álvarez (1968) as Polycheles granulatus Faxon, 1893) without specifying its distribution area, but d’Udekem d’Acoz (1999a) (following Bouvier 1917) mentioned the species in Galicia and Portugal. Pentacheles validus (=Polycheles validus A. Milne-Edwards, 1880) could very probably be found in Atlantic Iberian waters since it has been reported in the northern sector of the Gulf of Biscay (Saint Laurent 1985), the Azores and northern Africa (Galil 2000).

The infraorder Stenopodidea was represented in the book of Zariquiey Álvarez (1968) by only two species. The studies of Macpherson (1978), Pretus (1990), Cartes and Sardà (1992), Cartes et al. (1994), García Raso (1996), Goy and Cardoso (2014)and Cartes et al. (2014) have increased this list to seven species (in four genera and two families), most of them from deep waters. Odontozona addaia is an endemism of the Iberian Peninsula, described from a marine cave of Menorca (Balearic Islands) (Pretus 1990). Odontozona edwardsi (=Richardina edwardsi Bouvier, 1908) was captured in northern Spain (Goy and Cardoso 2014), Richardina fredericii in the western Mediterranean (Macpherson 1978, Cartes and Sardà 1992, Cartes 1993, Cartes et al. 1994) and Spongicoloides evolutus in southern Portugal (García Raso 1996). Other studies (e.g. Neves 1969, García Raso 1985a) have increased the distribution area of some species. In addition, Spongicoloides profundus Hansen, 1908 could be added to this list, since although there are no records for Iberian waters, it has been captured in northern Morocco (García Raso 1996) and its biogeographic range reaches northern Europe.

The infraorder Caridea is a very large taxonomic group, with 128 marine species in Iberian peninsular waters included in 51 genera and 14 families. Fifty of these species were not mentioned in 1968. The taxonomy/phylogeny of the Caridea is not yet fully resolved, as is the case of the Palaemonidae, which appear in two clades (Aznar-Cormano et al. 2015), although only one of them is in Iberian waters.

Within the Alpheidae family, Alpheus platydactylus was reported for the first time in Spanish waters by Forest (1965), Synalpheus africanus (as S. hululensis) by García Raso (1984a), Automate branchialis by Guillen Nieto (1996), and Alpheus talismani and Athanas amazone by García Raso (1996) and Marina et al. (2015). These species have subsequently been found in other peninsular areas (e.g. Box et al. 2007, García Raso et al. 2010, Duris 1996, Abelló et al. 2002) (Appendix 1). Recently two new records of Alpheus species (Palero et al. unpublished data) have been found in Iberian waters, thus increasing to 12 the number of marine species of this family. As potential additions, we can mention the African Athanas grimaldii Coutière, 1911, given its report in the Gulf of Biscay, France (Coutière 1911), although it needs confirmation (Nouvel 1941 in d’Udekem d‘Acoz 1999a).

De Grave et al. (2014) showed that the former family Hippolytidae is not a monophyletic group and proposed to split it into five families. We have followed this criterion. Therefore, this group at present contains the following families: Lysmatidae Dana, 1852, Thoridae Kingsley, 1879, Bythocarididae Christoffersen, 1987, Merguiidae Christoffersen, 1990 and Hippolytidae S. Bate, 1888 s.s. Four genera and 11 species belonging to three of these families have been added to Iberian peninsular waters since Zariquiey Álvarez (1968).

Within the Bythocarididae family, only the occurrence of Bythocaris cosmetops has been reported in the Alboran Sea (García Raso et al. 2011).

Six new records for the Hippolytidae family have been provided in Iberian waters. Caridion gordoni was reported on the Galicia Bank seamounts (Cartes et al. 2014) and Caridion steveni off Portugal (larvae) (Paula 1987), in the Balearic Islands (García Socias and Gracia 1996, as Caridion sp., Torres et al. 2014, larvae) and in the Ebro Delta area (Abelló et al. 2002). Merhippolyte ancistrota was captured off Morocco, both in the Alboran Sea and in the Atlantic Ocean (d’Udekem d’Acoz and Duriš 1996, García Raso 1996), and recently also in the Gulf of Cádiz (García Raso, unpublished data, INDEMARES-Chica Expedition). Hippolyte lagarderei was reported in Cádiz by López de la Rosa et al. (2002), Hippolyte niezabitowskii in Málaga by García Raso et al. (1998), Hippolyte leptometrae in the Gulf of Biscay (Gascogne) and in the Mediterranean (France and Turkey, d’Udekem d‘Acoz 1999a), and recently in the Alboran Sea (García Raso pers. comm.). García Raso and Manjón-Cabeza (1996) provided information on the expansion of the distribution area for other species. However, the presence of Hippolyte holthuisi in northern Spain (Anadón 1975) needs to be confirmed.

Within the Thoridae family Eualus drachi was captured in the Alboran Sea (Duris 1996, García Raso et al. 2014a), Eualus lebourae in the Strait of Gibraltar off Ceuta (García Raso 1996), and the boreal species Lebbeus microceros on the Galicia Bank (Cartes et al. 2014). Paula (1987) found larvae of Eualus gaimardii in southwestern Portugal during six months of an annual cycle.

The presence of three additional species in Iberian waters is very possible. These belong to the Lysmatidae family: Lysmata nilita Dohrn and Holthuis, 1950, known in the Azores, Canary Islands and Mediterranean Sea, see (d’Udekem d‘Acoz 1999a), Lysmata olavoi Fransen, 1991, reported in the Azores and Mediterranean Sea (d’Udekem d‘Acoz 1999a), and Lysmata uncicornis Holthuis and Maurin, 1952, captured in Atlantic waters of Morocco (Lagardère 1971).

The Ogyridae family is represented by only one species, Ogyrides rarispina, not mentioned by Zariquiey Álvarez (1968), recorded in Cádiz by Holthuis (1977b) and later by López de la Rosa et al. (2002), Cuesta et al. (2006) and González-Ortegón (2008).

The Bresiliidae family, not mentioned by Zariquiey Álvarez (1968), is represented in Iberian waters by Bresilia atlantica, and was caught recently on the Galicia Bank (Cartes et al. 2014).

The Crangonidae family has 15 species included in seven genera in the study area. Of these, five have been reported since 1968 in Iberian waters. Four of these are found exclusively in Atlantic waters: Sabinea hystrix and Metacrangon jacqueti in the Galicia Bank by Cartes et al. (2014); Parapontophilus abyssi in the Gulf of Biscay and in the Gulf of Cádiz by Saint Laurent (1985) and García Raso (1996), respectively; and Crangon allmani in the Bay of Cascais, Portugal by Santinho (2009), although only larvae of this species have been captured. Anadón (1975) provided information on the presence of Crangon crangon in Galicia. Pontophilus norvegicus was first mentioned in Spanish Mediterranean waters by Forest (1965), and has also been reported in the Atlantic by d’Udekem d’Acoz (1999a) and Cartes et al. (2007). Philocheras bispinosus, reported by Zariquiey Álvarez (1968), Neves (1975), and García Raso and Manjón-Cabeza (2002), is represented by two subspecies or species according to the WoRMS. The subspecies Philocheras bispinosus neglectus (Sars, 1883) has been only reported by García Raso and Manjón-Cabeza (2002)in the Alboran Sea, and by Santinho (2009) in Portugal. More data on the distribution of some species in Portugal are available in Paula (1987), Dos Santos (1999) and Dos Santos et al. (2008). In addition, the presence of the African species Parapontophilus talismani (Crosnier and Forest 1973) (=Pontophilus talismani) is possible because it has been recorded in the northern part of the Gulf of Biscay (Saint Laurent 1985).

The Glyphocrangonidae family (not mentioned in Zariquiey Álvarez 1968) is represented by three Atlantic species: Glyphocrangon atlantica, G. sculpta and G. longirostris, the first two captured in the Gulf of Biscay (Holthuis 1971and Saint Laurent 1985), and the third on the Galicia Bank (Cartes et al. 2014).

The Nematocarcinidae family is represented by three species, two of them cited since 1968: Nematocarcinus exilis in the Atlantic Ocean (Saint Laurent 1985, Cartes et al. 2014, García Raso 1996) and in the Mediterranean Sea (Abelló and Valladares 1988), and N. gracilipes only in Atlantic waters off western Portugal and in the Gulf of Cádiz (Neves 1982, García Raso 1996); the presence of N. ensifer is mentioned in Portugal (without location) by Cardoso and Serejo (2007).

The Acanthephyridae family, included in Ophlophoridae in Zariquiey Álvarez (1968), is represented by 14 species within seven genera. Five of these have been cited since 1968: Acanthephyra brevirostris in Galicia and Portugal (Crosnier and Forest 1973), Acanthephyra purpurea in Galicia and Portugal (d’Udekem d‘Acoz 1999a, Silvertsen and Holthuis 1956) and also in the Gulf of Cádiz (Portuguese and Spanish waters) (García Raso 1996, García Raso unpublished data, INDEMARES-Chica 0211), Heterogenys microphthalma (syn. Acanthephyra microphthalma Smith, 1885) in Portugal (Coutière 1911, see d’Udekem d‘Acoz 1999a), Hymenodora glacialis in the Gulf of Biscay (Saint Laurent 1985); and Kemphyra corallina in western Portugal (see d’Udekem d‘Acoz 1999a). Finally, Meningodora miccyla (Chace, 1940) could also be present in Iberian waters because it was reported between the Azores and Portugal by Burukovsky (1996), although without a specific locality.

The Oplophoridae family is represented by four species, in two genera; three of them not mentioned by Zariquiey Álvarez (1968). Oplophorus spinosus was captured in Galicia (Cartes et al. 2014), southern Portugal and Cádiz (García Raso 1996, and García Raso INDEMARES-Chica, unpublished data); Systellaspis braueri was found in the Cantabrian Sea by Saint Laurent (1985) and Crosnier and Forest (1973); and Systellaspis cristata has been reported from the Gulf of Biscay (Crosnier and Forest 1973) and in the west and south of Portugal (d’Udekem d‘Acoz 1999a).

The Gnathophyllidae family has only one species in Iberian Peninsula waters, Gnathophyllum elegans, with a Mediterranean distribution (Zariquiey Álvarez 1968, García Raso 1982b), although it could be found in the Gulf of Cádiz [see WoRMS: Gnathophyllum elegans (Risso, 1816), accessed at http://www.marinespecies.org/aphia.php?p=taxdetails&id=107608 on 2018-04-03].

The Bathypalaemonellidae family has not been recorded in Iberian waters, but Bathypalaemonella serratipalma Pequegnat, 1970 was caught in the central part of the Gulf of Cádiz (Ibero-Moroccan), in waters close to the coast of Morocco (García Raso 1996).

Concerning the Palaemonidae family, Grippa and d’Udekem d’Acoz (1996) reviewed the species belonging to the genus Periclimenes from the Mediterranean and the northeastern Atlantic. González-Ortegón and Cuesta (2006) made a very useful illustrated key to identify the species of Palaemon and Palaemonetes (=Palaemon, after De Grave and Ashelby 2013) from European waters. In total, 18 marine and estuarine species of palaemonids are known from peninsular waters. Four of them have been captured since 1968: Brachycarpus biunguiculatus in the Mediterranean (in the Alboran Sea by García Raso 1987a, 1990, and in the Balearic Islands by Torres et al. 2014); Periclimenes aegylios in the Balearic Islands (García Socias and Gracia 1996); P. kornii in the Gulf of Cádiz (Saint Laurent and García Raso 1993), the Alboran Sea (d’Udekem d’Acoz 1999b, and García Raso unpublished data) and, with doubts, in Catalan Sea waters (Cartes et al. 1994, as Periclimenes sp.). An introduced species, Palaemon macrodactylus, has also been reported in the Gulf of Cádiz, in the Guadiana, Guadalquivir, Guadalete, San Pedro and Salado estuaries (Cuesta et al. 2004, 2006, González-Ortegón et al. 2005), in Cascais, western Portugal (Santinho 2009) and in the Balearic Islands as larvae (Torres et al. 2012). Finally, González-Ortegón et al. (2006), Cuesta et al. (2012) and Cartaxana (2015) found, by analysing the variation in mtDNA, that there is no reason to recognize Palaemon garciacidi (a species described by Zariquiey Álvarez 1968) as a species distinct from P. longirostris. In addition, larvae of Ascidonia flavomaculata were captured in Portugal by Paula (1987). Finally, Balssia noeli Bruce, 1998 has not been reported in Spanish waters, but its presence in Iberian Peninsula waters is quite probable since it was described by Bruce (1998) from Banyuls-sur-Mer, in the NW Mediterranean.

The Pandalidae family is represented by 17 species (within six genera), five of which were not recorded in Iberian waters by Zariquiey Álvarez (1968). These are Dichelopandalus bonnieri captured off northern Spain (Fariña et al. 1997, Domenech et al. 1981); Heterocarpus grimaldii off southern Portugal (Crosnier and Forest 1973 and García Raso 1996); Plesionika ensis in the Alboran Sea and Gulf of Cádiz (García Raso 1981, 1996), Plesionika geniculatus off Portugal (d’Udekem d‘Acoz 1999a) and Plesionika williamsi off southern Portugal (García Raso 1996). Neves (1985) reported the occurrence of Chlorotocus crassicornis on the Portuguese coast, Fariña et al. (1997) in Galicia and Landeira et al. (2015) in the Gulf of Cádiz, while describing several larval stages. A rare pandalid whose presence is probable (although not verified) is Plesionika alexandri (A. Milne-Edward,s 1883). It was captured off northern Morocco during the Balgim Expedition (García Raso 1996 as Plesionika sp.) and in the Azores (d’Udekem d‘Acoz 1999a, Fransen and Biscoito 2006).

The Physetocarididae family is not mentioned in Iberian waters, but the species Physetocaris microphthalma Chace, 1940 has been reported between the Azores and Portugal (Burukovsky 1996), without indicating a specific locality. It is a similar case to that of Meningodora miccyla.

The Pasiphaeidae family is represented by eight species within four genera, but only four species were mentioned in Zariquiey Álvarez (1968). New records are Eupasiphae gilesii, quoted in the Gulf of Biscay (North) and Galicia (Abbes and Casanova 1973) and between the Azores and Portugal (Burukovsky 1996, d’Udekem d‘Acoz 1999a); Pasiphaea ecarina, recently captured by Cartes et al. (2014) on the Galicia Bank; P. hoplocerca, found along the Spanish and Portuguese Atlantic coasts (Cartes et al. 2014, d’Udekem d‘Acoz 1999a, Dos Santos and Moreira 2003) and also in the Gulf of Cádiz by the INDEMARES-Chica expedition (García Raso unpublished data); P. tarda, recently reported in Cantabria and Galicia by Cartes et al. (2007 and 2014 respectively); and Psathyrocaris fragilis (previously reported by Zariquiey Álvarez 1968 as Psathyrocaris fragilis var. atlantica Caullery, 1896), found in Cantabria by Cartes et al. (2007). Other species whose presence is possible are Psathyrocaris infirma, reported in the Gulf of Biscay, French waters (Lagardère 1970) and in Moroccan Atlantic waters (d’Udekem d‘Acoz 1999a); Eupasiphae serrata (Rathbun, 1902), whose northernmost reference is from northern Morocco, at 35°09’N, far from the coast (Crosnier 1988, García Raso 1996), which could be found in the southern waters of the Iberian Peninsula; and Parapasiphae compta Smith, 1884, mentioned between 48° and 44°N and 15° and 17°W (Crosnier 1988, d’Udekem d‘Acoz 1999a). Two other pelagic species of this group that may appear in peninsular Atlantic waters, but more rarely since their closest references are some distance offshore, are Parapasiphae cristata Smith, 1884 and Pasiphaea liocerca Chace, 1940 (Silvertsen and Holthuis 1956).

The Processidae family is represented in Iberian waters by 10 species, all of them belonging to the genus Processa. With the exception of P. macrodactyla, all other species were already reported by Zariquiey Álvarez (1968), although the names of some species have been modified (see Table 1). P. macrodactyla is an African species first recorded in European waters in the Alboran Sea, in Málaga (García Raso and Salas 1985); later it was found in the Bay of Cádiz (Ortega et al. 2005) and in the Catalan Sea (Aguirre and Sánchez 2005). Today there are references of this species even in Turkish waters (Ateş et al. 2004). The occurrence of P. acutirostris from Guipuzcoa (Martínez et al. 2007 as P. cf. acutirostris) needs to be confirmed. We should also mention that the capture of P. intermedia during the CALMEN07 expedition (Junoy 2008) is the first reference on the Iberian Mediterranean coasts. Finally, Williamson and Rochanaburanon (1979) described two subspecies of P. modica (reported as P. parva by Zariquiey Álvarez 1968): P. m. carolii and P. m. modica, the first reported from Cadaqués and southwestern Spain and the second in Atlantic waters, from the North Sea to NW France. The distribution of both subspecies in the Iberian Peninsula has not been studied.

Gurney (1942) considered Amphionides reynaudii (H. Milne-Edwards, 1833) (a species of the family Amphionididae Holthuis, 1955) to be in the Caridea, and although its taxonomic position is not yet fully elucidated, DNA analysis shows its close affinity with the Pandalidae (De Grave et al. 2015). This pelagic species is widely distributed, since it is present in the Pacific, Indian and Atlantic oceans, particularly in tropical and subtropical waters between 36°N and 36°S (De Grave et al. 2015, Fransen 2010), including waters south of the Iberian Peninsula. Consequently, it has not been included in the checklist (Appendix 1), but its presence is likely offshore the southern part of the Peninsula, especially considering the current temperature increase of the oceans.

The old Thalassinidea infraorder is presently considered paraphyletic and was therefore divided into two new infraorders: Axiidea and Gebiidea (Bracken et al. 2009, Dworschak et al. 2012). A study of the European and Mediterranean Thalassinidea was carried out by Ngoc-Ho (2003).

The Axiidea infraorder is represented in Iberian waters by six families: Axiidae, Calocarididae, Callianassidae, Eucalliacidae, Bathycalliacidae and Gourretiidae (Dworschak et al. 2012), containing 13 species within 12 genera, nine of them not mentioned by Zariquiey Alvarez (1968).

In the Axiidae family the new reported species are Calocarides coronatus in Cantabria (Cartes et al. 2007), the Alboran Sea (García Raso 1996, Abelló et al. 2002) and Catalonia (Cartes et al. 1994), and Levantocaris hornungae off the Catalan coast (pers. comm. by Saint Laurent in d’Udekem d’Acoz 1999a).

Concerning the Calocarididae family, Calastacus laevis was quoted in the Bay of Biscay (Saint Laurent 1972a) and Calocaris cf. templemani on the Galicia Bank (Cartes et al. 2014) (as Calocariopsis in Sakai 2011). Additional data on the distribution of other species such as Calocaris macandreae, the most common axiid in Mediterranean waters, are provided by Diez et al. (1994) and Rueda et al. (2012a).

The species of the Callianassidae family from the NE Atlantic and Mediterranean Sea were studied by Saint Laurent and Bozic (1976), but currently this family has been split into several families. In the Iberian Peninsula Zariquiey Alvarez (1968) reported four species in three genera, but since this work several more species have been reported: Necallianassa truncata (previously mentioned by Zariquiey in waters of Melilla as Callianassa truncata) in Guipuzcoa (Martínez et al. 2007), Chingudi Bay, Hendaye (Lagardère 1966), Cascais, western Portugal (Santhino 2009) and the Balearic Islands (Torres et al. 2014). In the last two areas the reports were based on the capture of larvae. Pestarella candida was reported in the Gulf of Cádiz (García Raso 1985b, as Callianassa pestae), Algeciras, the Strait of Gibraltar-Alboran (García Raso 1983 as Callianassa pontica) and Cadaqués, NE Spain (Ngoc-Ho 2003). In Sakai (2011) the last two species are reported as Trypaea truncata and Gilvossius candidus (Pestarella tyrrhena is also cited as Gilvossius tyrrhenus), but these changes have not been acceped in the WoRMS.

Two additional species of Axiidea have recently been captured associated with mud volcanoes in the Gulf of Cádiz: Vulcanocalliax arutyunovi (Dworschak and Cunha 2007), belonging to the Bathycalliacidae family, and Calliax lobate, captured during the INDEMARES-Chica expeditions (Rueda et al. 2012b, García Raso et al. 2018) and belonging to the Eucalliacidae family. The eucallicid Calliaxina punica (Saint Laurent and Manning, 1982) could also be found in Spanish Mediterranean waters, since it is a typical Mediterranean species whose presence is known in the South of France (Ngoc-Ho 2003), but it has not yet been recorded.

The Gourretiidae family is represented in the Iberian Peninsula by a single species, Gourretia denticulata, recorded only in the Gulf of Cádiz (López de la Rosa et al. 1998).

The infraorder Gebiidea is composed of four families, two of them present in Iberian waters.

The Upogebiidae family contains at present four Iberian species. Only one species has been added since Zariquiey’s work: Upogebia mediterranea, from the Balearic Islands (Ngoc-Ho 2003). However, three additional species could be found: Gebiacantha talismani (Bouvier, 1915), whose presence is known in Morocco (Saint Laurent and Loeuff 1979, García Raso 1996), and whose distribution includes the Mediterranean and East Africa (Ngoc-Ho 2003); Upogebia nitida (A. Milne-Edwards, 1868), captured in Toulon and Banyuls (S France), Italy, Algeria, and from Mauritania to the Gulf of Guinea (Ngoc-Ho 2003); and Upogebia stellata (Montagu, 1808), known from the North Atlantic to France and also with a single occurrence, in Greece (Thessalou-Legaki 1986, Ngoc-Ho 2003).

The Laomediidae family is represented by a single species, Jaxea nocturna, found along the whole Iberian coast.

The Anomura infraorder is represented by 85 species belonging to 26 genera and 10 families.

Five species of the Chirostylidae family have been recorded in Iberian waters, two of which have been cited since 1968. These new occurrences are Uroptychus bouvieri, mentioned only in the NW sector of Spain (Cartes et al. 2014), although it probably occurs along the entire Atlantic coast since it has also been captured in northern Morocco (García Raso 1996); and Uroptychus cartesi, a new species described from the Galicia Bank (Baba and Macpherson 2012). In addition, Uroptychus maroccanus Türkay, 1976 is a well-known deep-sea species found in northern Morocco (Türkay 1976, García Raso 1996) and could probably be found in the Iberian sector of the Gulf of Cádiz.

The Galatheidae family is represented in the Iberian waters by 10 species, all of them belonging to the genus Galathea. The geographical distribution of the species given by Zariquiey Álvarez (1968) has been expanded (see Türkay 1976, Neves 1977, García Raso 1987b, 1989), and the list has increased by two additional species: Galathea capillata, captured in the Gulf of Cádiz near the Strait of Gibraltar (García Raso and Manjón-Cabeza 2002); and G. machadoi recently captured on the Bank of Galicia (Cartes et al. 2014) and the Gulf of Cádiz (García Raso unpublished data, INDEMARES-Chica). Galathea intermedia could contain a complex of species. For instance, d’Udekem d’Acoz (1996) mentioned two subspecies (G. intermedia intermedia Lilljeborg, 1851 and G. i. parroceli Gourret, 1887), but these subspecies should not be included until their taxonomic position is clarified. The presence of Galathea rufipes A. Milne-Edwards and Bouvier, 1894 in the Gulf of Biscay also needs to be confirmed (see comments in d’Udekem d‘Acoz 1999a).

The Munididae family is represented by eight species, all of them belonging to the genus Munida. The nomenclature and diagnosis of the species of the genus Munida from the northeastern Atlantic was revised by Rice and Saint Laurent (1986), who considered Munida perarmata a synonym of Munida tenuimana. However, Rodríguez Flores et al. (2016), using both morphological and molecular data, showed that M. tenuimana is a species restricted to NE Atlantic waters, whereas the Mediterranean specimens belong to a different species: they recommended that the name Munida perarmata should be resurrected. Furthermore, two additional species have been recorded in the area: Munida microphthalma, on the Galicia Bank (Cartes et al. 2014), and Munida sanctipauli, captured in the Gulf of Cádiz (García Raso unpublished data, INDEMARES-Chica 0211). New references of some species for different sectors are given (e.g. García Socias and Gracia 1988) in Appendix 1. Finally, Munida rutllanti has been shown to be a junior synonym of M. speciosa von Martens (Rodríguez Flores et al. 2016).

The Munidopsidae family is represented in Iberian waters by 13 species. All of them except Galacantha rostrata are included within the genus Munidopsis. Ten of these species were reported later than 1968 (Zariquiey Álvarez 1968): M. abyssicola, M. acutispina, M. allae, M. antonii, M. aries (=Munidopsis sundi Sivertsen and Holthuis, 1956), M. curvirostra, M. marionis, M. parfaiti, M. thieli and M vaillantii. All of them were reported from Atlantic waters of northern Spain, and/or in western and/or southern Portugal (see Appendix 1) (Saint Laurent 1985, Macpherson and Segonzac 2005, Cartes et al. 2014, Türkay 1975 and 1976, d’Udekem d‘Acoz 1999a, and data from ORBIS). In addition, Abelló and Valladares (1988) recorded Munidopsis serricornis in the Catalan Sea.

The Porcellanidae family includes four species, all mentioned in Zariquiey Álvarez (1968). Pisidia longimama was cited in Portugal by Neves (1977) and Nunes-Ruivo (1961), and in Galicia by Anodon (1975). The taxonomy of the genus Pisidia is currently under review and, based on morphological and molecular studies, the three known species will be validated and a fourth new species will be described (Cuesta et al. unpublished information).

The Albuneidae family is represented in Iberian waters by just one species already mentioned by Zariquiey Álvarez (1968), Albunea carabus, which shows a Mediterranean distribution, although it has recently been found in southern Portugal (Pereira et al. 2008).

No species of the Lithodidae family were mentioned by Zariquiey Álvarez (1968), but three species have been captured in northern Spain, all of them on deep-sea bottoms (see Saint Laurent 1985 and Macpherson 1988): Neolithodes grimaldii, Paralomis microps and Paralomis bouvieri Hansen, 1908. The last species was located in the Cantabrian Sea according to Figure 33 in Macpherson (1988).

The six species of the Diogenidae family known in Iberian Peninsula waters were all mentioned by Zariquiey Álvarez (1968).

Six species of the Parapaguridae family are now known in Iberian waters, three of them since 1968. Parapagurus nudus and P. abyssorum in the Gulf of Biscay (Saint Laurent 1985), and Paragiopagurus ruticheles in southern Portugal (Lemaitre 1990 as Sympagurus ruticheles). Lemaitre (1989) did a review of the genus Parapagurus in the eastern Atlantic, completing the previous study by Saint Laurent (1972b).

The Paguridae family is the most diverse, with 28 species within six genera. New references for different geographic sectors have been given by Ingle (1993), García Raso (1982a, 1984b, 1985c), and the ECOMARG (IEO) 2005 and 2006 Expeditions, among others (see Appendix 1). Seven species of pagurids have been added since Zariquiey Álvarez (1968): Anapagurus alboranensis, described from the Alboran Sea by García-Gómez (1994) and later recorded in the Gulf of Cádiz (Manjón-Cabeza and García Raso 1998) and along the Spanish Mediterranean coasts (Macpherson and Raventos 2004); Anapagurus pusillus, captured in Atlantic waters in southern and western Portugal (García-Gómez 1994, Sampaio et al. 2016) and in northern Spain (Cantabria, ECOMARG IEO, Serrano et al. 2006); Catapaguroides iris, captured in the Gulf of Cádiz (one specimen from INDEMARES-Chica 0211, García Raso unpublished data), previously known in Azores and the north of Morocco (Ingle 1993, García Raso 1996); Pagurus pseudosculptimanus, described by García Muñoz et al. (2014) and recorded in the Alboran Sea; the African Pagurus mbizi, which is penetrating into the Mediterranean through the Alboran Sea (García Raso et al. 2014b); Pagurus pubescentulus, recorded in Cantabria (IEO ECOMARG, Serrano et al 2006); and Michelopagurus atlanticus, a deep-sea (790-1575 m according to Lemaitre and Tavares 2015) amphi-Atlantic species recently found off Aveiro, Costa de Prata (western Portugal) at a much shallower depth (41 m) by Sampaio et al. (2016), which is noteworthy. Also, the occurrence of Anapagurus smythi Ingle, 1993 in the Cantabrian Sea (IEO ECOMARG http://www.ecomarg.com/biodiversidad.html) should be reviewed (so it is not included in Appendix 1). The specimen of Catapaguroides iris from INDEMARES-Chica 0211 shows a mixture of morphological characteristics of the two species cited in the area. The cephalothoracic shield has the same length and width (L/W=0.93, as shown in Figure 81a, p. 367 in Ingle 1993, for the holotype of C. iris), no longer than wide (as in Figure 85a, p. 345 in Ingle 1993, for a paratype of C. megalops), but the carpus of the large cheliped has an incomplete row of five spines (not a simple row of setosed tubercles as shown in Figure 81f in Ingle 1993 for C. iris, nor a complete and well-developed row of small sub-acute setose tubercles as shown in Figure 85g in Ingle 1993, for C. megalops). To complicate matters further, Saint Laurent (1968) drew a specimen (L/W=0.98) designated ?C. megalops, with a shield similar to that drawn by Ingle 1993 for C. iris, but with a strong row (even double row) of acute tubercles-spines. However, the deep-sea species C. megalops A. Milne-Edwards and Bouvier, 1892 could also be found, since it is known in nearby areas of the Atlantic (Morocco and Azores, Ingle 1993).

Other studies have increased the distribution area of some marine species in Spanish (García Muñoz et al. 2008, García Raso 1985c, García Raso et al. 2014a, Serrano et al 2006) and Portuguese waters (e.g. Pagurus chevreuxi by Almeida 2008). Furthermore, larvae of Cestopagurus timidus have been captured in southwestern Portugal (Paula 1987).

To complete the information regarding the Iberian Peninsula decapod crustacean fauna, it is worth mentioning that up to nine strictly freshwater species have been reported in the Iberian Peninsula, including the Balearic Islands (excluding brachyuran crabs). These are four crayfishes (Vedia and Miranda 2013): Austropotamobius italicus italicus (Faxon, 1914) (recorded as A. pallipes lusitanicus in Zariquiey Álvarez 1968) and Pacifastacus leniusculus (Dana, 1852) (both within the family Astacidae Latreille, 1802); Procambarus clarkii (Girard, 1852) (family Cambaridae Hobbs, 1942); and Cherax destructor Clark, 1936 (family Parastacidae Huxley, 1879). The last three species have been intentional human-mediated introductions for aquaculture. One yet undescribed alpheid shrimp of the genus Bermudacaris was discovered in an anchialine cave in the Balearic Islands (Gràcia et al. 2003, Gràcia and Jaume 2011). The freshwater list also includes three atyids, Atyaephyra desmarestii (Millet, 18319), present in all Iberian freshwaters, Dugastella valentina (Ferrer Galdiano, 1924), present in the Gulf of Valencia (Sanz and Gómez 1984), and Typhlatya miravetensis Sanz and Platvoet, 1995, described from a cave in Ullal de la Rambla de Miravet in Cabanes, Castellón by Sanz and Platvoet (1995), and also one palaemonid, Palaemon zariquieyi Sollaud, 1938 (Sanz Brau 1986) (as Palaemonetes in Zariquiey Álvarez 1968). The latter three species are endemic to the eastern sector.

DISCUSSION

Species richness and distribution by geographic sectors

The number of marine decapod crustacean species (excluding Brachyura) reported by geographic sectors (Table 2) shows that the greatest specific richness is found in Atlantic waters, with 264 species, while a total of 178 species have been found in Mediterranean waters. These differences are the consequence of the relative extension of the areas, the depth ranges and the confluence of different water masses. The whole Atlantic sector has the greatest extension, the largest range of depths, and also the greatest diversity of water masses, with different characteristics and associated species. As an example, the various different water masses found in the Gulf of Biscay and Galicia are East North Atlantic Central Water, Mediterranean Outflow Water, which forms in the Gulf of Cádiz and progresses northwards, and Labrador Sea Cold Water (Cartes et al. 2007, 2014). In the Gulf of Cádiz there are clear influences of two marine ecoregions, the Lusitanian (with Atlantic and African sectors) and the Mediterranean (Spalding et al. 2007), or three provinces (Lusitanica, Mauritanica and the Mediterranean) (Bianchi et al. 2012), with different associated species. It must also be borne in mind that the fauna present in the Mediterranean has an ancient Atlantic origin (originated after the Messinian salinity crisis in the late Miocene 5.6 mya, Bianchi et al. 2012). Consequently, most Mediterranean species are also found in the Atlantic. The interaction of the water masses present and their dynamics, with the geological and environmental history, account for the present biogeographic structures. The greatest species richness is found in the Gulf of Cádiz and Gulf of Biscay - Galicia (GB-GA) sectors, with 194 and 176 species respectively. The Alboran Sea shows a lower species richness, probably because of its shorter coastal extension and lower range of depths, especially when compared with adjacent sectors.

Table 2. – Number of marine decapod, families (F), genera (G), species, total (sp) and new registers after Zariquiey Álvarez (1968) (spnr) (excluded crabs and freshwater species) ordered by families and geographical sectors. GB-GA, Gulf of Biscay and Galicia; WP, western Portugal; GC, Gulf of Cádiz or southwestern Iberian Peninsula; ALB, Alboran Sea; SWM, Spanish western Mediterranean; S, southern Iberian waters (GC+ALB); A, Atlantic Iberian waters (GB-GA+WP+GC); M, Mediterranean Spanish waters (ALB+SWM).

[Table omitted. Please see PDF.]

Geographically, a total of 222 species have been identified in southern Iberian areas (GC + ALB), 176 species in northern areas (GB-GA), 160 species in eastern Spanish waters (SWM) and 155 species in western Portugal (WP).

The total number of marine decapod species known in and around Iberian waters, excluding Brachyura, is 293. However, this number increases to 305 when the nine freshwater species and several unpublished new records (see above) are also included.

Introduced marine decapod species

The number of introduced species (excluding brachyuran crabs) in the Iberian waters is relatively low (four, see Appendix 1 “Int”) and appears to be mainly related to human activities such as aquaculture (Zenetos et al. 2010, 2012). The four species are Penaeus pulchricaudatus (Penaeidae family), previously reported as Penaeus japonicus Spence Bate, 1888 (Rodríguez 1989, Guerra and Gaudêncio 2016, Tsoi et al. 2014), Panulirus regius and Jasus lalandii (Palinuridae family) (Zariquiey Álvarez 1968, Guerra and Gaudêncio 1982, Holthuis 1991), and Palaemon macrodactylus (Palaemonidae family).

Tsoi et al. (2014) showed that the references of P. japonicus in the Mediterranean corresponded to the cryptic species P. pulchricaudatus, but they only analysed specimens from its eastern basin. Therefore, we assume that the Iberian specimens also correspond to P. pulchricaudatus. This should, however, be checked since it is likely that these reports may be due to escapes from aquaculture facilities.

No additional occurrences of Panulirus regius and Jasus lalandii have been reported in Iberian waters, suggesting that their introduction around the Iberian Peninsula has not been successful. However, Froglia et al. (2012) captured several specimens of P. regius in Italian waters in the Ligurian Sea and considered, taking into account the very long larval planktonic life of the species and the size of the specimens caught, that the occurrence of a successful population could be related to a population range expansion driven by global warming and Atlanto-Mediterranean currents rather than being a direct result of human introductions. In addition, Guerra and Gaudêncio (2016) mentioned the occurrence of the western Atlantic lobster Panulirus guttatus (Latreille, 1804) on the Portuguese coast, but without specifying any reference and and introduction pathway (this species has not been considered in this paper).

The shrimp Palaemon macrodactylus has been introduced most probably through ballast water and presently shows well-established populations (Cuesta et al. 2004, Torres et al. 2012, Chícharo et al. 2009, González-Ortegón et al. 2010).

Additionally, the cause of the presence of certain African species (whose previously known northern limits of distribution were far to the south of the Iberian Peninsula) is not clear. They could be human-mediated introductions, but the scarcity of studies along the NW African coasts may also explain the lack of knowledge concerning the precise distribution areas of the species. Range expansions may also be due to climate change effects because the warming of the waters favours the northern expansion of thermophilic Atlantic species along the Atlantic coasts, and their introduction and further expansion in the Mediterranean Sea (e.g. Pagurus mbizi, García Raso et al. 2014b). Thus, the presence of these species today could be the result of natural expansions of their geographic distribution ranges rather than human-mediated introduction.

General reflections on the total specific richness in the Iberian Peninsula

The total number of decapod species in the Iberian Peninsula (marine and continental), including the 140 species of crabs reported by Marco-Herrero et al. (2015) and other new species in process of description, comes to 449. Regarding the crabs mentioned by Marco-Herrero et al. (2015), some references need clarification. In this regard, there are cases under study of synonyms within the Calappidae and Leucosiidae families [e.g. Calappa tuerkayana Pastore, 1995, a synonym of Calappa granulata (Linnaeus, 1758) (Holthuis 2001); a similar case could be found within the genus Ebalia] that will decrease the number of species in it. However, other species have to be added, such as Inachus parvirostris (Risso, 1816), captured on the CALMEN07 expedition (Junoy 2008) in the vicinity of Alboran Island (García Raso, unpublished data, INDEMARES Alboran expedition) and Pinnotheres petunculi Hesse, 1872 (Pérez-Miguel et al. 2017). Within the Pinnotheridae family there are also new species of crabs in process of description.

Consequently, the total number of decapod crustacean species in Iberian waters could reach 448, much higher than the total reported in Italy, with 324 marine and freshwater species (Froglia 2010); in Turkey, with 265 (Ateş et al. 2010, Deval and Froglia 2016); and on the coasts of Algeria, with 253 (Grimes et al. 2016). However, when only the species richness of the Iberian Mediterranean coasts (with less km of coastline) is compared with that of Italy, the number of species (about 296) is lower, although still higher than those reported for Algeria and Turkey. The position of the Iberian Peninsula in the western Mediterranean Sea, surrounded by both Atlantic and Mediterranean waters, is probably the main reason for this greater richness of species. The wide range of depths and long coastlines occurring in the area may be important additional reasons for the high species richness.

Finally, we must mention that molecular analysis studies are currently being carried out to verify the validity of some species. Moreover, the exploration of deep habitats and ecosystems under the European Framework Directive for the Marine Strategy will report additional new occurrences and species. Accurate faunistic analysis of fishery discards may also become an increasing source of biodiversity knowledge that may help to understand biogeographic patterns and dynamic changes in distribution, including early detection of allochtonous and invasive species (Gorelli et al. 2016, Anjos et al. 2018). Another interesting and necessary activity, which is also carried out in some areas, is the study of meroplanktonic larvae in relation to hydrographic currents and climate change, because it improves knowledge of specific composition and biocenoses, and can explain the presence of new records and the increase of the biogeographical range of some species (Torres 2015, Landeira et al. 2017, Landeira and Lozano-Soldevilla 2018).

ACKNOWLEDGEMENTS

We would like to express our sincere gratitude to Isabel M. Muñoz for providing information on some species. We thank Royston F. Snart for the English revision of the manuscript. This work was partly supported by the INDEMARES/CHICA “Mud volcanoes from Gulf Cadiz” and INDEMARES/ALBORAN Projects, EC contract INDEMARES-LIFE+ (07/NAT/E/000732) and FEDER funding assigned to equipment of the R/V Cornide de Saavedra (FICTS-2010-01), CTM2017-88080 (AEI/FEDER, UE). Finally, thanks are due to Dr E. González-Ortegón and an unknown reviewer for fruitful comments and critical reading of the manuscript.

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