The Biodiversity of the Mediterranean Sea: Estimates, Patterns, and Threats

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- Cnidaria

We summarized available information for the Classes hydrozoans, scyphozoans and cubozoan [31,78], while a special emphasis is put on anthozoans. Anthozoans are common in most marine habitats from shallow to deep oceanic waters. Although most anthozoans occur on hard substrata (Gorgonacea, Scleractinia, etc.), a large number are adapted to life in mud, sand or gravel (Pennatulacea, Ceriantharia) in big range of current velocities but favoring for strong ones [e.g. 79,80]. They feed on nanoeukaryotes, dinoflagellates, diatoms, ciliates, as well as detrital POC and are generally considered microplankton suspension feeder [81]. For some the nutrition comes via their symbiotic relationship with small algae living in their gastrodermis and other are carnivorous [e.g. 82,83,84]. Although Aristotle (4th century BC) was the first to mention the presence of four anthozoans in the Aegean Sea [38], the knowledge on biodiversity of this group in the Mediterranean Sea rose after 17th century. Here we revise previous estimates of anthozoan diversity.

A total of 757 species of Cnidaria are currently quantified (Table 1 and Table S1). Anthozoa species recorded so far are 164 species including 51 Octocorallia and 113 Hexacorallia (Table S13). Two primary faunal components are recognized: Atlantic-Mediterranean (62.20% of the fauna by numbers of species) and endemic (21.95%). The boreal component contributes only minimally (1.83%), while the remaining fauna is made up of cosmopolitan (8.54%) and Amphi-Atlantic species (4.88%). There is only one confirmed Indo-Pacific component (A. erythraea) in the Mediterranean Sea. Introduced corals into the Mediterranean have been cited due to shipping via the Atlantic: Oculina patagonica and Diadumene cincta [85,86], and in the Red Sea: Acabaria erythraea [87].

The geographical and bathymetric distribution of anthozoan species and genera richness in the different regions of the Mediterranean is rather uneven (Table 2, and Table S13). Three major zoogeographical zones have been identified in the Mediterranean according to the affinities of their anthozoan fauna: a zone of high diversity (Alboran Sea, Western basin, Tyrrhenian basin), a zone of moderate diversity (Adriatic Sea, Aegean Sea), and a zone of impoverished diversity (Ionian basin, Levantine basin). The higher species diversity in the western basin can be explained by the occurrence of a number of Atlantic species with eastern boundaries in the Mediterranean, i.e. in various areas of the western basin [88,89,90]. Intensive research effort in the western Mediterranean may also be responsible for this difference in numbers. Another reason is the general impoverishment in the diversity of the eastern part biota due to the oligotrophic conditions in the area [91].

Corals have been used since antiquity for jewellery, costume accessories and souvenirs [92], and also produce diverse compounds useful in medicine [93]. Harvesting and ripping out of colonies, as well as chemical pollution, burial and sediments deposition, are harming too many species, particularly permanently attached, erect forms such as gorgonians, antipatharians and scleractinians. Red coral (Corallium rubrum) is a precious coral and has been widely sought after since ancient times in the Mediterranean. Nowadays it is one of the most over-exploited invertebrates in this sea [e.g. 92,94].

Less than 20% of the corals living in the Mediterranean have been included under the annexes of the conventions for the protection of animal life. The most of them (85%) are only protected under Annex II of CITES, which does not extend full protection but rather regulates their commercial trade. Six of them are protected under EC legislation or international conventions and two are included in the lists of maximum protection [95].

- Mollusca

Mollusks are important components of marine communities worldwide, making up to 15-25% of the benthic macrofauna. This is one of the few phyla that are routinely taken into consideration in marine biodiversity surveys and are considered to be an ‘appropriate indicator group’ for rapid assessment of diversity inhabiting a particular area [96]. Their importance did not escape early human settlers who clearly appreciated their value as seafood or the beauty of their shells. The Mediterranean molluscan fauna is one of the most anciently studied. Most malacologist, professional or amateur, are aware of the seminal works of Forbes (1844) for the Aegean Sea, Philippi (1836) for Sicily, or Bucquoy, Dautzenberg and Dollfus (1882-1898) from the French Mediterranean coast, among others. In the late 19th century, syntheses of available knowledge on the Mediterranean mollusks were provided by Weinkauff, Monterosato, Carus, among others.

General collecting procedures, techniques and gears are used to sample mollusks in the Mediterranean (towed nets to sample planktonic species, fishing gears to sample cephalopods, bottom sledges, dredges, grabs, box samplers or corers to collect members of the infauna, scuba diving sampling for littoral species living of rocky bottoms, and many other specialized methods). A useful and rapid method to give preliminary background information of the mollusks’ diversity of a particular area is the study of small samples of bioclastic sediments (a sediment type composed of fragments of organic skeletal materials and shells of micromollusks). Empty shells are a biodiversity indicator that points the difficulty of estimating the real magnitude of species richness for taxonomical groups that do not have post mortem remains, such as flatworms, polychaetes, meiofauna, or peracarid crustaceans [96]. One small sample of bioclastic sediment in the Mediterranean may contain more than 100 species of mollusks [97].

In the Mediterranean, 2113 species of marine mollusks are known. Table S14 shows the completed species list of mollusks, including cephalopods, based on Sabelli et al. [98] and Bello [99] and updated from CLEMAM database (Check List of the European Marine Mollusca: The class Gastropoda is the one with higher number of species (74% of all known species in the Mediterranean), followed by Bivalvia (19%), Cephalopoda (3%), Polyplacophora (1.5%), Solenogastres (1.4%), Scaphopoda (0.7%), Aplacophora (0.4%), and Monoplacophora (0.05%). In general, mollusks show a decreasing biodiversity from the west to the east, with about 45% of the species in the eastern basin (Table 2 shows data regarding cephalopods and gastropods). Two main ‘hot spots’ for mollusks are observed in the Alboran Sea and the area around Sicily. A mixture of Mediterranean and Atlantic species are present in the former, where Penas et al. [100] recorded up to 655 species of mollusks in a small area surrounding the Alboran Island. On the other hand, the central zone of the Mediterranean Sea around Sicily is an area with a high degree of endemic species. Moreover, the Mediterranean Sea currently hosts more than 150 exotic species of mollusks, of which about 90 forms established populations [101,102]. The bulk of the introduced species of mollusks in the Mediterranean (about 154) are species of Indo-Pacific origin, mostly as ‘Lessepsian immigrant’.

Mediterranean mollusks are highly diverse from a morphological and ecological point of view, ranging from minute wormlike interstitial animals (smaller than 1 mm) to giant squids (Architeuthis), and from minute snails (0.7 mm in Retrotortina fuscata) to giant fan shells (up to 90 cm in Pinna nobilis). Within gastropods (the most diverse class of mollusks) almost 35% of the species have an adult size smaller than 5 mm (micro-mollusks). By contrast, species larger than 50 mm account only less than 3%. Nevertheless, faunal surveys and inventories have a tendency to focus on the large species of ‘seashells’ and neglect the smaller species.

While members of most classes of mollusks are adapted to a particular environment or life-stile (caudofoveates, scaphopods, and most of bivalves are members of the infauna of sedimentary bottoms, monoplacophorans and polyplacophorans live on rocky surfaces, solenogastres are epifaunal members that prey upon cnidarians, cephalopods are highly specialized predators), gastropods are present in any marine environment (from hydrothermal vents to the pelagic realm), and they show great disparity in external form and behaviour. All kind of feeding habits (micrograzers, deposit feeders, herbivores, filter feeders, parasites, generalist or specialized predators, and scavengers) can be found within gastropods. All species of the order Acochlidiacea are interstitial animals, and all members of the orders Thecosomata and Gymnosomata and few others are holoplanktonic animals. Within the pelagic realm, cephalopods are the only group of mollusks that have nektonic species, reaching large adult size, are placed as predators competing with fishes. Most of the benthonic mollusks live in littoral areas or in the continental shelf (about 96% of the species) and only about 4% belong to the bathyal fauna. Some groups of mollusks evidence significant degree of endemics, for example, nearly 10% of the Mediterranean cephalopods are considered endemic or quasi-endemic, a characteristic observed in the Family Sepiolidae [99].

Although mollusks have been more intensely studied in the Mediterranean than almost in any other sea since the antiquity, an average of five to ten new marine species of mollusks are still being described each year in this sea. As an example, Penas et al. [100] recently described eight new species of gastropods near the Alboran Island. Currently, large-scale DNA sequencing provides the view of biodiversity being underestimated in all parts of the tree of life due the existence of cryptic species. Therefore, probably many more Mediterranean new species of mollusks will be recognized in the next years [e.g. 103]. Besides, many exotic species are being added to the native species because the global process of ‘bioinvasions’ is affecting notably the Mediterranean Sea, as it has been commented before.

Habitat loss and degradation is the main threat that impacts the molluscan diversity in the Mediterranean coasts, following by over-exploitation and fisheries (due to cephalopods and bivalves are important fishing resources for Mediterranean countries), pollution, introduction of new species, climate change and others. These threats are frequently cumulative and cause biotic homogenization and impoverishment. Mass mortalities have also known on some species, for instance the bivalve Spondylus gaederopus suffered widespread mortality in 1981 and 1982 [104] probably because a viral, bacterial or fungal infection. Besides, anchor causes notably damage the populations of the large fan shell Pinna nobilis [105]. In some cases, fishing periods coincide with reproductive seasons for cephalopod species, increasing the impact on their populations [106,107].

Seventeen Mediterranean species of mollusks have been included in the Anex II of the Barcelona Convention (Annex II and III) and Appendix II of the Bern Convention as worthy of protection. One of the species included in these lists is the data mussel Lithophaga lithophaga, because its overfishing by scuba divers causes serious damage along calcareous coasts of the Mediterranean Sea [108,109]. This boring bivalve is harvested by scuba divers, who smash the rocks with chisels or pneumatic hammers to detach the specimens from the walls into which their live. The major consequence is the removal of the biological cover (macroalgae and zoobenthos), which ranges from bare patches to complete desertification of the bottom communities [110].

- Polychaeta

Polychaeta are truly segmented worms belonging to the phylum Annelida, among which they represent the class with the largest number of described Mediterranean species. The Mediterranean Polychaeta fauna has always been among the most intensively studied. A wide panoply of sampling and analytical methods are involved in the study of polychaetes [111,112,113,114], while major insights in revealing cryptic species have been obtained, and more are still expected, from the use of fine morphological [e.g. 115], and molecular techniques [e.g. 116]. Based on an extensive taxonomic revision of all known European polychaetes, in which all existing literature up to 2008 has been checked and contrasted, we updated the total estimates for the Mediterranean polychaete species (see Table S16), which is then compared with the respective estimates for geographically restricted areas (western and eastern basins, and the Iberian Mediterranean coasts).

Polychaetes have a wide size range, extending from small meiofauna (less than 1 mm long) to big megafaunal (reaching about three m long) organisms, and are among the most common inhabitants of marine benthic bottoms, from shallow-waters to deep-sea and from brackish to hypersaline waters, with well-consolidated incursions into the plankton, as well as in continental environments [117]. They inhabit all types of substrata, from rocky bottoms to muddy sediments, where they may be among the most dominant organisms, both in terms of abundance and biomass, and also often in diversity [118]. Most of them are free-living, showing a variety of feeding strategies [119], but there are numerous cases of more or less specialized symbionts, from parasites to mutualists, living in association with many marine taxa, including other polychaetes [120].

There are numerous works in the Mediterranean Sea comprising different geographical ambits, which include more or less complete taxonomic compilations of polychaete species (see Table S16). A first attempt to estimate the total number of benthic Mediterranean polychaetes was done by Bellan [121] who reported 950 species. More recently, Arvanitidis et al. [122] extensively analysed the Mediterranean and Black Sea polychaete biodiversity patterns, reporting 1036 species as valid (based on a 1999 inventory). Our analysis reveals that the whole Mediterranean polychaete fauna currently includes 1122 species, grouped in 452 genera belonging to 72 families.

According to Arvanitidis et al. [122], the Mediterranean polychaetes are dominated by cosmopolitan species (more than 30% of the total), while the Mediterranean endemics represented 18.8%. However, the former trend (often attributed to the presence of opportunistic species), may be confusing, as many supposed cosmopolitan species revealed to have, after recent accurate morphological and/or genetic studies, more geographically restricted distributions, and the Mediterranean cosmopolites (e.g. Haplosyllis spongicola) are certainly not an exception [see 123]. In turn, a few Mediterranean endemics have to be removed from this category after looking at the adequate habitats in other biogeographical regions, or as a consequence of more precise taxonomic revisions. Haplosyllis chamaeleon (a symbiotic species associated to the gorgonian Paramuricea clavata, another Mediterranean endemics), has been recently found to live in association with Paramuricea grayii in the Atlantic coasts of Galicia, NE Iberian Peninsula [123], while Acanthicolepis costeaui was shown to be a junior synonym of Acanthicolepis asperrima, a species with a wider distribution along the European Atlantic coast [124]. Thus, the number of species known only from the Mediterranean Sea may be currently estimated as 210, taking into account that it is not yet possible to assess how many of them are truly Mediterranean endemics.

The western Mediterranean is the richest basin for polychaetes, and most likely the best studied too (Table 2). It is harbouring 85% of the species. The Central Basin, Adriatic Sea and Aegean Sea harboured a 50%, respectively, while the Levantine Basin harboured less than 45%. A total of 946 polychaete species are known from the western Mediterranean, 877 (78%) from the eastern Mediterranean, and 601 (54%) from the Iberian Mediterranean coasts. Therefore, the western basin is a 6% richer than the eastern one, but we have to consider the spectacular increase of the diversity of the polychaete fauna from the eastern basin lately, probably associated with the increasing number of studies in this biogeographic region. The Iberian Mediterranean has a reduced set of species, but its variation in taxonomic distinctness (283.4), shows a significant departure (p = 0.08) from the upper limit of the simulated distribution [125,126,127], indicating that some higher taxa include more species than those expected at random. A possible explanation may be the recent faunistic efforts concentrated in a few families carried out within the frame of the Fauna Iberica project [128,129].

New species are still being described either as a result of studying common habitats using new approaches [e.g. 130], or unexplored environments or regions in any way [e.g. 114], while an additional source of diversity may be invasive species [131] including Lessepsian migrants [e.g. 132], which were estimated to be around 6-7% the known Mediterranean species (i.e. 70 - 80 known invaders nowadays). The same occurs with a significant part of the southern coasts, and major advances in terms of the knowledge of the diversity of the group may be expected from the exploration of these regions. The average number of polychaete species that have been newly described from Mediterranean waters is 3.8 per year, since the first one described by Linné [133]. This rhythm of descriptions was almost four times higher (12.4 new species per year) during the 1860s, with the concurrent works of R. E. Claparède, A. E. Grube and E. H. Ehlers, among others, and was almost twice as high (5.7 new species per year) during the late 1960’s and early 1970’s. In the last five years (2005-2009), there has been an average of 4.2 newly described Mediterranean polychaete species per year (Figure 13a). Although the deep-sea Mediterranean bottoms seem to be less rich that those from the nearby seas (e.g. the Atlantic ones), they are still largely unexplored. The same occurs with a significant part of the southern coasts, both of the western and eastern basins. Major advances in terms of the knowledge of the diversity of the group may be expected from the exploration of these regions.

Major threats to polychaete diversity in the Mediterranean Sea may be linked to the increasing anthropogenization of the coasts, the global increase of temperatures, changes in the water quality (e.g. acidification, turbidity), overfishing, or the presence of introduced/invasive species, sometimes in a massive way like in the case of the reef-building serpulid Ficopomatus enigmaticus [134]. Polychaetes can also play a harmful role as fouling organisms in ship-hulls or harbours and other marine structures, as pests in natural and cultured oyster populations, or as introduced species, with the subsequent economical losses [e.g. 135].

Moreover, some polychaetes are eaten by humans, mainly those known as “palolo” worms [136], while others have been traditionally used in local pharmacopoeia [e.g. 137]. The presence of toxins or venom glands in some groups (e.g, Amphionomidae, Glycera, Metaxypsamma), and the fact that other groups are chemically defended [e.g. 138], opens the possibility of new investigations and applications in pharmacology and medicine. Polychaetes have also a significant economical relevance, as revealed by the growing commercial activities and the international market for polychaete species that are dug up or farmed, mainly for being used as fishing bait and as a food item in aquaculture, with the implied risk of introducing foreign species and associated pathogens or other non-native organisms in the wild [e.g. 139,140]. Threats to polychaete diversity may be also linked to overfishing. However, polychaetes are ubiquitous and, unless major perturbations occur, the only expected consequences would be the replacement of some species by others locally or regionally. An exception may be species associated with restricted habitats, like coastal lagoons, or with specialized life habits, like the symbiotic Ichthyotomus sanguinarius [an external parasite of eels never found again since its original description, 141].

- Arthropoda

This phylum is highly diverse and species can be found all along the Mediterranean Sea and among the most common habitats. The estimation of total number of species is difficult because species vary enormously in size and habitats, as well as they go through metamorphic changes that have confused taxonomists for centuries. Information for this synthesis came from several sources (Table 1 and Table S1). Crustaceans are the dominant group in terms of biodiversity and they include some commercial groups that have been better studied (such as decapods or stomatopods). Estimates for several arthropod groups of the Mediterranean Sea have been listed in regional lists along the basins, and several checklists exist (Table 1 and Table S1). Here we summarize main species estimates and we revised in detail the estimates available for i) cumaceans, ii) mysidaceans and iii) decapods.

A total of 2244 species of crustaceans have been registered so far in the Mediterranean Sea (Table 1 and Table S1). Cumaceans and mysidaceans are important groups belonging to the suprabenthos communities. Studies in the western Mediterranean showed that they can comprise from 42% of total suprabenthos in deep waters [862-1808 m, 163] to 61.8% in shallower areas [1-3 m, 164]. A total of 871 species are listed here to be the principal components of the Mediterranean suprabenthos communities, including Cumacea, Mysida and Lophogastrida, Isopoda, Amphipoda, Tanaidacea, and Euhpausiacea.

(i) Cumaceans: Accurately sampling the near-bottom invertebrate swimming species (in the suprabenthic habitat) is a difficult task due to the size of the species and their behaviour. Cumaceans, for example, were considered during long time larval stages of other crustaceans. Adult cumaceans range from 1.5 to 35 mm total length but they grow into small development instars before reaching the adult form. Fractions of a same population are lost if they are sampled or sieved with standard mesh size such in studies of macrofauna [142]. Cumaceans live in the water-sediment interface, they burrow into the sediment, and some littoral species migrate to the water surface during night-time [143]. Studies to quantify their biodiversity use dredges, and when densities are low, as frequently it happens in deep water, suprabenthic or epibenthic sledges that collect animals from the nearest sea floor water layer are used.

It was only in 1870 when cumaceans were recognized as an independent order. Since then, the knowledge of this crustacean group has grown slowly and taxonomic effort has varied greatly among biogeographical regions. The Mediterranean Sea, together with the northeast Atlantic Ocean, is likely one of the best-studied regions in the world. Our estimates show that cumaceans from the Mediterranean are composed of 99 species (Table S17). Six of the eight cumacean families are present in the Mediterranean Sea, lacking the family Gynodiastylidae, distributed in the southern Hemisphere and the family Ceratocumidae, the species of which are restricted to deep waters. Nannastacidae is the most specious family (28 species), followed by Bodotriidae and Diastylidae (27 and 23 species respectively). Leuconidae is represented by 13 species while Lampropidae and Pseudocumatidae have only four species each. The level of endemics is relatively high reaching to 32.3% for the whole Mediterranean Sea. In addition to the still unknown cumaceans Mediterranean species, it is also very probable that other foreigners could invade this sea. This fact has been already observed in the Levantine Sea where a Red Sea species was recently found [165].

In the NW Mediterranean there are 78 cumacean species known that account the 91.8% of those recorded for the western basin, and the 78.8% of those in the whole Mediterranean Sea. Conversely, in the Tunisian Plateau/Gulf of Sintra and the Adriatic Sea only few species have been recorded (four and 13, respectively). A total of 85 species are recorded in the western basin versus 74 in the eastern (Table S18). However, high differences in species richness observed in the four regions considered of the western basin are mostly due to the different taxonomical effort instead to an actual difference between the fauna of these seas. Both basins have 15 endemic species in common, while there are 11 endemic species exclusives from the western basin, and seven from the eastern. In terms of percentage, endemics are very similar in both basins (30.6% in the western and 29.7% in the eastern).

The taxonomic effort to describe cumaceans diversity varied also greatly among regions and areas such as the Adriatic Sea or the Gulf of Gabés are practically unknown. The accumulation curve of cumaceans species discovered (described or first recorded) in the Mediterranean Sea (Figure 13b) shows that no asymptote has been reached so far, and therefore, there has been no slowing in the rate of discovery of Mediterranean cumaceans since late 19th century. This represents an annual increase of about 0.5% of the known Mediterranean cumaceans. This rate of discovery, despite it seems low, is very similar to observed in most groups of crustaceans [172].

Human activities manly focused on littoral waters such as fishing, coastal engineering, sandy beach restoration, fish farming, etc., are affecting and will affect the area where cumacean diversity and level of endemics of these crustaceans are highest. Cumaceans, and especially those living in shallowest bottoms, are very sensitive with the mean size grain of the sediment [173] and small changes in its composition may lead to the disappearance of certain species. Changes in the cumacean assemblage structure were already observed in the Spanish coast induced by massive influxes of waste water and sludge discharges [174]. Increment in shipping, as well as in water temperature because the global warming, will favour the establishment of aliens that could threaten the autochthonous cumaceans fauna.

(ii) Mysidaceans: They are regularly represented in the Mediterranean Sea, from the Alboran Sea to the Marmara Sea and from the coastal lagoons and beaches to the bathyal environments. The majority of the species live as well in the suprabenthic habitat, but some can be found in hard bottoms (or caves) or performing vertical migrations between the bottom and the surface, mainly at night. The studies of Mediterranean mysidacean began in 1837, when the first two species were described by Milne-Edwards. During the decades of 1860-1941 several species were described as a result of intensive sampling by diverse authors (G.O. Sars, W. Tattersall, V. Czerniavsky, M. Bacescu). From Bacescu [144] contribution, the knowledge of the mysidacean biodiversity has grown regularly until now.

Several methods are used to sample the majority of lophogastrid and mysid species [145,146]. The majority of world species [74%, 147] live in the suprabenthic habitat (or hyperbenthic, composed of near-bottom swimming species), which need specific gears to sample. The most efficient mysidacean samplers, and in general for suprabenthos, are nets mounted on suprabenthic sledges that are towed over the surface of the sediment [148,149]. The choice of a suprabenthos sampling equipment depends largely on local conditions (e.g. depth of the samples, size of the ship, power and capabilities of the lifting gears, bottom conditions, etc.). Coastal mysids, especially in hard bottoms (or caves) can also be sampled by Scuba using diver-operated specially-designed suction bottles or plankton nets. Many mysidaceans species exhibit pelagic phases and perform vertical migrations between the bottom and the surface, mainly at night. Pelagic samples can be taken with plankton nets (such as rectangular mid-water trawl, obliquely Tucker trawl, or vertically plummet net). However, coastal and oceanic strictly pelagic species are relatively few in number.

Mediterranean mysidaceans are composed of 38 genera and 102 species. Four of the seven mysidacean families are present in the Mediterranean Sea, lacking the family Petalophthalmidae (order Mysida) whose species are restricted to deep waters. The two families of the order Stygiomysida are a small group of cavernicoulus mysids endemic of caves in Central America and Mediterranean fresh waters caves but without known representation in the Mediterranean marine environment. A complete list of genera and species of the Orders Lophogastrida and Mysida at present known from the Mediterranean marine waters, their geographical distribution, and their habitat based on current information are provided in Tables S19-20. The Mediterranean mysidacean fauna consist of 37 endemic species, in addition to 48 species that also occur in the north-eastern Atlantic, and 18 cosmopolitan species. At present there are few invasive species in the Mediterranean Sea: the Ponto-Caspian mysid Hemimysis anomala, and the Atlantic species Neomysis integer, both recently detected and confirmed, respectively, in the estuary of the Grand Rhône [166]. There is a decrease in mysidacean species richness from the west to the east basins (90 known species in the western basin versus 55 in the eastern) (Figure S3). Isopoda, Cirripeda, Amphipoda and Decapoda also show a general higher species richness in the western basin (Table 2).

The mysidacean fauna of the Mediterranean Sea is also considered one of the best known faunas of the world [145]. However, while the marine mysid fauna of the north-western Mediterranean and the Tyrrhenian Sea (especially the Gulf of Naples) are the best known, various parts of the African coasts and of the eastern sector of the Mediterranean Sea have been little or no studied (Figure S3). The temporal trend of new species descriptions show a climbing curve that suggest that there is still a large number of unknown species to describe in this sea (Figure 13c).

The impact of anthropogenic activities in the Mediterranean mysidacean is poorly documented. There are some evidences of significant changes in the mysidacean fauna in areas under strong anthropogenic pressure, thus most Mediterranean coasts. In some areas of the north-western basin there are indications of species substitutions of the genus Hemimysis, probably caused by global climate warming [175]. In the gulf of Naples, as a result of the eutrophication of waters and other anthropogenic coastal pressures, local population extinctions or strong area regressions have been described [176]. Unfortunately, many of the supposed new species could likely become extinct before we even know of their existence [147,176]. Biodiversity of Mediterranean lophogastrids and mysids is therefore immersed in a critical point of knowledge and scientists must work against time with the aim of not loosing such valuable biological information.

(iii) Decapods: regarding their diversity, several early regional studies included substantial information [e.g. 150,151,152,153,154,155,156,157]. The first treatise on the whole Mediterranean decapod fauna was published by Heller [158] and listed 154 marine species. In the 20th century important monographs updated the knowledge of regional faunas [159,160,161]. In the last twenty years our knowledge of the decapod fauna has progressed further, often as a by-product of the fishery surveys carried out to monitor the status of Mediterranean fishery resources that include high valued shrimps. Currently, there is a complete compilation on decapods of the Mediterranean Sea and north east Atlantic Ocean [162].

Overall 383 species, of those 309 autochthonous species in 63 families, have been reported in the Mediterranean Sea for decapods (against 480 species in 79 families in the Ibero-Mauritanian sectors of the Atlantic Ocean). The present autochthonous Mediterranean decapod fauna mirrors that of the temperate east Atlantic, even if it is significantly impoverished. This is due to the main connection of the Mediterranean to the eastern Atlantic Ocean through the Gibraltar Strait and that Mediterranean decapod fauna, after the Messinian salinity crisis, was rebuilt mainly from the east Atlantic stock of species. Forty Mediterranean autochthonous decapod species are endemic, and 12 of these are still known only for the type locality and the original description (Table S21). The cut of the Suez Canal in 1869 has restored the connection with the Indian Ocean, and since then, we have witnessed an exponential increment in the number of Indo-pacific decapod species recorded in the eastern Mediterranean [167]. Some of them have become locally valuable fishery resources [168], others are deemed responsible for the reduction of the populations of some autochthonous species [169]. The arrival of a large number of Lessepsian immigrants in the Levantine Sea has significantly modified the biodiversity of its decapod fauna. Currently, the Mediterranean is hosting 74 non indigenous decapod species. The autochthonous decapod species recorded in the Levantine Sea are only 60% of those reported for the whole Mediterranean Sea, even if the intensified research effort in the Aegean and Levantine Seas in the last thirty years has proved the species richness of the autochthonous decapods fauna is higher than previously supposed.

Even if decapods of the Mediterranean Sea are also regarded among the best known, new taxa continue to be discovered as a result of revisions of “difficult” genera, as Anapagurus by García-Gómez [170], or of the use of more appropriate techniques to collect burrowing species that are seldom obtained with traditional sampling gears [171].

Multiple anthropogenic causes, such as over-exploitation of fishery resources, coastal pollution, maritime traffic, etc., threaten the Mediterranean decapod fauna. The joint research effort of taxonomists and ecologists may help to mitigate the losses of the original Mediterranean biodiversity. Global warming may in the next future significantly affect both the distribution of these species living in coastal waters and the arrival of new Lessepsian migrants in the eastern basin. Actually a small population of Ocypode cursor has already been recorded on a Sicilian beach and the stock of Crangon crangon, a fishery commodity, in the northern Adriatic collapsed in the last twenty years. The dramatic decrease of this boreal species has been in parallel by the increment of the penaeid shrimp Melicertus kerathurus a thermophilic species unknown in the area until a century ago [see 158,159]. The changes we witness may be explained also by other factors that act synergistically, and highlight the importance of a continuous monitoring of the biodiversity in the Mediterranean Sea.

- Bryozoa

Species of this phylum are abundant and diverse in the sea bottom of the Mediterranean Sea, mainly on hard substrata. However, due to its comparatively small size, they are very often overlooked or misidentified. Bryozoans live in different substratum on which the colonies are settled: rocks, algae or other animals like mollusk shells or corals. Few species live on soft bottoms, where they usually settle on small hard pieces of substratum (small rocks or shells) that lie on the bottom. The relatively well studied western European coast of the Mediterranean has produced an abundant bibliography (Table S22).

Bryozoans have been sampled by extraction of the substratum on which the colonies are settled: rocks, algae or other animals like mollusk shells or corals, because the detaching of the colonies often destroy or severely damage them to the point of becoming unrecognizable. The observation and annotation of colonies without extraction, directly or by photography, is suitable for middle to large sized colonies of well known and easily recognizable species, which are not the most. Therefore, the sampling by trawls or dredges that do not include hard substrata tends to underestimate the bryozoans diversity of an area. Most species are determined in the laboratory with a stereomicroscope, because their usually well developed calcareous skeleton is very distinctive, but the use of a Scanning Electron Microscope (SEM) is necessary in many cases due to the small size of the zooids and the overall similarity of some species.

A total of 389 species of bryozoans have been cited in the Mediterranean and 88 species (23%) are endemic (Table S22). Of these ones, 53 species belong to the order Cyclostomata (with 17 endemic species), 44 species belong to Ctenostomata (with five endemic species), and 292 species are of the order Cheilostomata (with 66 endemic species). However, some citations may be doubtful and the taxonomic status of some of the species cited may need updating. In addition, the qualification of endemic is provisional, given that the neighbouring Atlantic waters and a great part of the Mediterranean ones are not well studied.

The most extensive works about Spanish bryozoan are those by Zabala [177] and Zabala and Maluquer [178], which, although mainly devoted to Catalonia, suppose a revision and updating of most previous information about the Mediterranean bryozoan. The Straits of Gibraltar and Alboran Sea have been studied in the last 20 years by several authors [179,180,181,182,183,184,185,186,187,188,189,190]. The French coasts are also well studied, with continuous new descriptions since the 19th century [191,192,193,194,195,196,197]. The Italian coast has been less studied but there is some noticeable work [198,199,200]. Moreover, the bryozoans of the Adriatic Sea have been recently revised [201]. On the contrary, the African coasts have been scarcely studied, although some works, mainly from French authors were done along these coasts in the first half of the 20th century [e.g. 202]. The eastern basin has been poorly studied as well, except from some occasional works [203,204].

However, not only a geographical bias exists, but also a taxonomic one, which is not exclusive of the Mediterranean Sea, but extensive to the general studies about bryozoans. The order Cheilostomata is the best studied due to it is the most diverse one, with about 80% of the known species, and species are easier to identify due to their high polymorphy of zooids, which are modified for different uses, and differ greatly between species even when they are closely related. On the contrary, the other two marine orders, Ctenostomata and Cyclostomata, have technical difficulties that make them harder to identify. Ctenostomata species are soft and give few external characters. Cyclostomata are well calcified, but the colonies are usually very small and cryptic, and often, if they are not reproductive, they cannot be morphologically identified because some distinctive characters occur only in the gonozooid, which is not present in young specimens or no reproductive colonies. The works on these two orders are thus very valuable, like the worldwide revision of Ctenostomata [205] and the works on Cyclostomes [185,187,188,195]. In addition, some general works such as Zabala [177], Zabala and Maluquer [178], Álvarez [186]; Álvarez [189], and Hayward and McKinney [201] include interesting revisions.

Little is known about threats to bryozoans populations in the Mediterranean Sea because many species have been seldom found. The cryptic habitats and small size of most colonies make an evaluation of risks difficult. But they share the fate of their habitats and where the coast is environmentally degraded, the populations of bryozoans may suffer the same effect. A case of massive mortality of invertebrates, including bryozoans, has been documented in the Provence related with a month of high temperature [206]. Besides, some species (i.e. Pentapora fascialis, Retepora spp) are also impacted by diver frequentation [e.g. 207]. However, there are also some species that are favoured by human activities, mainly the fouling ones like Bugula neritina, Schizoporella errata or Watersipora subovoidea. These and other ones are usually intertidal or shalow sublitoral species that may settle on artificial substrata and be dispersed by ships and debris. Bugula neritina, for instance, is a currently cosmopolitan species of Indo-Pacific origin which was spread by ships during the 19th and 20th centuries, and is now among the most abundant bryozoans in European and Mediterranean shallow-waters, especially well installed in harbours. One species of bryozoan, Hornera lichenoides is included in the Annex II of the Barcelona Convention.

- Echinodermata

A detailed revision of the relevant literature was carried out and information regarding the taxonomy and geographical distribution of the Mediterranean species of echinoderms was collected (Text S4). Tortonese [208] and Koukouras et al. [209] reviewed the Mediterranean echinoderm fauna. Since then, some additional information on the taxonomy and the geographical distribution of the Mediterranean echinoderm species has been published [210,211,212,213,214]. We also used the data collected from 190 stations in the Aegean Sea and Cyprus (0-1,250 m depth). Samples were obtained using fishing nets, dredges, grabs and by free or SCUBA diving. All echinoderm specimens were identified to species level and deposited at the Museum of the Department of Zoology, Aristotle University of Thessaloniki (Greece). Based on the literature and sampling data we updated the available checklist and their general distribution in the Mediterranean Sea.

Tortonese [208] reviewed the Mediterranean echinoderm fauna and reported 143 species (five Crinoidea, 30 Asteroidea, 34 Ophiuroidea, 26 Echinoidea and 48 Holothuroidea). Koukouras et al. [209], based on new information, raised the number and updated their distribution in the Mediterranean Sea. We estimated that the echinoderm fauna of the Mediterranean Sea is currently composed of 154 valid species (five Crinoidea, 33 Asteroidea, 34 Ophiuroidea, 28 Echinoidea, 54 Holothuroidea) (Table S23 provides with information on their depth distribution and detailed references). Species richness in each one of the main geographical areas of the Mediterranean Sea is given in Table 2 and Figure S4. Differences between areas can be discussed in terms of water masses and circulation [215,216] along with data on temperature and salinity variations [217], and geographical aspects [218,219]. Most of the Mediterranean echinoderm species (67.7% of the total Mediterranean species number) have an Atlantic-Mediterranean distribution, while 37 species (24%) are Mediterranean endemics. The echinoderm fauna of the Levantine is enriched by 5 Lessepsian migrant species, one of which Synaptula reciprocans has expanded its distribution in the Aegean Sea (Figure S4 and S5).

Echinoderms richness showed that the western Mediterranean hosted 93.5% of the known Mediterranean species and displayed the highest species richness among all other areas (Figure S5). The Central Mediterranean came fourth in echinoderm species richness among the Mediterranean areas (63.6%). However, it should have had a higher species number compared to the Aegean Sea and the Adriatic due to its direct neighbouring with the western Mediterranean. The rather low number of echinoderm species from this area is attributed to the limited sampling effort. The Adriatic Sea hosts 101 echinoderm species (65.5%). The Aegean Sea, although more distant from Gibraltar, displayed a higher echinoderm species richness in relation to the Adriatic and the Central Mediterranean (69.4% of the species). The Levantine Basin displayed the lowest richness (47.4% of the species).

In general terms, the Mediterranean echinoderm fauna is well studied, however there is a lack of relevant information from the southern Mediterranean coast due to less intensive sampling effort [219], as well as there is a lack of knowledge concerning the Mediterranean deep-sea echinoderm fauna [212]. The acquisition of a concise, detailed view on the Mediterranean echinoderm fauna is often limited by certain taxonomical problems. The descriptions of certain rare endemic species, such as the ophiuroid Pectinura vestita, are old and incomplete and these species have not been re-collected ever since. Thus, various records of echinoderm species from the Mediterranean should be considered doubtful since the respective identifications have been carried out in the framework of benthic ecological studies and have not been checked by taxonomy experts. In addition, many records in the literature are given under older, invalid names [209].

Climate change is considered a major threat for the Mediterranean marine biodiversity. Recently, it has been demonstrated that the entrance rate of the Lessepsian decapod, mollusk and fish species in the Mediterranean has been accelerating as a result of the increase in the mean temperature of the Mediterranean waters, a reflection of the global climate change [220,221,222]. Furthermore, for the same reason, the dispersal rates of the Lessepsian species towards higher geographical latitudes are also increasing. In this context, the Lessepsian holothurians species Synaptula reciprocans seems to be quickly expanding its distribution in the Mediterranean since it has been recently reported from the Dodecanese and Cyclades Islands (south Aegean Sea) [223], while till 2007 it was known up to Rhodos I. [209]. Future research may be focused to study the entrance and dispersal rates of the Lessepsian echinoderms in the Mediterranean and their potential impact on the native fauna.

Certain echinoderm species constitute an important fishery resource. The sea urchins Paracentrotus lividus and Sphaerechinus granularis can be usually found in Mediterranean fish markets since their gonads are regularly consumed. Certain holothurians, such as Holothuria tubulosa, are commonly used as fishing bait, while different species of asteroids and echinoids are used for decoration. However, there is no enough information about threats to echinoderm species, although they can have important ecological roles: for example, sea urchins are important in structuring the assemblages in shallow hard-substrate areas through grazing, and they may drive the transition from erect macroalgal assemblages to coralline barrens [224,225, and references therein].

- Sipuncula

The phylum Sipuncula is one of the minor worm phyla, closely related to annelids. The last comprehensive revision of the Mediterranean sipunculan fauna was by Pancucci-Papadopoulou et al. [226]. Since then, only a few additions have been published [227,228,229,230,231]. Here we reviewed available information to update the list of sipunculans in the Mediterranean with published and unpublished identifications.

A total of 34 species and 4 subspecies of sipunculans arranged in 9 genera and 5 families were recorded in the Mediterranean Sea (Table S24). The more ubiquitous species are Sipunculus nudus, Golfingia elongata, G. vulgaris, Onchnesoma steenstrupii, Phascolosoma granulatum and Aspidosiphon muelleri. By contrast, very rare species are Nephasoma constricticervix, Phascolosoma perlucens, Apionsoma trichocephalus, N. sp. cf. flagriferum, P. turnerae and the subspecies G. vulgaris antonellae. The last subspecies is endemic of the Mediterranean Sea, whereas the last species is recorded for the first time for the investigated area by using unpublished material (J.I. Saiz, personal communication, Table S24).

Cluster analysis (Figure S6) shows that Mediterranean biogeographical sectors can be placed together into two main groups with a similarity level of 61%. The smallest group includes the three Adriatic areas plus the ‘Gulf of Lyon and Ligurian Sea’ sector. By contrast, the largest group of the dendrogram comprises the remaining 6 sectors located both at the western and eastern Mediterranean. The main species responsible for this dichotomy are N. diaphanes diaphanes, A. murinae bilobatae, and Thysanocardia procera, which are well represented in the largest group of the dendrogram, whereas G. margaritacea is almost the only species well represented in the smallest group.

Mediterranean sipunculans represent almost the 25% of the global sipunculan diversity. This percentage is relatively low, since the large diversity of the phylum corresponds to warm shallow tropical areas [232]. At the level of families, almost all sipunculan families are represented in the Mediterranean, with the exception of Themistidae. Concerning genera, nine of the 17 genera of sipunculans are represented in the Mediterranean. Some of the absent genera are monotypic or very restricted in their global distribution to warm waters [233]. Only two exotic species have been described so far, and there is only one subspecies considered to be endemic: Golfingia (Golfingia) vulgaris antonellae [227].

Regarding species data by region (Table 2) we observed a latitudinal gradient from the north to the south of the Mediterranean and west to east, linked to the temperature of the water masses along the year [234]. Thus, the dendrogram obtained (Figure S6) may be reflecting a physiological barrier for sipunculans with cold versus warm species. In fact, G. margaritacea is mainly a temperate and boreal species [226] and its presence in the Mediterranean may be indicating the prevalence of colder water masses. By contrast, other termophilic species, such as Ph. convestitum and A. elegans have been proposed as Lessepian migrants [226,230]. In this way, some other rare records of Phascolosoma and Apionsoma could be further candidates to migrants.

The spatial analysis of sipunculan diversity shows that African coast is especially undersampled. This also applies to the abyssal zone (> 3,000 m) where only three single records (N. diaphanes corrugatum, Ph. tuberculosum and A. murinae murinae) are published [235,236].

The only reported endangered sipunculan species is Sipunculus nudus, which is collected massively along the Spanish littoral as bait for fishing [237].

- Meiobenthos

Marine sediments hold an abundance of microscopic life, the smallest of which attach to individual sand grains or live in the interstices between grains. A variety of bacteria, archaea, and protists share this habitat with minute metazoans, the meiofauna, a major component of seabed ecosystems, particularly in the deep-sea. About half of the animal phyla are represented in the meiofauna, and some (e.g., Loricifera, Kinorhyncha) are confined to it. Nematodes are typically the most numerous component, with harpacticoid copepods, and foraminiferans also important.

The study of the diversity of main species in the large meiobenthos group (mainly composed of small benthic invertebrates that that can pass through a 0.5 mm mesh but will be retained by a 32 μm mesh and live mainly in the sediment) is a difficult and time consuming tack and therefore most studies have been dealt with higher taxonomic levels [238]. The Mediterranean Sea is not an exception, and species or genus level ecological data are scarce. In this study, nematodes and benthic (Harpacticoida) copepods were investigated, since these two groups are the dominant ones in the most of the cases (Text S5). In addition, among the various meiobenthic organisms, living soft and hard shelled benthic Foraminifera are usually equally important with nematodes, and these two taxa together usually account for over 90% of the meiobenthic community [239]. We therefore include some sparse information available about Foraminifera, as well as about Gastrotricha.

Most of the early qualitative work on free-living marine nematodes in the Mediterranean was summarized by Allgen [240] and Schuurmans Stekhoven Jr [241]. Schuurmans Stekhoven Jr [242] compiled a list of all the species found to the date from the Mediterranean and reported 143 species, 106 of which were new to science. Significant work in the past provided a series of taxonomic works on nematodes, which contributed to the knowledge of nematode biodiversity [e.g. 243,244,245]. De Bovée [246] studied the nematode populations in sublittoral terrigeneous muds off Banyuls-sur-Mer during an annual cycle, and reported a much higher number of species (184). More recently, Danovaro and Gambi [247] investigated the nematode assemblages in a Posidonia oceanica bed of the north-western Mediterranean over an annual cycle. High diversity values were correlated with high concentration and high heterogeneity of the food sources indicating that biodiversity is closely coupled with changes in food availability. In the eastern Mediterranean, only few studies have dealt with the lower taxonomic composition of nematodes (family, genus, or species), and several are yet to be published. Wieser [248] studied the meiobenthic nematodes of the Piraeus harbour area (south Aegean Sea) and identified 44 different species of which nine were new to science. Lampadariou [249] studied the nematodes of the continental shelf of the Cretan Sea and some deep-sea areas of the north and south Aegean Sea and found approximately 280 different species, many of which were undescribed. Nematodes currently listed more than 700 species in the Mediterranean Sea [31], and the most important taxonomic papers that contributed to the knowledge of nematode biodiversity are listed in Text S5.

Regarding harpacticoid copepods, there is no clear picture on their actual biodiversity since most studies have been mainly of taxonomic nature [e.g. 250,251,252]. However, the taxonomic studies from the Mediterranean suggest that copepod diversity might actually be high and that many new species are yet to be described. Steuer [253] investigated five different stations near El Shatby in Alexandria. Mitwally and Montagna [254] also studied the harpacticoid copepods along three sandy beaches in Alexandria and found nine species, among which, two members of the Ectinosomatidae, were new to science (Arenosetella bassantae and Noodtiella toukae). In the western Mediterranean, Soyer [255] studied the populations of harpacticoid copepods on the continental shelf off the coast of Albères between t and 130 m depth and found 254 species. In a study on the harpacticoid copepod populations from the eulittoral zone of a sandy beach on Crete, Stobbe [256] found 12 species. The community was dominated by Psammotopa phyllosetosa. Sevastou [257] also studied the harpacticoid copepod populations from two geographically spaced sandy beaches in Crete applying a 13-month sampling design and found 96 species, which outreached by far any species richness recorded in previous studies of comparable habitats.

Until the late 1990s, the study of benthic foraminifera in the Mediterranean has been restricted to a small number of samples collected from specific sites [258,259]. More recently, a larger number of investigations have considered the importance of living benthic foraminifera in ecological studies [e.g. 260,261,262], however they are still far from being well studied. The knowledge recently gained indicated that in the Mediterranean the foraminifera are highly diverse and consist with more than 600 species. A large number of these species belong to Lessepsian invaders [11], which is however still a matter under debate since data on the biogeography, diversity and ecology of autochthonous shallow-water faunas from various carbonate environments of the Mediterranean Sea are limited [263].

In addition, the studies on marine gastrotrichs in the Mediterranean were conducted mainly through a programme of faunistic and taxonomic research of the Italian seas [264,265,266,267,268,269,270] and in Crete [271]. All these studies revealed approximately 150 species from the Mediterranean from a total of 280 and 580 which are known from Europe and world wide, respectively. Hofrichter [31] mentioned 165 species in the Mediterranean Sea.

A small number of pollution studies in the Mediterranean Sea considered the use of meiofauna as potential indicators of anthropogenic disturbance. These studies concerned mostly domestic sewage discharges, pollution in harbours, and fish farm impacts. Marcotte and Coull [272] identified copepod species from five stations placed on transect in the northern Adriatic Sea along a gradient of municipal raw sewage discharge and showed that diversity was very low near the outfall and increased with increasing distance from the source of pollution. Keller [273] investigated the meiofauna communities in a marine area which was highly polluted by the sewage outfall of Marseille. Lampadariou et al. [274] investigated the nematode and copepod community structure along a grid of seventeen stations covering an area from the innermost polluted to the outer clean area of Heraklion harbour, and showed that the nematode community showed a clear zonation according to the degree of pollution and physical disturbance from shipping activities. They concluded that, besides physical factors such as depth, the high level of organic carbon or pollutants such as copper and cadmium played an important role in structuring the nematode communities. A decline in diversity as a result of disturbance caused by fish farming activities was also reported by Mirto et al. [275] from the Gulf of Gaeta in the NW Mediterranean. Setosabatieria, which was the dominant genus, was found to be highly sensitive to organic disturbance as it disappeared completely three months after the deployment of cages. In contrast, other nematode genera, such as Dorylaimopsis, Sabatieria, and Oxystomina, proved to be tolerant and benefited from the new organically enriched conditions.

Information on other marine invertebrates is summarized in Table 1 and Table S1. Total updated registries for the other invertebrate species are of 2168 species (1393 species, excluding Arthropoda).

- Ascidiacea

Ascidiacea comprises the largest Class of the Subphylum Tunicata, a part of the phylum Chordata. Although most abundant at sublittoral rocky communities, they are also adapted to live in abyssal plains [276], and few species are intertidal [277]. The Mediterranean fauna is an important part of the global ascidian fauna of the Atlantic and Mediterranean coasts, which totals approximately 500 species [278]. The ascidians of the Mediterranean Sea have been explored since the second half of the 19th century [e.g. 279,280]. In the first half of the 20th century, important taxonomic works were published [e.g. 281,282,283] that improved the knowledge of Mediterranean ascidian diversity. Pérès [284] listed 130 species in the Mediterranean Sea (although since then some species have been synonymised or have split in several species). Later faunistic and taxonomic studies analysed the diversity of the ascidian fauna in particular areas, especially in the western Mediterranean. In this revision we build on Pérès’ work and updated the records of ascidian species in the Mediterranean Sea, as well as their distribution and affinities.

Fiala-Medioni [285] reported 77 species from the SE of France. Turon [286] found 107 species in the NE Spanish littoral. Ramos-Esplá [287], similarly, listed 117 species in the Mediterranean shores of Spain. Naranjo [288] found 84 species in the area around the Gibraltar Straits, 79 of which were present in the Mediterranean side. Koukouras et al. [289] first reviewed the long-neglected ascidian fauna of the eastern Mediterranean, providing a check-list of 86 species in this basin, out of an estimate number of 187 for the whole Mediterranean. Mastrototaro and Tursi [290] provided a check-list of the Italian fauna of ascidians, including 128 species. Aside from these authors, most recent work on ascidian diversity in the Mediterranean has been done by specialists such as Jean-Marie Pérès, Claude and Françoise Monniot, Françoise Lafargue, and Riccardo Brunetti, among others. Building on the work by Pérès [284], and exhaustively searching all posterior reports, we listed 229 ascidian species for the Mediterranean Sea. Although some of the records are dubious, we have eliminated from our list only those species that were clearly invalid or synonyms of other species (Table S25). Overall, the number of new citations incorporated to the database since 1960 is 22.4 species per decade (Figure 13d). Of these, 52 species (46.8% of additions) corresponded to newly described species, not just new reports.

From a biogeographic point of view, Pérès [284] reported a major contribution of endemic Mediterranean species (50%). The second major component consisted of species of Atlantic-Mediterranean distribution (37.7%). A further 11.5% of the species were cosmopolitan or had a circumtropical range. Only one species (Herdmania momus) was detected as possible Lessepsian migrant at that time. In the area close to the Straits of Gibraltar, the number of Mediterranean endemics falls to 22% of species, while those of Atlantic-Mediterranean distribution rise to 60% [278]. We currently estimated that the endemic ascidians accounted for 34.9% of the species (Table S25). The species with an Atlantic-Mediterranean distribution accounted for another 46.7% (of which 12.2% were found in the Mediterranean only in the western basin); cosmopolitan and circumtropical species made up 14.4% of the total, and nine species (3.9%) were identified as probable Lessepsian migrants. Izquierdo-Muñoz et al. [291] reviewed introduced ascidian species in the Mediterranean, listing 14 species, and Shenkar and Loya [292] reported the occurrence of seven non indigenous ascidian species in the Mediterranean coasts of Israel, many of which are most likely Lessepsian migrants. Particularly worrisome is the recent report of the clubbed tunicate Styela clava in the lagoon of Thau, France [293], given the problems that this species has posed to shellfish industry elsewhere [294]. Although not generally recognized as such, many forms adapted to live in man-made structures, even if they have been present for long times in the Mediterranean, are probably introduced, e.g. Styela plicata and Botryllus schlosseri [295,296].

The composition of the ascidian fauna of the Mediterranean, with species of temperate and subtropical affinities, makes it sensitive to ongoing global warming [77], and a displacement of species of cold-water affinities in favour of termophilic species [297] is expectable. The arrival of introduced ascidian species to the Mediterranean has received considerable attention in recent years. Many introduced species are confined to artificial environments, but other spread outside and colonize natural substratum, turning into invasive forms. One clear instance is the solitary ascidian Microcosmus squamiger, which can reach high densities and carpet natural substrates [298]. This species is now common in western Mediterranean [299], and Streftaris and Zenetos [300] listed this species (as M. exasperatus) within the 100 “worst” marine invasive in the Mediterranean.

The trends observed in the known distribution of ascidians from the sixties on indicate that the percentage of endemics is now considerably lower (from 50% to ca. 35%), which is partly caused by the finding of Mediterranean species in adjacent Atlantic waters [e.g. 301]. The number of species known only from one or another basin, which was high and highly skewed towards the western basin, is now much lower and more balanced between basins (Table 2). This is the result of increased knowledge of intra-Mediterranean distribution of many species and, particularly, the rise of studies performed in the eastern basin, which is compensating the previous neglect of this area [e.g. 289,290,302]. Endemic ascidians present at both basins represented 16.9% of Mediterranean ascidians, while 11.3% and 7.4% were endemics reported only in the western and eastern basin, respectively (Table S25).

Ascidian diversity is underreported due to sampling effects, the lack of taxonomists, and inherent difficulties in the taxonomy of the group. Rocky shores in the northern Mediterranean can be considered reasonably well studied, but the Mediterranean African shores still present few studies available [e.g. 303], and may be a hotspot of ascidian biodiversity. This is because this region acts as a refuge for Atlantic-Mediterranean species of tropical affinities, while the northern region of the Mediterranean harbours species with temperate and even boreal affinities. Moreover, much remains to be known about the ascidian fauna in deep waters, or in soft-bottom communities. The ascidians themselves pose important problems for identification, due to the lack of distinctive features in some groups, overlapping of characters between species and, particularly, because observation of characters requires specimens adequately preserved in a relaxed state and ripe, which are hardly available. It is a group where molecular tools can substantially contribute to help taxonomic work. Available studies that have incorporated genetic markers to the research on Mediterranean ascidians showed that cryptic speciation may be much commoner than usually recognized [e.g. 304,305,306].

Ascidians are subjected to different threats such as habitat destruction, degradation, and pollution, as well as global warming, arrival of invasive species, exploitation and other factors [e.g. 307,308]. Mass mortalities have also occurred at some occasions; for instance, an undescribed illness devastated the populations of Microcosmus sabatieri in the early 1990s at least along Spanish and French shores. This species was once the most abundant solitary form in the shallow sublittoral and reached abundances of approximately one individual per square meter in NE Spain (X. Turon, personal communication). It is nowadays extremely rare. This is significant because in the Mediterranean, large pyurid species such as M. polymorphus, M. sabatieri and M. vulgaris have been consumed since the 1st century AD [309]. M. sabatieri is still abundant in the eastern Mediterranean, where it sustains a small-scale fishery [309,310]. Threats to ascidian biodiversity have also biotechnological implications, as ascidians are producers of some of the most promising anticancer compounds found to date in marine invertebrates [311]. They have originated one drug already marketed (Trabectedin, sold under the brand Yondelis®), obtained from Ecteinascidia turbinata, a circumtropical species that used to be farmed in the Balearic Islands before a synthetic production of the drug was achieved in the 1990s). A second compound, Aplidin®, obtained from the Mediterranean species Aplidium albicans is in advanced clinical trials.


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