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DEEP RESULTS OF THE 2016 EXPEDITION EXPLORING SUBMARINE CANYONS Towards Deep-Sea Conservation in Lebanon Project

March 2018

DEEP SEA LEBANON RESULTS OF THE 2016 EXPEDITION EXPLORING SUBMARINE CANYONS Towards Deep-Sea Conservation in Lebanon Project

Citation: Aguilar, ., García, ., Perry, A.., Alvarez, ., Blanco, ., Bitar, . 2018. 2016 Deep-sea Lebanon Expedition: Exploring Submarine Canyons. Oceana, . 94 p. DOI: 10.31230/osf.io/34cb9 Based on an official request from Lebanon’s Ministry of Environment back in 2013, Oceana has planned and carried out an expedition to survey Lebanese deep-sea canyons and escarpments. Cover: membranaceus © OCEANA All photos are © OCEANA 06

Introduction Index 15 14 14 15 12 11

Methods Data analyses Infaunal surveys Rov surveys Collaborations measurements Oceanographic 42 42 42 42 40 40 38 36 31 30 29 27 20 20 19 16 16 16 24 22 22

Results bottoms Sandy-muddy Annelids Bryozoans Molluscs Cnidarians Sandy bottoms beds and rhodolith/maërl Coralligenous habitat Habitat types Bathyal muds Canyon heads Rocky bottoms 46 46 45 44 44 44

Areas St. George Sayniq Tarablus/ 49 49 47 47

impacts Other observations Fisheries Marine litter 58 57 56 52 67 65 64 64 59 58 56 54 53 52 51 50 65

conclusions Discussion and Survey ar Fossil r concretions Oyster aggr Sea penfields Gorgonian/sea fan ) Scleractinians (Stony aggr Key community findings Main expedition Acknowledgements Conclusions Other thr Thr Main thr communities dominated - communities rhodolith/maërl beds. concretions and Coralligenous ecological importanc types andtheir eats observed eefs eats eats eas egations/ egations

84 able A1.Listof 84 68 68

Annex 1 species identified of threatened species identified group. broad taxonomic according to identified taxa, Number of T Figur able A2.List A1. 85 85 86 86

Annex 2 survey areas interest of conservation assesing r Methodology for Annex 3 References elative

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Biodiversity of the Eastern

The Mediterranean Sea is a semi-enclosed basin, surrounded by 21 African, Asian, and European states. It extends over an area of 2.5 million km2 and is connected to the Atlantic though the Strait of , to the through the Dardanelles and Bosforus Straits, and to the through the artificially opened Suez Canal. When compared to the average depth of the world’s (3733 ), the Mediterranean Sea is relatively shallow, with a mean depth of ca. 1650 m1, yet it is the deepest of the enclosed . The entire Mediterranean Sea is considered a biodiversity hotspot, hosting up to 18% of the global total of macroscopic species (in less than 1% of the global ocean surface), with 25% to 30% of them considered endemic.2,3

The Eastern basin of the Mediterranean is defined as lying to the east of the . It encompasses the Adriatic, Aegean, Ionian, and Levantine Seas, and includes the deepest waters of the Mediterranean (5121 m), in the Hellenic Trench.4 The nutrient concentration of the Mediterranean is relatively low, and it is more

Introduction oligotrophic or ultra-oligotrophic in the Eastern basin, where the exhausted surface layer is thicker and deeper due to stronger stratification.5 In the , the easternmost of the Mediterranean, temperatures are also more extreme than in the rest of the Mediterranean, such that it is also referred to as the ‘warm-water corner of the Mediterranean’.6 This characteristic appears to have become more pronounced during recent decades, possibly due to climate change.7,8,9 The relatively warmer waters of the Levantine may be responsible for the fact that, even though community composition throughout the Mediterranean Sea is relatively homogenous, some important habitat‑forming species are absent from Levantine waters (e.g., seagrass and sea fans such as Paramuricea clavata and Eunicella spp.).10,11,12

Within the Levantine Sea, the waters of Lebanon host a wide variety of , in an area representing 0.005% of the global marine surface. These ecosystems range from shallower features such as coralligenous habitat, seagrass meadows and vermetid reefs, to deep-sea ones such as underwater canyons.13 Lebanese offshore waters are characterised by a narrow , which is perpendicularly crossed by various canyon systems that connect coastal zones to deep‑sea habitats. These geographic features range approximately from 50 to 1600 m deep in Lebanese waters, and are important ‘keystone structures’ due to their role in supporting deep-sea communities acting as nursery and shelter habitats, benefiting fisheries, enhancing carbon sequestration, and providing other goods and services to human society.14 pipelines andthree powerstationsrunonfuel. four commercial ports,15fishing12oil concentrated onthecoast.Intotal,Lebanonhas Active industrial and commercial areas are unregulated manner, particularlyduringtheperiod progressed incoastalareas inachaoticand of resorts andindustrialdevelopment,has Coastal development,including theconstruction represents 8%oftheterritorialarea ofthecountry. activities are concentratedalongthecoast,which entire Mediterranean.Majoreconomicandsocial perspective, similarpressures andthreats asthe Lebanese marine ecosystems face, from a general toLebanesemarinelife Threats and newspeciescontinuing to bediscovered. species richnessestimatedinthethousands, supports adiversityofmarinelife,withtotal The arrayofmarinehabitats in Lebanesewaters common. to occur, withthefirsttwobeing relatively and sea turtles(i.e.,Carettacaretta, irregularly spotted.Three speciesofthreatened and monk seal (Monachus monachus) has been coeruleoalba) are foundinLebanese waters, physalus), andstripeddolphins(Stenella macrocephalus), fin (Balaenoptera cetaceans suchasspermwhales(Physeter be atriskofextinction.Forexample,threatened are variousiconicspecies that are considered to Among thespeciesknowntooccurinregion of coastalareas unsustainable. from different origins,makingthecurrent uses declining duetoover-exploitation andpollution of theLebanesecoasthasbeendescribedas this area and compete for space. Thus, the status As aresult, manycommercial interests influence population, concentratedincoastalurbanareas. , asithosts55%ofthetotalLebanese The coastal zone suffers severe anthropogenic rhinobatos). oxyrinchus) andcommonguitarfish(Rhinobatos Endangered speciessuchasshortfinmako(Isurus are also reported from the area, including Critically impacts suchastheeffect ofclimatechange. urbanisation, invasive species, and larger-scale habitat degradation,recreational uses,coastal dumping, unsustainable and illegalfisheries, andgravel)extraction,sewageoil include, among others: marine aggregate (i.e., Dermochelys coriacea) have been reported 13 Various threatened sharksand rays 16 19 Chelonia mydas These activities 18 13,15 18 18 17

gear), withdeleteriouseffects onmarinefauna. (i.e., plasticbags,otherdebris, lost fishing discarded, including non-biodegradable plastic Most solid waste along the Lebanese coast is regular waterflux,increasing sanddeposition. while structures like marinas have disrupted affected landscapesandthreatened biodiversity, of theLebaneseCivilWar. problem Nevertheless, overexploitation is arecognised in theMediterraneanare overexploited. 90% oftheassessedcommercial fishstocks Commission fortheMediterranean(GFCM),over fisheries stockstatusfrom theGeneralFisheries According tothemostrecent report on of Lebanon. damaged, includinganature reserve inthenorth A large portion of the coastline was severely release Mediterranean. accidentintheEastern major oil spill, which constituted the largest oil- during the2006conflictwithIsrael resulted ina example, airstrikesonaJiyehoilstoragedepot resulted insignificantimpacts tomarinelife.For pollution, whileepisodicpollutioneventshavealso effluents represent themainsources ofchemical Untreated domestic sewage and industrial data are typicallyscarce andunderestimated. there hasbeenalackoflogbooks,andlandings largely due to a paucity of information; traditionally, precise statusofLebanesestocksisnotknown, overfishing. decreased insizeandabundanceasaresult of suggested thatsomenative fishspecieshave vulgaris 15 andsurveysoflocalfisherman have 23 17 This developmenthas 21 The 22 20 13

7

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The artisanal or traditional fisheries sector Current marine protection represents the dominant fishery sector in Lebanon, and the main gears used include trammel nets, The Lebanese government has signed or ratified gill nets, longlines, purse seines (used together various international conventions and agreements with lampara – floating lamps used at night to that directly affect the protection of its waters. attract larval and juvenile fishes to the nets24) and These conventions include: the Convention for beach seines.22 Some of the most destructive the Protection of the Mediterranean Sea Against types of fisheries are not carried out in Lebanese Pollution (Barcelona Convention); the Convention waters (i.e., ), as the is not on Biological Diversity (CBD); the United Nations suitable for it. Convention on the Law of the Sea (UNCLOS), the Convention on Wetlands (Ramsar Convention); the The fishing fleet comprises around 2600 African-Eurasian Migratory Waterbird Agreement registered fishing boats (based on the most (AEWA); the Agreement on Conservation of recent available data, from 201521) with total Cetaceans of the Black Sea, Mediterranean Sea lengths of less than 12 m.22 There are more and contiguous Atlantic area (ACCOBAMS); than 80 species of commercial interest, and an and the UNESCO Convention Concerning the average of 3574 t were landed annually from Protection of the World Cultural and Natural 2000-201321). Moreover, the general disorder that Heritage (World Heritage Convention). prevailed during the period of the war resulted in an anarchic use of marine resources, and some At the national level, Lebanon has protected two illegal practices reportedly still take place (i.e., marine areas, covering only 0.22% of national dynamite fishing, spear fishing, and undersize waters (Map 1): Palm Islands Park and Nature mesh), contributing to the depletion of stocks Reserve and Tyre Coast Nature Reserve. The and habitat destruction.13,22 former was created in 1992 and covers an area of 5 km2. It is formed by a group of three islets The Levantine part of the Eastern Mediterranean (Palm, Sanai and Ramkin) that lie roughly 5.5 km is particularly susceptible to the invasion of alien northwest of and are important for marine species from subtropical environments, due to turtles that are Red Listed as threatened by the its aforementioned warm temperatures.10 The International Union for Conservation of Nature opening of the Suez Canal in 1869 constituted a (IUCN),26 as such loggerhead turtle (. caretta; corridor for the spreading of thermophilic species, Vulnerable) and green turtle (C. mydas; largely affecting local biodiversity.8 Other activities Endangered), and migratory birds. Endangered such as aquaculture, shipping and leisure boating monk seals (M. monachus) were also regularly are increasing the magnitude of this problem, recorded in the area in the past, but such sightings which is also worsening under climate change, have since become very rare27. It was recognised particularly in the Eastern .13,25 as a Ramsar Site in 200127. The Tyre Coast Nature The spread of invasive species in the region has Reserve was declared in 1998. It occupies an resulted in observable impacts on the productivity area of 3.8 km2 and consists of private agricultural of Lebanese coastal marine ecosystems. For land, freshwater springs and the last remaining example, 37% of total catches in the region of sandy beach ecosystem in the country28. As with Tyre (south Lebanon) in 2008 were alien species, the Palm Islands, its geographical location places whereas this proportion did not exceed 14% in it along a major migratory route for birds, and for the entire Eastern Mediterranean in 2012.22 nesting sea turtles28. It has been designated as a Ramsar Site29, an Important Bird Area (IBA), and a UNESCO World Heritage Site.

The artisanal or traditional fisheries sector represents the dominant fishery sector in Lebanon. Map 1. MarineProtected Areas inLebanon. 9

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Peltaster placenta

Despite the demonstrated effectiveness of MPAs A general obstacle to MPAs designation and as a tool for marine protection30, the percentage of management is a lack of detailed information about marine area under this designation remains below and species. In the Mediterranean, internationally agreed targets, globally, in the such information is still relatively deficient, despite Mediterranean Sea. According to the CBD Aichi advancements in marine research technology Target 11, at least 10% of coastal and marine and recent efforts to prioritise the study of marine waters should be protected by 2020;31 however, ecosystems. In the Levantine area, the need for in the Mediterranean Sea, only 7.14%21 has been marine data is particularly pronounced for the legally protected (and only 4.29% if the extensive northeastern basins of the Levantine area, in Pelagos Sanctuary is excluded). Most marine comparison with the central and western basins11. ecosystems represented in the Mediterranean This lack of data is even more pronounced when network of MPAs are those associated with it comes to deep-water ecosystems, which are shallow, coastal waters, while valuable deep- known to contribute significantly to the total sea ecosystems are typically excluded, which biodiversity of the Mediterranean.34 leads to an MPA network which is neither coherent nor ecologically representative. This Closing this knowledge gap is the primary aim of situation is especially prevalent in the Eastern and the Deep-sea Lebanon project. Providing first- Southern basins of the Mediterranean, which are hand information about deep-sea ecosystems to characterised by small and dispersed MPAs along the Lebanese authorities will help to increase the the coast.32 Nevertheless, the deep-sea canyons protection of Lebanese waters, and specifically from the Lebanese and Syrian have already of ecosystems found in deep areas. Ultimately, been identified as an Ecologically or Biologically this increased protection will also contribute Significant Area (EBSA) by the CBD, namely to reaching Aichi Target 11 by 2020, and to East Levantine Canyons, thus acknowledging strengthening the natural marine biodiversity their important role in ecosystem functioning, corridor in the Eastern basin. their uniqueness and rarity, and their special importance for -history stages of the species Closing the knowledge that they host, among other characteristics.33 gap about the deep-sea was the primary aim of the expedition. Methods based partlyontheirinclusionwithinthe2011MedNetpr Oceana. Protected Area Strategy canyon, andJouniehcanyonwere includedinLebanon’s Marine as potentialMPAs. Forexample,theBeirutescarpment,St. George areas hadpreviously beenhighlightedasprioritiesforconsideration scientific informationandconsultationwithproject partners.Several Potential siteswere selected priortotheexpedition,basedonavailable 11.4 mextreme breadth. board the coast of the country. Surveys of these zones were carried out on five areas coveringsome of themainunderwatercanyonsthatlieoff period ofnearlyfourweeks(3 Oceana carriedoutaresearch cruiseinLebanesewatersduringa Figure 1. Remotelyoperatedvehicle(ROV)onboard theresearch vessel. description ofthesamplingmethodsisexplainedbelow. conductivity, temperature, anddepth(CTD)instrument.Adetailed areas. Inaddition,oceanographicparameterswere recorded witha sediment sampleswere collectedwithaVan Veen grabinsoftbottom obtain more detailedinformationabouttheinfaunalcommunities, operated vehicle(ROV)withahigh-definitioncamera.Inorder to carried outbynon-intrusive(visual)methods,usingaremotely In eachofthefiveareas, explorationoftheseabedwasprincipally and Sayniq. to south):Tarablus/Batroun, Jounieh,St. George, Beirutescarpment, fromwaters. The final areas north surveyed were as follows (Map 2, Lebanese for datacollectionacrosscentral,andsouthern northern, These areas were selectedtoensure theinclusionofhighpriorityareas meeting inwhichtheyjointlyagreed theareas thatwouldbesurveyed. the LebaneseNationalCouncilforScientificResearch (CNRS)helda UNEP © OCEANA/EnriqueTalledo ‑ Sea Patron,afully-equippedvesselof42moveralllengthand MAP RAC/SPA 36 InSeptember2015,representatives ofOceana,IUCN, 37 , theLebaneseMinistryofEnvironment, and 35 as proposed deep ‑ 27 October 2016),across atotalof 27 October ‑ sea sitesforprotection, oposal by 11

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ROV SURVEYS

Data were collected using a Saab Seaeye Falcon depth, course and time. Scientists onboard Sea DR ROV (Figures 1 and 2), equipped with two Patron viewed the video feed in real time, to carry forward-facing video cameras: (1) a high-definition out preliminary species identification and select camera with 1920x1080 resolution, 1/2.9” Exmor species, habitats, and seabed features of interest R CMOS Sensor, minimum scene illumination of for more detailed investigation. In total, 51 ROV 3-11 Lux, and a 4‑48 mm, /1.8‑3.4 zoom lens; dives were carried out, ranging in depth from and (2) a low-definition camera with resolution 36 to 1050 m. These surveys yielded 71 h 15 min of 540 TVL, 1/2” interline transfer CCD sensor, of video of the seabed, and 4601 still images. sensitivity of 0.35 Lux, and a 1/2” aspherical, Samples of key habitat‐forming species were wide‑angle, fixed-focus lens. During ROV also collected (by means of the robotic arm of the transects, Sea Patron sailed at an average speed ROV) for detailed analyses to confirm preliminary of 0.1‑0.2 knots, filming both in high- and low– species identifications based on the live video definition, and simultaneously recording position, feed.

© OCEANA/ Enrique Talledo

Figure 2. ROV‑launching manoeuvre.

Exploration of the seabed was principally carried out using a remotely operated vehicle down to 1000 m depth. Map 2. Areas surveyedduringthe2016OceanaexpeditioninLebanesewaters,andsurveymethodsapplied. 13

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Figure 3. Preparation of the Van © OCEANA / Enrique Talledo Veen grab for seabed sampling.

INFAUNAL SURVEYS OCEANOGRAPHIC MEASUREMENTS

Infaunal community composition was examined In order to take measurements of different by collecting samples in soft-bottom areas, with oceanographic parameters, a Valeport MIDAS a 12 L Van Veen grab. In total, 12 grab samples CTD + instrument was used, down to a maximum were collected from a depth range of 76‑1015 m depth of 1076 m. The following parameters were and were processed on board. Specimens recorded: conductivity, temperature, pressure, retained on 0.5 mm and 1 mm mesh sieves were turbidity, dissolved oxygen, pH, and kept for identification (Figure 3). chlorophyll-a. The CTD was deployed in all five of the sampling areas, recording parameters at 1 m depth intervals while the CTD unit was descending and ascending, yielding a total of approximately 7300 measurements for each parameter. Examples of the temperature results are presented in Figure 4.

Figure 4. Outcome plot extracted from CTD device, showing the bottom temperature gradient across the areas sampled and the temperature gradient across the depths sampled. of ascidianspecieswere compiled inareport; Dr. Alfonso A. Ramos Esplá, whose identifications experts forconfirmation.Theseincluded: specimens and/orfootagewere senttotaxonomic identification required additional expertise, to thefinest resolution possible. Incaseswhere were alsoidentified (=94 specimens;Figure 5), collected withtheROVandingrabsamples to thefinesttaxonomiclevelpossible.Specimens scientists. All of the visible species were identified recorded bytheROVwascarriedoutOceana Following theexpedition,analysisoffootage DATA ANALYSES CNRS of Environment, and the Lebanese Ministry UNEP project partnersIUCN, and supportof the fullcollaboration was carriedoutwith The research expedition and Ph.D.GhaziBitar, representing UNEP. Rahman HassounandAliBadreddine from CNRS, AbedEl collaborators onboard SeaPatron:Dr. was greatly supportedbythefollowingscientific of theLebanesemilitary. Research workatsea and withpermanentrepresentation onboard Lebanese MinistryofEnvironment, andCNRS, partners IUCN, UNEP the fullcollaborationandsupportofproject The research expeditionwascarriedoutwith COLLABORATIONS various phyla Ph. D. Ghazi Bitar, who identified specimens from JoséTemplado and ÓscarOcaña,Dr. and Dr. documented, andprepared atechnicalreport; Pérez, whoidentifiedmost ofthespongespecies Thierry JeanVaceletMarie Grenier, andDr. Dr. ‑ 40 MAP RAC/SPA, . ‑ MAP RAC/SPA, the 39 38

research cruise. on board SeaPatronduringtheDeep-seaLebanon left) andDr. Ghazi Bitar (UNEP Figure 6. Project collaborators,AliBadreddine (CNRS, sample. Figure 5. Examplesofspecimenscollectedinagrab © OCEANA/EnriqueTalledo © OCEANA/EnriqueTalledo ‑ MAP RAC/SPA, right) 15

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HABITAT TYPES

In total, six main habitat types were documented over a broad depth range (36‑1050 m). These habitats and the key associated communities and species are described below and are presented according to increasing depth. Coralligenous habitat and rhodolith/maërl beds In all of the shallow-water surveyed areas, the presence of dead and live coralligenous habitat and rhodolith/maërl beds was documented (Map 3).

A belt of coralligenous habitat was found between approximately Results 70 m and 90 m depth (Figure 7), which coincided with the edge of the continental shelf and the heads of the submarine canyons, in all surveyed areas. In some cases, coralligenous habitat occurred at shallower depths (roughly 50‑60 m), extending to where it joined with that found on the edge of the continental shelf.

Figure 7. Coralligenous concretions in Tarablus/Batroun, at 74 m depth.

Although some large sponges occurred on these algal concretions, the highest density of massive sponges was found on the rocks and coralligenous habitat on the shelf edge. The most common and representative sponge species was oroides (Figure 8), which usually occurred together with other sponges, such as Fasciospongia cavernosa, Ircina spp., Spongia (Spongia) officinalis, Sarcotragus spinosulus, clathrus, and Calyx nicaeensis.

Figure 8. Agelas oroides on coralligenous formations at the edge of the continental shelf, in Tarablus/Batroun, 77 m depth. Map 3. Distributionofcoralligenousandrhodolith/maërlbedsacross sampledareas. 17

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A diversity of fishes was found in association with maintain the reef structure that is characteristic of these reef formations. Examples of documented coralligenous formations, and therefore provide fish species include anthias, Coris julis, substrate and shelter for numerous species. Muraena helena, Mycteroperca rubra, Seriola dumerilii, and Serranus cabrilla, as as various Other noteworthy calcareous red algae invasive fish species, such as Pterois miles, formations were rhodolith/maërl beds found on Siganus luridus and Sargocentron rubrum. sedimentary bottoms in the upper circalittoral zone, in the areas of Tarablus/Batroun, Jounieh, Significant concentrations of hatpin Sayniq and St. George. As in other parts of the (Centrostephanus longispinus) were found to Mediterranean, these beds represent valuable occur on this of seabed, with the result habitat for many species; other species of algae that it was the dominant species in some areas. and attach themselves to the rhodoliths This species is included in Annex II of the SPA/ (see Figure 9), and numerous small- and medium- BD Protocol of the Barcelona Convention, the sized live within the beds. These list of endangered or threatened species in the include, for example: species of green, brown, and Mediterranean that require strict protection.41 red algae (e.g., Valonia macrophysa, Lobophora variegata, Amphiroa rigida and Rhodymenia sp.); In some areas (i.e., Tarablus/Batroun, St. George bryozoans (e.g., Adeonella pallaisi and Hornera and Beirut), ancient calcareous formations were frondiculata); ascidians (e.g., Halocynthia documented below the coralligenous habitat at papillosa, sp. and Pseudodistoma the shelf edge. These formations may have been cyrnusense); and a variety of other organisms, the result of coralligenous habitat that extended to such as , sponges, hydrozoans, and a greater depth in the past. Although they may no echinoderms. longer contain living algae, they may nevertheless

Figure 9. Valonia macrophysa on a coralligenous bed in St. George, 60 m depth. as those within the genera the order Dyctioceratida (Family:Irciniidae), such species includedmassivesponges belongingto common onsandysubstrate. Inparticular, these associated withhard bottoms,theywere also Although massivespongesare more frequently ellissii. Reteporella spp., particular interest, suchasHornerafrondiculata, several larger species stand out as being of Of thebryozoanspresent inthishabitattype, bryozoans, andhydrozoans. tunicates (particularlyofthefamilyDidemnidae), served assubstrateforavarietyofspecies by meansofbudding.Remainsthisspecies smaller elongatecupshaddeveloped,possibly elongate cup-shapedstructure uponwhichother, species, which in many cases consisted of a large, not uncommontoobservesmallcoloniesofthis hard substrates,eithernaturalorartificial.Itwas muddy bottoms, as well as attached to various wasfounddetachedonsandyandsandy- as , gurnardssuch Serranus cabrillaandS. hepatus sandy bottoms,includingseabassessuchas intermixed with native species that are typical of recorded from thishabitattype.Theywere (Cheilodipterus novemstriatus),were frequently possibly, theIndianOceantwospotcardinalfish cardinalfish (Ostorhinchusfasciatus ) and, (Torquigener flavimaculosus)andbroadbanded Invasive species,suchaswhite-spottedpufferfish types ofbiogenicorartificial remains. algae, smallrocks and/or rubble, orwithvarious coralligenous habitat,andsiteswithisolated combination withrhodoliths,inareas ofscattered They were alsofoundinareas ofmixedhabitat,in and in areas at the head of shallower canyons. zones, indifferent regions ofthecontinentalshelf throughout theinfralittoral anduppercircalittoral Sandy bottomswere widelydistributed Sandy bottoms it maybelongtothegenusAnomocora. Its identification isstill pending,but itappears that previously been described from the Mediterranean. coral was frequently observed which has not In somesandyareas, aspeciesofscleractinian Synodus spp. Lepidotrigla cavilone,andlizardfishes suchas Adeonella pallaisi and Ircinia and Sarcotragus . Caberea 42 This . S. cabrilla andPagelluserythrinus Pyuridae andDidemnidae,fishessuchas mediterranea), ascidiansbelongingtothefamilies such as Mediterranean feather star ( certain organisms. These animals include species created that permit or augment the presence of bottoms, artificialhard substrateshave been and otherobjectsofhumanoriginonsandy Due tothehighabundanceoflitter, metallicwaste, the MediterraneanSea(L. ciliarisandL. sarsi). the twospeciesofthisgenusthatare knownfrom identification, anddoesnotcorrespond toeitherof deserves mention is a sea star belonging to the Another echinodermfrom thishabitattypethat their armsabovethesedimenttocapture food. hundreds orthousandsofindividualsthatlifted Ophiopsila aranea, which formed colonies of most abundantechinodermwastheophiuroid sea starEchinaster(Echinaster)sepositus.The (Hacelia attenuata),andMediterraneanred star (Antedonmediterranea),smoothsea observed included Mediterranean feather inhabitants of sandy-bottomareas. Species Echinoderms were also amongthetypical Anomocora. belong tothegenus Mediterranean. Itmay described fromthe not previously been which has observed was frequently scleractinian coral A speciesof Luidia. Theprecise speciesis pending 19

OCEANA | Deep sea Lebanon 20 In some areas of the deep , ancient reefs had formed hard substrate. These fossil reefs were found in Jounieh and Tarablus Batroun canyons

Sandy-muddy bottoms Rocky bottoms The transition zone between the upper and lower Rocky substrate was found in all of the sampled circalittoral was where muddy bottoms typically areas (Map 4). At the continental shelf edge, in began to appear mixed with sandy ones, before the deep circalittoral, rocky formations were becoming the dominant habitat type across most observed that delimited the beginning of the drop- of the Lebanese sea bottom. off or , as was the case with coralligenous habitat. However, these formations In this mixed habitat type, bioturbation by could be found at greater depths, extending into small gobies of the genus Lesueurigobius was the upper bathyal zone. Many of these formations apparent. Two such species were observed to be were found to be covered with sponges such as present: Fries’ goby (L. friesii) and Lesueur’s goby , Phorbas fictitius, and other non- (L. sueri). In other regions of the Mediterranean identified that were white, cream, and nearby waters of the , these yellow, orange, or red in colour. fishes typically share the holes that they create with crustaceans such as Goneplax rhomboides. In the deep bathyal zone, rocky formations However, no such observation was made during were scarce, although they were occasionally these surveys. observed on muddy substrate. In most cases they were found on the steepest walls of the This transition zone between sandy and muddy canyons, where the fine sediment could not bottoms in the lower circalittoral and upper fully cover them. In these areas, some deep-sea bathyal zones was the preferred area for extensive coral species (e.g., Caryophyllia (Caryophyllia) communities of red sea pens (Pennatula rubra). calveri) had settled, together with various serpulid In some cases, these were mixed communities polychaetes, numerous scyphozoan polyps of the of red and slender sea pens (Virgularia mirabilis), genus Nausithoe, crustaceans (e.g., Bathynectes although the latter species also formed maravigna and Munidopsis marionis), and, monospecific facies. more rarely, gorgonians (i.e., Swiftia pallida) or As was the case with sandy bottoms, sponges. In one of the deepest surveyed areas anthropogenic waste in this habitat type was of the Jounieh canyon, even a glass sponge was observed to have altered some areas, both observed (i.e., Farrea bowerbanki). through establishing new biotopes and supporting In some areas of the deep bathyal zone, ancient the presence of species more typically associated reefs had formed hard substrate (Map 4). These with hard bottoms. fossil reefs were found in Jounieh and Tarablus/ Batroun canyons (and possibly in other areas surveyed, such as St. George, where hard substrates documented looked similar to these formations), and provided a substrate that was functionally equivalent to areas of rocky bottom.

The most commonly recorded fish species in rocky areas were swallowtail seaperch () (Figure 10), parrot seaperch ( ruber), and deepwater cardinalfish (Epigonus constanciae). Although their depth ranges were distinct, in some cases these species were found © OCEANA / Enrique Talledo to overlap. Map 4. Distributionofrocky bottomsandfossilreefs insurveyedareas. 21

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Figure 10. Swallowtail seaperch (Anthias anthias) and parrot seaperch (Callanthias ruber) over rocky bottom, with the Megerlia truncata and star coral ( pharensis).

Crustaceans, such as pandalid shrimp of the Bathyal muds genus Plesionika (particularly P. narval) were The bathyal zone was predominantly covered also present in rocky areas in high densities, by muds, which gave rise to different facies and in some cases mixed with P. edwardsii. Less communities. These habitats were dominated by commonly, individuals of small European locust echinoderms, large foraminifera, brachiopods, lobster (Scyllarus arctus) and golden coral shrimp tube worms, tube-dwelling anemones, and some (Stenopus spinosus) were also observed. sea pens. Canyon heads The foraminifera Pelosina cf. arborescens was In addition to the canyon heads that begin distributed throughout a wide bathymetric range immediately beyond the coralligenous formations, and was the most abundant species on these others dropped off suddenly from areas of beds. Not surprisingly, large foraminifera were soft bottom, such as in the case of the Beirut intermixed with the tubes of some sabellid worms, escarpment. The Beirut slopes were characterised which may have been Spiochaetopterus sp. by rocks covered with star coral (Madracis pharensis), deep-sea oyster (Neopycnodonte The brachiopod Gryphus vitreus was observed to cochlear) and/or the brachiopod Megerlia be distributed both in a dispersed manner, and in truncata, and various sponges that overlaid large small groups, at depths below 450 m. areas of rocks. The cerianthiarians, particularly Cerianthus Among the fishes found in these areas, leopard- membranaceus, were documented throughout spotted goby (Thorogobius ephippiatus) were the range of depths surveyed. They were found frequently observed, along with some isolated iris to be abundant on substrates as disparate as wrasse (Lappanella fasciata) individuals. coralligenous habitat and bathyal muddy bottoms.

Various echinoderms occurred on both soft and Although sea pens were more abundant in the rocky bottoms at the head of the canyons, such domain between the circalittoral and bathyal as smooth sea star (Hacelia attenuata) and long- zones, their distributions also extended to greater spine slate pen sea urchin ( cidaris). depths. Pennatula rubra continued to be the most hundred individualspersquare metre. could reach severaltens,orevenmore thanone Penilpidia ludwigi.Densitiesofthelatterspecies intestinalis andthesmallelasipodidholothurian two mostcommonspecieswere Mesothuria Holothurians were alsofrequently observed.The their remains were visibleonthesubstrate. animals, giventhattheylivewithinthesediment, lyrifera; althoughitisdifficult to observethese muds, suchasirregular sea urchins Bryssopsis urchins were also known to occur on bathyal down tothedeepbathyalzone.Othersea a verywiderangeofdepths,from thecircalittoral pen seaurchin (Cidariscidaris ) wasfoundacross echinoderms. For example, the long-spine slate organisms onthesesoftbottomswere After theforaminifera,mostdominant were foundatapproximately 300mdepth. of 500m,andremains of dead F. quadrangularis quadrangularis) were observedstartingat depths while someindividualsoftallseapen(Funiculina abundant oftheseapenspeciesintheseareas, Jounieh canyon,at593mdepth. Figure 11. CommonAtlanticgrenadier (Nezumiaaequalis)andglassheadgrenadier (Hymenocephalusitalicus)in (Madracis pharensis) by rocks covered withstarcoral The Beirutslopeswere characterised longirostris). were deep-waterrose shrimp(Parapenaeus were abundant(particularlyP. edwardsii),as muds, shrimpbelongingtothegenusPlesionika Among the crustaceans documented on bathyal linguatula), andSymphurus spp. (Lepidorhombus scaldback (Arnoglossusrueppellii),megrims and included species such as Rüppell’s Flatfishes were alsocommonlyobserved, rosefish (Helicolenusdactylopterus ). (Hoplostethus mediterraneus),andblackbelly spiderfish ( dubius), silver roughy sorcerer (Nettastomamelanurum),Mediterranean 11),blackfin Hymenocephalus italicus; Figure spp., Coelorinchus caelorhincus,Nezumia (Chlorophthalmus agassizi), (e.g., with this habitat type were shortnose The mostabundantfishspeciesassociated spp.), spottedflounder(Citharus 23

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SPECIES © OCEANA/ Enrique Talledo In total, 619 taxa were identified: 352 to the level of species, 121 to genus, and 146 to order or higher. In the Annex 1 to this report, Table A1 provides a full list of recorded species by survey area, and Figure A1 shows the number of recorded species by broad taxonomic groups.

Dozens of species identified during these surveys had never previously been recorded from Lebanon. Key species of interest among these first-ever records include the glass sponge Farrea bowerbanki, rabbitfish (Chimaera monstrosa), velvet-belly lanternshark (Etmopterus spinax), and cnidarians such as the gorgonian Swiftia pallida, sea pens (e.g., Pennatula rubra, Virgularia mirabilis, and Funiculina quadrangularis), and species of the subclass , such as Sideractis glacialis and tree coral (Dendrophyllia ramea). In addition, three species were discovered that are likely to be new to science: the sponge sp. (Figure 12)39, the sea star Luidia sp., and the stony coral cf. Anomocora sp.

Observed species also included those that Figure 12. Sponge sample (cf. Axinella sp. nov.) have been assessed as threatened or at risk of collected with the ROV. becoming threatened. Seven of the identified species have been listed under Annex II of the SPA/BD Protocol of the Barcelona Convention, while six species are Red Listed by IUCN as Seven of the identified threatened (Table A2). These species were distributed throughout the five sampling areas species have been (Map 5). In addition, three recorded species are listed under Annex III of the SPA/BD Protocol,43 listed under Annex II which requires that their exploitation be managed (i.e., Epinephelus marginatus, Scyllarus arctus, of the SPA/BD Protocol and Spongia (Spongia) officinalis) (Table A2). Nineteen of the organisms identified from these of the Barcelona surveys are considered exotic; all of them are of Indo‑Pacific origin (Table A1). Exotic species Convention. were present in all five surveyed areas (Map 6), although, in general, fewer such species were identified during the expedition than expected. One possible explanation relates to the deep-sea range of the surveyed areas, since Lessepsian species tend to be associated with shallow or pelagic waters.44,45,46,47 SPA/BD Protocol oftheBarcelona Convention. Map 5. Distributionofidentifiedspeciesthatare listedasthreatened, eitherontheIUCNRedListorAnnex II ofthe 25

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Map 6. Exotic species identified in the sampling areas. in the easternmost partoftheMediterranean” in theeasternmost of anancientthermophilousfaunathatsurvived waters, someofthisfaunacouldbe“remnants proposed forseveralsponges foundinLebanese even Lessepsianspecies.Ashasalready been Atlantic speciesthantootherMediterraneanor appeared to have more taxonomic affinity to the speciesidentifiedduringthisexpedition Beyond the known exotic species, some of marine scientists, This opinionisshared by severalMediterranean diverse group ofanimals. molluscs represented themostabundanceand the taxadocumentedfrom infaunalsamples, and serpulidsonrocky substrates).Among bathyal zone (both sabellids on soft substrates even onsandybottoms);andpolychaetesinthe stones and rubble in the circalittoral zone (or turf formedbyhydrozoans andbryozoanson pharensis), andthebrachiopodMegerliatruncata); cf. Pelosina of animals: colonial species (e.g., the foraminifera numbers were exceededbyrelatively fewtypes richness, aswellhighabundance.Their groups withthehighestrecorded species fishes andcrustaceanswere thetwobroad Of alltheidentifiedtaxafrom ROVsurveys, (Tursiops truncatus). ( surinamensis),andbottlenosedolphin (Hirundichthys rondeletii),Atlantictripletail (Onychoteuthis banksii),blackwingflyingfish vessel. Theseincludedcommonclubhooksquid water’s surface,from thedeckofsurvey other specieswere observedbychanceatthe In additiontothebenthicspeciesrecorded, Echinoderms). sea areas ofLebanon(see sectionbelowon like smoothseastar(Haceliasuperba)indeep- This factcouldexplaintheoccurrence ofspecies involved inthisproject. arborescens, star coral (Madracis 49 someofwhomhavebeen 48 . various flatfishes(order Pleuronectifores). silver roughy (Hoplostethusmediterraneus)and 13), rosefish (Helicolenusdactylopterus;Figure spiderfish (Bathypteroisdubius),blackbelly (Nettastoma melanurum),Mediterranean common speciesincludedblackfinsorcerer bioturbation. Onsoftbottomsindeepwaters, leaving theseabedwithwidespread evidence of numerous burrows excavatedinthesediment, The lattergroup formedlargecolonieswith S. hepatus) and gobies (i.e., Lesueurigobius spp.). were speciesofserranids(e.g.,Serranuscabrilla, the upperbathyalzone,mostabundantfishes sedimentary bottomsofthecircalittoral zoneand greeneye (Chlorophthalmusagassizi ). On (Hoplostethus mediterraneus) and shortnose spp.),silverroughy italicus, andNezumia Coelorinchus caelorhincus,Hymenocephalus bottoms were different speciesofgrenadiers (e.g., Among thenoteworthyspeciesfrom bathyalmud chondrichthyans. the exceptionofasmallnumberindividuals fishes belongedtotheclassActinopterygii,with pending identification.Nearlyallofthe recorded to thelevelofgenus,andvariousothersstill 84 identifiedspecies,11furthertaxa group amongthespeciessurveyed withROV, with Fishes were thebestrepresented taxonomic Fishes Beirut, 650mdepth. Figure 13. Blackbellyrosefish (Helicolenusdactylopterus), 27

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The most common fishes in rocky areas were gregarious species (e.g., Anthias anthias, Callanthias ruber) and deepwater cardinalfish (Epigonus constanciae), as well as leopard- spotted goby (Thorogobius ephippiatus). The latter species was essentially restricted to the rocky bottoms at the edge of the continental shelf and canyon heads, mainly between 100 m and 200 m depth. Other frequently recorded species included large‑eyed dentex (Dentex macrophtalmus), which demonstrated great mobility between bathyal hard and soft bottoms, as well as gurnards and longspine snipefish (Macroramphosus scolopax), which typically Figure 14. Velvet-belly lanternshark (Etmopterus occurred on soft bottoms of the upper bathyal spinax) in Beirut canyon, 953 m depth. zone. Within the , abundant fishes included those belonging to the family Stomiidae, (e.g., Chauliodus sloani and Stomias boa) and lanternfishes – especially jewel lantern fish (Lampanyctus crocodilus). Spine (e.g., Nemichthys scolopaceus) were also observed.

Many of the exotic fish species observed were found in association with coralligenous habitat, partly reflecting the shallower depth range of this habitat type. Examples include dusky spinefoot (Siganus luridus), redcoat (Sargocentron rubrum), common lionfish (Pterois miles) and (Fistularia commersonii). However, others were also found in sandy-bottom areas, such as broadbanded cardinalfish (Ostorhinchus Figure 15. Longnosed skate (Dipturus oxyrinchus) in fasciatus), white-spotted pufferfish (Torquigener St. George canyon, 425 m depth. flavimaculosus), Randall’s threadfin bream (Nemipterus randalli) and Equulites klunzingeri.

Other species with more sporadic distributions were Scorpaena spp., Capros aper, Serranus scriba, Lophius piscatorius, Pagellus bogaraveo and Electrona rissoi.

Only three species of chondrichthyans were recorded during surveys: velvet-belly lanternshark (Etmopterus spinax), longnosed skate (Dipturus oxyrinchus) and rabbitfish (Chimaera monstrosa) (Figures 14‑16). All of these species were observed in the central zone of Lebanon, in canyons close to Beirut. The closest known records of this species are from Egypt50 and Israel51.

Figure 16. Rabbitfish (Chimaera monstrosa) in St. George canyon, 893 m depth. Crustaceans (2015) The checklist published by Goren and Galil longirostris), Jounieh,440mdepth. Figure 17. Deep-waterrose shrimp(Parapeaneus P. edwardsii. In somecases,theseindividualswere mixedwith groups ofseveraltensorhundreds ofindividuals. shallower waters (approximately 100 ) wasobservedin shrimp (Plesionikanarval muddy bathyalsubstrates. Meanwhile, narwal approximately 250 m depth, on both rocky and highest abundance.Itwasdocumentedfrom with themostwidespread distributionandthe edwardsii) wasbyfarthecrustaceanspecies species. Stripedsoldiershrimp(Plesionika beingthemostabundant longirostris; Figure 17) deep spp.)and with pandalidshrimp(Plesionika Surveys revealed anabundance ofcrustaceans, new records forLevantinefauna. shallower depthrange,whileothersconstituted already knownintheLevantine seas,butata and Facciolella oxirhyncha,Lepidorhonbusboscii sphyraena,Dalophisimberbis this expedition, except for Nezumia aequalis, the speciesfoundinLebanesewatersduring found deeperthan500m)includesmanyof . whiffiagonis.Someofthesefisheswere L. ‑ water rose shrimp (Parapeaneus 51 ofdeep-seafishspecies(i.e.,those ‑ 250 m), in individual crabsbelongingto thegenusEbalia. taken during surveys, with the exception of some No crustaceanswere found in thegrab samples carrying symbioticanemones. genus abundant were hermit ,primarily of the Other speciesthatwere observedtobelocally sedimentary bottoms. be abundantwithin the watercolumn overdeep shrimp Additionally, insomeareas themesopelagic and P. edwardsiisoldier shrimp(Plesionikanarval ). bathyal rocks togetherwithdenseaggregations of lobster (Scyllarusarctus)were alsoobserved,on (Stenopus spinosus)andsmallEuropean locust In isolation,speciessuchasgoldencoralshrimp 441 m depth. Figure 18. Spinolambrusmacrochelos , Jounieh, Figure 18), althoughonslightlyshallowerbottoms. ‘orangutan’ (Spinolambrusmacrochelos; was observed in the case of the pattern on sedimentarybottoms.Asimilardistributional at different depths,especiallybelow400mand Neomaja goltziana (or its remains) were found Various liveindividualsofthespider crab than theothershrimpspeciesmentionedabove. than 500m,althoughtheywere lessabundant (Aristeus antennatus)occurred inareas deeper abundant species.Inaddition,blueandred shrimp to the genus (Parapeaneus longirostris)andshrimpbelonging on muddybottoms,deep-waterrose shrimp and thecrabBathynectesmaravigna.Incontrast, the smallgalatheidcrabMunidopsis On rocky bottoms,themostcommonspecieswere Pagurus. These animals were frequently Eusergestes arcticuswasobservedto Plesionika were by far the most cf. marionis 29

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Echinoderms The dominant faunal species on bathyal found on both soft and hard bottoms, particularly muddy bottoms were echinoderms, particularly at the edge of the continental shelf. Of these holothurians such as Mesothuria intestinalis and two species, the former was more commonly Penilpidia ludwigi, which were both abundant in observed. Some individuals of Ceramaster that habitat type. Neither of these two species had granularis and Peltaster placenta were also found previously been recorded from Lebanon. Another in these areas, although in smaller numbers. echinoderm, the long-spine slate pen sea urchin (), was also widely distributed. Of particular interest were the occurrences of three species of sea stars that have not previously The small elasipodid Penilpidia ludwigi was been recorded from the Mediterranean Sea. previously known from the and the Surveys revealed Hacelia superba on bathyal Western Mediterranean52, and here was observed rocky bottoms, Leptasterias sp. on ancient fossil forming aggregations with densities of tens to reefs, also in the bathyal, and a species of the hundreds of individuals per square metre on the genus Luidia (Figure 20), which was abundant on sea bottom in deep bathyal waters. These high sandy bottoms in the circalittoral zone. The latter concentrations of individuals were typically found species was distinct from the two other species at depths greater than 800 m. of the genus that are known to occur in the Mediterranean (L. ciliaris and L. sarsi). Other significant aggregations of echinoderms occurred in the circalittoral zone. The main such aggregations were formed by: the brittle star Ophiopsila aranea on detritic bottoms; hatpin urchin (Centrostephanus longispinus) on coralligenous formations; and Mediterranean feather star (Antedon mediterranea; Figure 19) in elevated locations – such as on top of rocks or marine litter found on sandy bottoms of the circalittoral zone.

Figure 20. Luidia sp., St. George, 71 m depth.

Other species of echinoderms were observed sporadically. For example, the sea star Hymenodiscus coronata and urchin (Gracilechinus acutus) occurred on bathyal muds, while blue spiny (Coscinasterias tenuispina) and common brittlestar (Ophiothrix Figure 19. Mediterranean feather star (Antedon fragilis) were documented from circalittoral detritic mediterranea) on a coralligenous concretion, Beirut, bottoms with rubble. This expedition marked the 64 m depth. first time that H. coronata was recorded from Lebanese waters. Sea stars did not usually form large aggregations, Grab samples also contained various individuals but they were nevertheless frequently observed of the small green urchin (Echinocyamus pusillus), on certain substrates, especially in the circalittoral which were collected from sedimentary bottoms, and upper bathyal zones. Smooth sea star and the irregular sea urchin Bryssopsis lyrifera. (Hacelia attenuata) and Mediterranean red sea star (Echinaster (Echinaster) sepositus) were , These included some species of the genera hydrozoans, whichwere occasionallyabundant. abundance ofcnidarians,withtheexception Lebanese watersdonothostagreat diversityor Cnidarians Figure 21. Swiftiapallida,Jounieh,607mdepth. that belonged to two species of dead man’s to softoctocorals,individualswere recorded respect 21).With within Jouniehcanyon(Figure in smallgroups ofindividuals,onbathyalrocks gorgonian speciesSwiftiapallidawasobserved had notbeenrecorded from Lebanesewaters.The anthozoans, priortothesesurveys,gorgonians respect to With all of the sampled areas (Map 7). Lebanese waters,withseveralgenerapresent in Various octocoralswere documentedin rocky bottoms. bottoms, and some scyphozoans on bathyal or canyonheads,someseapensonsedimentary as certain scleractinians on the continental shelf some exceptionalcasesare outlinedbelow, such on softbottomsorrubble.Asforothercnidarians, Nemertesia antennina andLytocarpia myriophillum hard substrates),aswell asspeciessuch Sertularella and Eudendrium(on with which occurred ondetritic bottomstogether depths, andgrey seapen (Pteroeidesgriseum), was more sparsely distributed and at greater tall seapen(Funiculinaquadrangularis),which in someplaces.Alsorecorded inthesewaterswas abundant thanP. rubra,butwaslocallyabundant (Virgularia mirabilis),whichwasgenerallyless sometimes seenalongsideslenderseapen it usuallyoccurred asa single species,itwas circalittoral and upper bathyal zones. Although facies on sedimentary bottoms of the lower is thered seapen(Pennatularubra ), withlarge Another speciesofoctocoralthatwasabundant occur inthecircalittoral zone. spinulosum. Bothspecieswere observed to fingers: 40% duringthelasttwodecades. estimated to have decreased by approximately quadrangularis populations ofF. Vulnerable intheMediterraneanbyIUCN,with Thesefourspeciesare considered as in Map 8. . rubra.Theirdistributionsare presented P. palmatumandParalcyonium 53 and . rubra P. 31

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Map 7. Distribution of octocorals across the sampled areas. Map 8. Distributionofseapensacross surveyedareas, byspecies. 33

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Surveys also revealed at least six different species of scleractinian corals: star coral (Madracis pharensis), Caryophyllia (Caryophyllia) calveri, Devonshire cup coral (C. smithii), tree coral (Dendrophyllia ramea), cockscomb cup coral (Desmophyllum dianthus), and a fifth species that may belong to the genus Anomocora. This species had not yet been described from the Mediterranean, and may potentially be a new species. All occurrences of scleractinians are represented in Map 9.

Star coral (Madracis pharensis) was the most abundant hard coral species. It formed dense Figure 22. Dendrophyllia ramea on soft sediments in aggregations on rocky bottoms of the deep Tarablus/Batroun, 172.4 m depth. circalittoral and upper bathyal zones, frequently at the edges of the continental shelf and canyon Other hexacorals present in the areas surveyed heads. In addition, it was found in small colonies, included anemones, such as cloak anemone on scattered rocks on the continental shelf, and ( palliata), although they were not on anthropogenic waste. abundant. Hormathia alba was also recorded and The next most common hard coral species is was always associated with the shells that were still pending identification. Some individuals were carried by hermit crabs of the genus Pagurus. It found detached, on sandy bottoms, as well as is also worth highlighting the occurrence of some attached to small rocks, and objects of artificial individuals of the corallimorpharian Sideractis origin, such as litter, oil drums, tyres, cans, and glacialis. metallic objects. It did not form large colonies Ceriantharians were distributed across a wide and in many cases appeared as solitary corals range of depths and habitat types, with some scattered on the seabed. This species was only areas of high abundance. Most of these animals observed to occur on the deep continental shelf, were found on bathyal muddy bottoms, but they typically below 40‑50 m depth. were also common in coralligenous beds of Corals belonging to the genus Caryophyllia were the continental shelf edge. The most common always found on hard substrates. Devonshire cup species was the cylinder anemone (Cerianthus coral (C. smithii) was seen in shallower zones, and membranaceus; Figure 23); some lesser cylinder even in deep circalittoral waters, while C. calveri anemone (C. lloydii) was also found, as well as could be abundant on bathyal rocky bottoms a few Arachnanthus sp. and Pachycerianthus sp. at different depths. It is also possible that other species of this genus may be present in Lebanese waters, such as C. inornata and other deep-sea species. However, sampling would be necessary in order to verify their identification.

In the case of Dendrophyllia ramea, it was only observed in one location in the Tarablus/Batroun area, at 172.4 m (Figure 22), a record depth for this species. Small colonies were found on sedimentary substrate, similar to those which were recently discovered in the waters of .54

Figure 23. Cylinder anemone (), Beirut, 648 m depth. Map 9. Distributionofscleractiniancoralsinthesamplingareas. 35

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Finally, were not observed to In some cases, other species were locally be abundant, except in the case of small abundant, such as aaptos, massive scyphozoans in the polyp phase, on vertical walls beige horny sponge (Biemna variantia), Crambe of the bathyal zone (Figure 24). These animals crambe, and various Axinella spp. One such belonged to the genus Nausithoe, and colonised Axinella species, found and collected in the the small overhangs and protrusions of deep-sea Sayniq area at 525 m depth, was of particular rocks and ancient fossil reefs. interest as it appears to be a newly discovered species. Its morphology is similar in some aspects to the genus Axinella, but differs in other specific characteristics.39

A variety of other sponges occurred on smaller rocks, rubble, and sandy bottoms. These species included Chodrosia reniformis, Corticium candelabrum, Dysidea fragilis, Haliclona spp., Oscarella lobularis, Petrosia (Petrosia) ficiformis, and Ulosa stuposa. Meanwhile, a diversity of Figure 24. Scyphozoans, vermetids and corals on other sponges covered deep rocky surfaces. deep-sea rocks, St. George canyon, 683 m depth. Among these species were Hexadella dedritifera, Phorbas fictitius, simple‑rayed membrane sponge Sponges (Plakina monolopha), Pleraplysilla spinifera, and Large sponges were common on the continental many other unidentified species. shelf and, in particular, on the rocks and Another species of interest, the glass sponge coralligenous formations at the shelf edge; in the Farrea bowerbanki (Figure 26), was observed on latter areas, Agelas oroides was the dominant vertical walls formed by fossil reefs in deep waters species (Figure 25). It was generally found of the Jounieh canyon.39 together with other characteristic species, such as various sponges belonging to the family Ircinidae. In particular, these species included Sarcotragus spinosulus and Ircinia variabilis, as well as Fasciospongia cavernosa and Chlathrina clathrus. These sponges were the largest and most representative habitat-forming species and were present in all the sampling areas (Map 10).

At one location in the Sayniq canyon, Calyx nicaeensis was also present. This cup‑shaped has a fragmented distribution across Figure 26. Hexactinellid sponge (Farrea bowerbanki) the Mediterranean Sea, and concerns have been on fossil reef in Jounieh. canyon, 857 m depth. raised about the need for urgent conservation measures to protect it.55,56,57 It was possible to confirm the presence of additional sponges in surveyed areas, via samples that were collected from sediment grabs or with the hydraulic arm of the ROV. Species that were sampled in this way included Hymedesmia sp., Plakortis simplex, Prosuberites longispinus, Timea geministellata, T. stellata, and Topsentia calabrisellae.57

Figure 25. Agelas oroides, Clathrina clathrus, Dysidea fragilis and other sponges on rocky bottom, Jounieh, 77 m depth. Map 10. Distributionofselectedspongespeciesinthesamplingareas. 37

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Molluscs There is still a lack of detailed knowledge regarding molluscs in the Eastern Mediterranean. Some groups are well represented, such as the fauna, which comprises 81% of the known cephalopod species recorded for the entire Mediterranean.58 However, there are no comprehensive lists for other mollusc groups in the Eastern Mediterranean or, specifically, in the Levantine Sea. Some previous studies carried out in Lebanon11,59,60 have provided valuable information about molluscs, but the results from this expedition underscore the knowledge gap that still exists regarding in this regard, both in shallow and deep waters.

Via the ROV videos recorded during the expedition, it was possible to document the occurrence of a diversity of , bivalves, gastropods, and scaphopods. Grab samples also provided Figures 27 and 28. Up: Cuttlefish effs in Jounieh additional data on bivalves, gastropods, and canyon, 522 m depth. Below: Sepiidae eggs in a tube, scaphopods which, due to their size or behaviour Beirut, 818 m depth. (e.g., living within the sediment) would have been impossible to identify via direct visual methods. Other cephalopods which occurred on bathyal Among the cephalopods observed, it is worth muddy bottoms were gastropods such as highlighting the occurrence of species such as sp., Naticarius stercusmuscarum, stout bobtail (Rossia macrosoma), fourhorn Semicassus granulata, and an unidentified octopus (Pteroctopus tetracirrhus), common species belonging to the family . A large octopus (Octopus vulgaris), cuttlefish (Sepia giant tun individual (Tonna galea; Figure 29) was officinalis), pink cuttlefish (S. orbignyana), neon also observed in the Jounieh area. This species is flying squid (Ommastrephes bartramii) and greater listed as threatened, on Annex II of the SPA/BD (Argonauta ). In the case of the Protocol of the Barcelona Convention.41 latter species, only remains were observed on the seabed.

Octopus vulgaris and Sepia officinalis were found in sedimentary areas of the continental shelf, using anthropogenic waste (such as old tyres and plastic) to hide themselves. The other documented cephalopod species were observed on bathyal muddy bottoms, although some of the most abundant squid that could not be identified were found in the water column. Figure 29. Tonna galea, Jounieh, 239 m depth. In some of the deepest areas surveyed, clusters of cuttlefish eggs were seen on both rocky bottoms One noteworthy finding was the presence of large and on pieces of marine litter (Figures 27, 28). quantities of deep‑sea oysters (Neopycnodonte cochlear) at the continental shelf edge, in the canyon heads of Jounieh, St. George, Beirut and Sayniq (Map 11), and on the continental shelf (Figure 30). In some cases, they were found to be covering marine litter, such as tyres. Map 11. Oysteraggregations (Neopycnodontecochlear)insurveyedareas. 39

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pallasii, Caberea ellisii, Hornera frondiculata, and Reteporella sp. Other identified species were Dentiporella sardonica, Exidmonea atlantica, Frondipora verrucosa, Margaretta cereoides, Rhynchozoon neapolitanum, and Scrupocellaria cf. scrupea. All of these species were found on the continental shelf, both in coralligenous habitat and sedimentary bottoms.

Harmelin et al.61 described 93 species of Figure 30. Deep‑sea oyster (Neopycnodonte bryozoans in Lebanese waters, including some cochlear) in Jounieh, 70 m depth exotic ones; all of those species were collected at depths shallower than 42 m. Because this In some areas of the continental shelf, individuals expedition focused on sampling in waters below of rough penshell (Pinna rudis), a species included that depth, new records were expected to be within Annex II of the SPA/BD Protocol of the found. Only 13 species of the bryozoans found Barcelona Convention, and muricids (e.g., Murex have been identified to the species level so far, forskoehlii and ) were observed. and seven of these constitute new records. The At approximately 60 m depth, in continental species Hornera frondiculata was of particular shelf areas, some vermetid aggregations or reefs interest since, despite its abundance, it had not were found which were covered by sponges, previously been mentioned from either Lebanon or polychaetes, and hydrozoans (Figure 31). These in a recent Israeli compilation.62 Further samples communities were on sedimentary bottoms where are still pending analysis and revision and may add some scleractinian corals were also common. to the number of known Lebanese bryozoan taxa. Brachiopods The most common brachiopod species on hard bottoms was Megerlia truncata (Figure 32). It could reach high densities on the edges of the continental shelf and in canyon heads, in some cases forming colonies together with various polychaetes or corals (e.g., Madracis pharensis). Specimens obtained from grab samples also revealed the presence of brachiopods, such as Figure 31. Vermetid reef in Jounieh, 60 m depth. cuneata and Megathiris detruncata.

Additional data about molluscan fauna were obtained via grab samples, which revealed considerable species richness within the seabed of the continental shelf. Dozens of mollusc species were identified from these areas, as well as from bathyal muds. Among these species were bivalves such as Anadara sp., European clam (Corbula gibba), Nucula spp., Parvicardium sp., Similipecten similis, and oval venus (Timoclea ovata), and gastropods such as inaequicostata, spp., Figure 32. Megerlia truncata on the border of pyramidata, necklace shell (Euspira catena), Taranis St. George canyon, 173 m depth. moerchii, and Turritella turboma. Many of the molluscan species identified from the grab sampling On bathyal bottoms, below 450 m depth, had not previously been recorded in Lebanese Gryphus vitreus was also documented. waters, even from the coastal shelf. Although densities were not observed that were comparable to those discovered in other parts of Bryozoans the Mediterranean,63,64,65,66 they were nevertheless locally important in some areas. Through the ROV videos and grab samples, it was possible to detect the presence of some Both M. truncata and G. vitreus were distributed in large bryozoan species, such as Adeonella all sampled areas (Map 12). Gryphus vitreus)insampledareas.Map 12. Distributionofbrachiopodspecies(MegerliatruncataandGryphus 41

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Tunicates been described as inhabiting fossil corals in the Eastern Mediterranean.67 All of the observed tunicates were found on the continental shelf or in canyon heads, on hard Also of interest was the wide distribution of the substrates. In some cases, they were observed echiuroid Bonnelia viridis, in terms of both its on dead corals or bryozoans or on anthropogenic geographic and bathymetric ranges. It was found waste. This was the case of those belonging to to be abundant in the lower circalittoral and upper the families Didemnidae (Figure 33) or Pyuridae, bathyal zones, but also was observed in high while Halocynthia papillosa had a wider range of densities in some bathyal areas. It even occurred distribution and preferred natural substrates. at depths of nearly 1000 m. Foraminifera Particular mention should be made of the foraminifera, as it is an important taxonomic group in Lebanese waters, due to its high abundance. Large astrorhizid foraminifera, such as Pelosina cf. arborescens (Figure 34), occupied extensive areas of all bathyal muddy bottoms surveyed (Map 13).

In areas of the continental shelf, Miniacina Figure 33. Didemnidae growing on dead Sarcotragus miniacea was also documented, while other spinosulus, Tarablus/Batroun, 102 m depth. species (e.g., Ammodiscus cretacicus, Quinqueloculina sp., and Pyrgo inornata) were Annelids recorded from grab samples. In some places, large concentrations of sabellid Algae or serpulid annelids were found. In the case of sabellids, these aggregations occurred on Various species of algae were found in all of the sedimentary bottoms and were often mixed with surveyed areas. Due to the fact that most of large foraminifera. Meanwhile, serpulids were the areas surveyed were below 50 m depth, most common on both hard bottoms of the continental of these algae are sciaphilic, such as calcareous shelf and upper circalittoral, and in escarpments, red algae. However, some green algal species, overhangs, and rocks in canyon heads and in the like Valonia sp. and Caulerpa scalpelliformis were bathyal zone. also documented.

The majority of annelids observed could not be identified, with the exception of a small number of species, such as lacy tubeworm (Filograna implexa), bearded fireworm (Hermodice carunculata), sand mason worm (Lanice conchilega), red-spotted horseshoe (Protula tubularia), Mediterranean fanworm (Sabella spallanzanii), peacock worm (Sabella pavonina), and red tube worm (Serpula vermicularis).

Although it was not possible to take samples of some of the deep-sea polychaetes to verify their , one of the most common polychaetes observed to have settled on deep-sea rocks and fossil reefs appeared to be the serpulid Vermiliopsis monodiscus. This tentative identification seems Figure 34. Giant foraminifera (Pelosina cf. arborescens) probable, given that this species has previously with slender rockfish (Scorpaena elongata). Map 13. ObserveddistributionoftheforaminiferaPelosina cf. arborescens. 43

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TARABLUS/BATROUN

The area between Tarablus and Batroun was characterised by many different types of substrates, including rhodolith/maërl beds, coralligenous concretions (living and dead), hard rocky bottoms including fossil reefs, detritic sand, and mud beds.

This survey area was the only place where the scleractinian coral Dendrophyllia ramea was observed, on muddy bottoms at 172.4 m depth, the deepest location known for the species.

Areas Deep‑sea populations of D. ramea such as these have only been discovered very recently in the Mediterranean; they have been documented from another Levantine area, on muddy bottoms in Cyprus.54 From a conservation perspective, the presence of D. ramea in Lebanese waters is of particular interest. It is considered Vulnerable on the IUCN Red List of Mediterranean Anthozoans, on the basis of population declines in the Mediterranean,53 and was recently granted strict protection under Annex II of the SPA/ BD Protocol of the Barcelona Convention. Lebanon, like other signatory countries, is therefore required to take measures to safeguard this species in its waters.

Tarablus/Batroun was also the area with the highest abundance of typical Mediterranean habitat-forming species that are considered vulnerable or sensitive. For example, tall sea pen (Funiculina quadrangularis) and the brachiopod Gryphus vitreus were present in higher numbers in that area than in the other four. Other sea pens (i.e., Pennatula rubra and Pteroeides griseum) were also found there, in association with sandy bottoms.

Other species of special interest were recorded from the Tarablus/Batroun area. Small European locust lobster (Scyllarus arctus), which is listed in Annex III of the SPA/BD Protocol of the Barcelona Convention, was found on rocky bottoms at 287 m depth, together with other crustaceans (e.g., Stenopus spinosus, Plesionika edwardsii and P. narval). Large bryozoans (e.g., Hornera frondiculata or Caberea spp.) were commonly found in rhodolith/maërl beds, detritic sand and coralligenous beds, while a population of the Annex II‑listed large mollusc rough pen shell (Pinna rudis) was found in detritic sediments. JOUNIEH

The Jounieh area supported coralligenous habitat, rhodolith/ maërl beds and sandy bottom with high biodiversity. Jounieh also hosted other interesting deep-sea substrata, such as rocks, fossil reefs and muddy bottoms.

Overall, Jounieh comprised the highest number of total species identified during the expedition. This canyon was found to host some interesting and rare species, like the Atlantic Hacelia superba and Leptasterias sp., the protected mollusc species Tonna galea, the recently discovered glass sponge Farrea bowerbanki and the octocoral Swiftia pallida. In the case of the latter tyres. deep objects were colonisedbyfauna,especially inside thecanyons.Manyofanthropogenic Marine litterwasabundantbothontheshelfand were documented. parts oftheserocks, structures likefossilreefs and solitarystonecoralswere hanging.Onsome were visible,from whichscyphozoans,serpulids, However, insomesteepzones,emergentrocks most ofthecanyonwascovered bymuds. by brachiopods and sponges.In deep waters, where somerocks appeared thatwere covered bottom extendedtotheheadofcanyon, coralligenous andrhodolith/maërlbeds.Detritic In shallowwaters,St. George canyonhostedboth ST. GEORGE found livingondeep-searocks orfossilreefs. Both the glass sponges and the gorgonians were gorgonian hadbeenfoundinLebanesewaters. species, itsdiscoverymarkedthefirsttimethata ‑ sea oysters (Neopycnodontecochlear)on sabellid wormsandlargebioturbations. which insomecaseswere mixedwithunidentified cf. by giantforaminifera(Pelosina Areas ofmuddybottomwere heavilycovered be NearThreatened byIUCN. canyon. Bothofthesespeciesare considered to oxyrinchus), were recorded from theSt. George expedition (Chimaera monstrosaandDipturus three chondricthyansfound duringtheresearch aggregation onlyfoundin this location.Two ofthe in areas close to 700 m depth, an exceptional of unidentifiedtopshells(Trochidae) were filmed canyon,alargenumber dives intheSt. George of interest. Forexample,during oneoftheROV Surveys inthearea alsorevealed specificfauna the area, especiallyamong coralligenousbeds. spinefoot (Siganusluridus)were alsocommonin such as redcoat (Sargocentron rubrum) and dusky shrimp (Plesionika edwardsii ). Exotic fish species, (Parapeneus longirostris)andstripedsoldier St. George area, suchasdeep-waterrose shrimp Shrimp were amongthecommon speciesinthe © OCEANA/EnriqueTalledo 68 arborescens), 45

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BEIRUT SAYNIQ

The head of the Beirut canyon had a steep The Sayniq canyon has a very well-represented gradient and rocky bottom with many corals, deep-sea muddy bottom community, with many brachiopods, oysters and sponges covering the species that are characteristic of this kind of hard bed. Surveys of the area were complicated substrate. by both the rocky bottom and the fact that the walls of the canyon head were densely covered While the deep‑sea elasipodid Penilpida ludwigi by fishing lines and nets and other garbage. was common around all of the Lebanese canyons surveyed, it was most abundant in The most common fishes in this canyon were Sayniq, together with other holothurians (e.g., swallowtail seaperch (Anthias anthias), parrot Mesothuria intestinalis) and the giant foraminifera seaperch (Callanthias callanthias), iris wrasse Pelosina cf. arboresecens. Striped soldier shrimp (Lappanella fasciata) and leopard-spotted goby (Plesionika edwardsii) was also abundant and was (Thorogobbius ephippiatus). widely distributed in the canyon.

Coralligenous habitat in the Beirut area was Coralligenous habitat was also important in the occupied by many large sponges and was the head of the canyon and along the continental site of the largest concentration of hatpin sea shelf break. The benthic community associated urchin (Centrostephanus longispinus) recorded with coralligenous formations in these areas was in Lebanon. This species is protected in the characterised by large sponges (e.g., Agelas Mediterranean, as it is listed in Annex II of the oroides, Fasciospongia cavernosa, and Spongia SPA/BD Protocol of the Barcelona Convention. (Spongia) officinalis). In addition, the rare sponge species Calyx nicaeensis was also found in The only recorded during the research this area, and the dusky (Epinephelus cruise, the velvet-belly lanternshark (Etmopterus marginatus), which is listed in Annex III of the spinax), was found at nearly 1000 m depth. SPA/BD Protocol of Barcelona Convention. This small species of lantern shark has not been previously recorded from Lebanese waters. The Mediterranean fanworm (Sabella spallanzanii), a worm species that has been decreasing in The largest sea pen fields documented during Lebanese waters, appeared to still be abundant this survey were also recorded in the head of in the Sayniq canyon. the Beirut canyon. The threatened red sea pen (Pennatula rubra) was the most abundant species, In the sandy bottom close to the coralligenous although slender sea pen (Virgularia mirabilis) was beds, two introduced species were found: the also present. The latter species was abundant in algae Caulerpa scalpelliformis, and reticulated specific sites, and also in mixed aggregations with leatherjacket (Stephanolepis diaspros). P. rubra. C. scalpelliformis is a species that has been known from Levantine waters since 192969 and The potentially newly discovered sea star species has been recorded in Lebanon at depths of up belong to the genus Luidia was also very abundant to 44 m.70 In the case of the surveys presented around the Beirut canyons, specifically on sandy/ here, the species was recorded between 66 m detritic bottoms. and 82 m depth. Similarly, S. diaspros was also first recorded in Palestinian waters in the 1920s.71 Since then, it has expanded along the shores of The potentially newly the Eastern Mediterranean, even reaching the discovered sea star Central Mediterranean.72 species belong to the genus Luidia was also very abundant around the Beirut canyons. Human Impacts (St. George, 70mdepth). Figure 36. Tyres andglassbottlesondetriticbottoms (see below). metallic waste such as appliances,and fishing nets (Figure 36), encountered in sites that appeared to be disposal areas for tyres Meanwhile, inmanypartsofthecontinentalshelf,marinelitterwas Figure 35. Barrel inJouniehcanyon,596m depth. urban waste,andoildrums(Figure 35). these deepareas, itwasusual to findamultitudeofplasticdebris, depths, formingtrueunderwaterlandfillsitesinsomezones.In Household wastewasdiscovered inallofthecanyons,andat into theseawere observed inalloftheareas surveyed(Map 14). quantities ofgarbageandotherdebristhathadbeenthrown widespread presence ofmarinelitterontheseafloor. Abundant The most visible human impact on the Lebanese seabed was the MARINE LITTER 47

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Map 14. Distribution and abundance of marine litter in surveyed area. habitats andecosystems. some ofthemostproductive –butsensitive – indicates aworryingconcentrationofeffort on Thisfinding edge, andcanyonheads(Figure 38). found inareas ofcoralligenoushabitat,theshelf The highestdensity of lostfishinggearwas traps (Figure 37). discarded fishinggear, includinglines,nets,and impacts were widespread, intheformoflostor were observedontheseabed. However, indirect a result, nodirect impactsofsuchfishingactivities trawling) donotoccurinLebanesewaters,andas damaging to benthic ecosystems (i.e., bottom fisheries. Thefishingtechniquesthat are most surveyed wasrelated tosmall-scale orartisanal All ofthefishingactivityobservedinareas FISHERIES Batroun canyon. Figure 38. FishinglinesattheheadofTarablus/ 307 m depth. Figure 37. LostfishingtrapinSt. Georgecanyon, Jounieh canyon(Figure 40). an amphorawasobservedontheseabedof on only one occasion during the surveys, when Marine archaeological remains were encountered OTHER OBSERVATIONS fishing gear. of lostordiscarded widespread, intheform impacts were Indirect fisheries Figure 40. AmphorafoundinJounieh,88 mdepth. Batroun, 267mdepth. Figure 39. Lostfishingnetinthearea of Tarablus/ the associatedriskofentanglingROV. seabed, butalsohindered surveys,becauseof such netsnotonlycovered extensiveareas of 39).Forexample,around Beirut, heads (Figure formed dense covers over some rocky canyon in coralligenousareas, while abandonednets Remains offishinglineswere mostnumerous 49

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The importance of the biodiversity of the deep-sea canyons off the Lebanese coast is well recognised, with various recommendations for their conservation, including from Oceana’s MedNet proposal36, Lebanon’s Marine Protected Area Strategy,35 and the CBD process to identify ecologically or biologically significant marine areas (EBSAs) in the Mediterranean Sea.33 The importance of deep-sea canyons has also been emphasised under the UNEP-MAP RAC/SPA ‘Dark Habitats Action Plan’73 for the conservation of these and other sensitive Mediterranean ecosystems that require protection.

Designating MPAs to protect such deep-sea areas would represent an important step towards strengthening the marine protected area network within Lebanon, and with the Mediterranean Sea as a whole, by extending coverage to key offshore areas that harbour unique biodiversity and serve as important connections between coastal and deep-sea ecosystems. Proper management is also urgently needed, particularly in the case of those species protected under the Barcelona Convention, and species considered by IUCN as threatened that have been found to inhabit these waters.

Advancing the protection of these canyons, however, depends on having high-quality, scientific data about the that they support. Until now, such data have been lacking. Prior habitat mapping studies of the canyons had relied mainly on technologies such as multibeam echo sounders, but biological surveys had only been carried out within the first few hundred metres of depth. The 2016 Deep-Sea Lebanon Expedition sought to close this knowledge gap. The research expedition represented the first time that an ROV was used to document benthic habitats and communities at greater depths.

Rhodolith bed Discussion and conclusions respect tohabitattypesinclude: The highlightsoftheexpeditionfindingswith and variousassociatedcommunites. taxa withinsixbroad habitattypes 619 identified of marinelifeinthefiveareas surveyed,with The four-week expeditionrevealed adiversity MAIN EXPEDITIONFINDINGS forming species. forming and vulnerable habitat- coralligenous concretions Key findingsinclude – – – – documented. gorgonians (Swiftiapallida)were also sponges (Farreabowerbankii)and spp.),glass dianthus andCaryophyllia stony corals(e.g.Desmophyllum species ofinterest were found,including organisms. Intheseplaces,particular reefs builtbycorals,wormsandother bottoms were related toancientfossil while indeep-seaareas, somerocky sponges, oysters,andbrachiopods, covered withorganismssuchascorals, surrounding canyon heads, rocks were important benthiccommunities.Inareas Barcelona Convention. their conservation,assignatoriesofthe Mediterranean statesare committedto essential tohundreds ofspecies,andall beds are considered asvulnerable and Both coralligenoushabitatandrhodolith productivity, whichattractsfishers. to thecoast,anditsrelatively high gears, probably duebothtoitsproximity affected bylostordiscarded fishing species. Italsoappeared to bethemost supported a high diversity of in turn formed well-developedreefs, which also documented. The coralligenous locations where rhodolith bedswere the headsofcanyonssurveyed, coast, onthecontinentalshelfandat habitat extending along the Lebanese Rocky bottom Extensive ar eas ofcoralligenous areas supporting species, theexpeditionrevealed: Among theobservationsregarding individual – – – – – – – – (e.g., been recorded from theMediterranean additional speciesofscientificinterest. pending identificationandmayyet reveal obtained duringtheexpeditionare still genus identification, butlikelybelongingtothe and astonycoralwhichispending two knownMediterraneancongeners), (which appearstobedifferent from its a sea star belonging to the genus a spongebelongingtothegenusAxinella, Thr should beputinplace. protection andmanagement measures in theMediterraneanbyIUCN,andso species are assessedasthreatened bathyal muddyareas. Most ofthese quadrangularis) occurred incertain with mud,whiletallseapen(Funiculina areas where sandybottomwasmixed mirabilis) were widely distributed in Multiple speciesthathavenotpr assessed byIUCNasEndangered in and Scyllarusarctus);twospecies marginatus, Spongia(Spongia)officinalis Barcelona Convention(Epinephelus oftheSPA/BD Protocol ofthe Annex III whose exploitationisregulated under Sarcotragus foetidus);three species Tonna galea, Desmophyllum dianthus,Pinnarudis longispinus, Dendrophylliaramea Barcelona Convention(Centrostephanus oftheSPA/BD Protocol ofthe Annex II measures ofprotection are required under including: seven species for which strict threatened intheMediterraneanSea, Swiftia pallida,Dendrophylliaramea). bowerbanki, previous Lebaneserecords (e.g.,Farrea species forwhichthere were no and dozensofknownMediterranean (i.e., forming species.Forexample,seapens that supportedvulnerablehabitat- Muddy andsandy-muddybottoms Thirteen speciesthatar ee speciesthatmaybenewtoscience: Hacelia superba, Pennatula rubra Anomocora. Additionalspecimens Tursiops truncatusand Chimaera monstrosa, Leptasterias and e considered Virgularia eviously Luidia sp.), 51

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the Mediterranean Sea (Epinephelus KEY COMMUNITY TYPES AND THEIR marginatus and Desmophyllum ECOLOGICAL IMPORTANCE dianthus), five assessed as Vulnerable (Dendrophyllia ramea, Funiculina Discussed below are the most important quadrangularis, Merluccius merluccius, community types found during the expedition, Tursiops truncatus, and Pennatula from a conservation perspective. We therefore rubra). Two additional species were focus on those communities that: are considered observed that are considered Near threatened or vulnerable; are protected under Threatened (Chimaera monstrosa and international agreements; or play significant Dipturus oxyrhincus). ecosystem roles, either by building habitat or providing other ecosystem services. Coralligenous concretions and rhodolith/ maërl beds. Rhodolith/maërl beds occur across the entire Mediterranean basin, from 25 m to 150 m depth, with some exceptional observations from deeper areas in the .74,75 Coralligenous concretions are also present throughout the Mediterranean. They are found within a depth range of 12 m to 120 m76 and, like rhodoliths/ maërl beds, also reach deeper depths in the Balearic Sea2, due to the transparency of those waters. Both habitat types are considered hotspots of biodiversity; hundreds of marine species depend on their three‑dimensional structures, for refuge, hard substrate upon which to settle, and food and nursery grounds.1,3,77 Together with Posidonia oceanica meadows, these calcareous concretions are among the most biodiverse habitats of the Mediterranean Sea, and also represent key habitat for threatened and commercial species. During the Lebanese expedition, these communities were documented in all of the survey areas, in shallower waters, and were associated with a diversity of other marine life.

Coralligenous concretions and rhodoliths/maërl beds are vulnerable to anthropogenic impacts, such as damage from trawling and other fishing activities, pollution, sedimentation, warming and invasive species.4,78 Both habitat types are listed as habitats of interest under the Barcelona Convention, and so sites with presence of these habitats are to be considered for the creation of marine protected areas (MPAs). Also, under the Barcelona Convention, the Action Plan for the Conservation of the coralligenous and other calcareous bio-concretions in the Mediterranean Sea was adopted in 2008, and then updated Filograna implexa in 2017.79 Under the Action Plan, signatories of law inEUMediterraneanwaterssince2006. vulnerable communitieshasbeenprohibited by dredges, shore seines and similar nets over these Mediterranean. Therefore, fishingwithtrawlnets, is oneofthemainthreats to thesehabitatsinthe coralligenous habitats and rhodolith/maërl beds Union laws.Forexample,trawlingabove these importanthabitatsare foundinEuropean best examplesoflegislativemeasures toprotect As highlightedwithintheActionPlan,someof habitats. management planstoavoidanydamagethese the creation ofMPAs) andscience-based monitoring), andtoestablishlegislation(including of thesehabitats(through datacollectionand the convention are urged to improve knowledge and forhuman society. marine environment, for thehealthof that are essential ecosystem services a widevariety of species thatprovide Sponges are key Barcelona Convention. maërl should be made, usingthe tools of the strengthen legalprotection ofcoralligenousand EU, theActionPlannotesthatsimilarefforts to and LIFE-IP INTEMARES projects). Outside of the the frameworkofSpanishLIFE+INDEMARES been underdebateinseveralscientificfora(e.g., I has beds as protected habitats under Annex meadows. ListingMediterraneanrhodolith/maërl degree oflegalprotection asPosidoniaoceanica coralligenous habitatandmaërlbedsthesame fisheries, theActionPlanpointstoneedgrant In additiontoaddressing thedirect threat from of theseimportanthabitatsattheregional scale. Mediterranean basin,toenhancetheprotection This prohibition shouldbe appliedtotheentire 80 the generaIrcinia, compounds havebeendiscovered inspongesof pharmacological effects. with antimicrobial, for .Theyproduce compounds sponges themselvesare importantresources In additiontotheservicestheyprovide insitu, species andtheirprey. many different species,includingcommercial density aggregations thatserveashabitatfor deep-sea sedimenterosion; andforminghigh- building andconsolidatingsubstrata;reducing for marinenutrientsandorganicproduction); nitrogen andsiliconcycles(whichare important communities; playingadecisiverole inthe linking nutrientsfrom thewatercolumntobenthic for humansociety. Thesecontributionsinclude: for thehealthofmarineenvironment, and variety ofecosystemservicesthatare essential Sponges are keyspeciesthatprovide awide and coralligenousformationsattheshelfedge. areas, andinparticular, were foundontherocks communities were observedinallofthesurvey and softbottoms.Duringtheexpedition,sponge communities ondeep-,bothinhard sponge speciesthatcancreate important In theMediterranean,there are hundreds of Sponge aggregations coastline degradation). eroded riverbasins,untreated wastewater, and increased turbidity and sedimentation (e.g., communities, suchasland-basedsources of more distalthreats to coralligenous and maërl consider measures toreduce theofimpacts them, suchasanchoring.Itisalsoimportantto other activitiesthatdirectly orindirectly damage systems, includingimpactsfrom fisheriesand to limit localised threats to these important would bethecreation ofmarine protected areas they support.Onekeyelementoftheseplans damage these habitats or thecommunities that place toavoidanyactivitiesthatmaypotentially possible andmanagementplansshouldbeputin Lebanese watersshouldbemappedassoon of coralligenousandmaërlassemblagesin in linewiththeActionPlan,distribution should prioritisetheirconservation.Specifically, both biodiversityandforfisheries,Lebanon Given theimportanceofthesehabitatsfor 90 87

Spongia, allelopathic, 81,82,83,84,85,86 89 Many of these 91,92

Fascispongia, 88 andother 93

53

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Agelas,94,95 Sarcotragus, and Axinella;96 these Another scleractinian species, star coral (Madracis taxa were the most commonly observed ones pharensis), is a colonial, mainly encrusting during the research expedition. Compounds of coral with a wide distribution in the tropical and potential relevance for cancer treatment have temperate waters of the Atlantic Ocean, from the been confirmed in many of these species.97 to the Mediterranean98. This species mainly occurs in shallow waters. In the Western Most of the sponge species that were Atlantic, it is generally zooxanthellated, while in the documented during the expedition were living in Mediterranean, it typically lacks zooxanthellae.99 association with coralligenous habitat. Therefore, In Lebanese waters, it represents one of the many of the recommendations for protecting most characteristic communities growing on coralligenous would also be beneficial for the hard bottoms in the region of the continental protection of sponge aggregations. Those shelf break, in canyon heads, and in the upper sponge aggregations that were not linked part of the bathyal zone. Madracis pharensis has with coralligenous assessmblages should be also been found close to hydrothermal vents in considered as vulnerable marine ecosystems , covering a high proportion of the primary (VMEs), and appropriate measures be taken for rock, and representing one of the main and their protection (either under the General Fisheries characteristic species of constructing corals100. In Commission for the Mediterranean, or under the the deep circalittoral and upper bathyal areas off 73 Dark Habitats Action Plan of UNEP-MAP RAC/ Lebanon, this species must also be considered as SPA). In addition, in the case of one particular one of the main bio-constructors, which frequently species (Sarcotragus foetidus), its protected occurs together with oysters (Neopycnodonte status under Annex II of the SPA/BD Protocol of cochlear) and sponges and provides an important the Barcelona Convention means that Lebanon habitat for many species. The primary threats to must take legal steps to ensure the protection of this species are those related to global climate both the species and its habitat. change, such as ocean acidification and the 101 Scleractinians (Stony corals) increased incidence of coral diseases. Only a few Mediterranean coral species belong to Tree coral (Dendrophyllia ramea) is a tree- the scleractinians (also known as madreporarians shaped coral found in the upper bathyal zone. or stony corals), which are the type of corals This species is distributed in the Mediterranean that forms the extensive reefs of the tropics. In Sea and in the eastern Atlantic Ocean, where it the Mediterranean, the great majority of these occurs around the African coast and the Iberian species are only represented by small colonies or Peninsula.102,103 It has also been recorded from the in isolation (or as part of reefs created by other , in Mozambique.104 In the Western species). Mediterranean and the Eastern Atlantic, it on hard rocky surfaces, from very shallow waters Five main types of stony coral communities until the shelf break, and on top of .105, were found in Lebanese waters during this 106,107 In the Eastern Mediterranean, the species survey (cf. Anomocora sp., Madracis pharensis, has only been recently described from Greece,108 Dendrophyllia ramea, Caryophyllia (Caryophyllia) Cyprus,54 and now Lebanon. Contrary to what calveri and Desmophyllum dianthus). has been observed in Western Mediterranean communities of D. ramea, the eastern ones seem One type occurred in relatively shallower waters in to prefer soft bottoms, in deep waters close to the circalittoral zone, composed by small colonies the shelf break. Therefore, they may be vulnerable of a scleractinian coral that has not yet been to fishing gears that directly impact the seabed, described and may be new to science (possibly such as bottom trawls, bottom longlines, set nets, Anomocora sp.). This species was found on soft and traps, whether actively used or discarded on sandy bottoms or, in some cases, attached to the seabed. This species may also be affected natural and artificial hard bottoms between 60 m by other types of marine litter, as well as by and 90 m depth. Vulnerable, is considered threatened. ItisRedListedas sedimentation resulting from terrestrial run-off were found. colonies, upto44suchspeciesofepibionts can liveintheirbranches.InthenearbyCypriot the colonies.Also,manyassociatedspecies coral allowmanyspeciestolivewithinornear The three-dimensional structure created bythis etc. deployment ofsubmarinecablesandpipelines, of waste,energydevelopmentandexploitation, fishing lines,climatechange,mining,dumping human activities,includingbottomtrawling, Both speciesare considered vulnerabletodifferent Chile. the Mediterranean Sea, and the Pacific coast of has awidedistribution,across theAtlanticOcean, coral community on deep-sea reefs. can formdensecoloniesandevenformspartofthe not usually occurs in large numbers, D. dianthus sediment ‘rain’from does above.WhileC. calveri in deep-,eventoleratinghighlevelsof fauna foundonverticalwallsandfossilreefs grow insmallcolonies.Theyare themainerect corals from thedeepbathyalzonethatsometimes cup coral(Desmophyllumdianthus)are solitary calveri andcockscomb (Caryophyllia) Caryophyllia (i.e., Tarablus/Batroun). introduced forthearea where itwasdocumented its habitat. including prohibiting damageanddestructionof possible protection andrecovery ofthespecies, Lebanon islegallyobligedtoensure themaximum Therefore, its protection is a legal obligation, and SPA/BD Protocol oftheBarcelona Convention. North-East AtlanticandMediterraneanSea. resulted initbeingRedListedasEndangered, 114 112 . dianthus have Population declines in D. isonlyfoundinthe Incontrast,C. calveri 41 53 Legalmeasures shouldtherefore be 110 and is also listed on Annex II ofthe andisalsolistedonAnnex II . ramea IntheMediterranean,D. 111 D. dianthus 109 113 53 .

Habitats ActionPlanofUNEP should therefore be includedunder the Dark waters, and over a wide bathymetric range. It other partsoftheMediterranean),butindeeper caespitosa (themainscleractinianreef-builder in a bioconstructing role similar to that of However, inLebanesewatersitappeared toplay habitat-forming speciesintheMediterranean. latter coral is not usually considered to be a major pharensis) formed large communities. The (M. structures ontheseabed,whilestar coral ) provided important three (D. ramea diverse benthiccommunities.Inparticular, tree coral species clearlyformedhabitatsthatsupported In theLebanesedeep and managedaccordingly. conservation, inventories ofplacesnaturalinterest for their the selection of sites to be included in the national Reference ListofMarineHabitatsproposed for described aboveare includedintheUNEP-MAP Many ofthecommunity types ofstonecorals (i.e., St. George). introduced forthearea where itwasobserved and so legal measures of protection should be this speciesthemaximumpossibleprotection, ramea,Lebanonisrequired togrant with D. BD Protocol oftheBarcelona Convention.As IIoftheSPA/ and inits2017listingonAnnex – – – – – – – – (Caryophyllia) calveri (Caryophyllia) Bathyal mudwith Bathyal r Bathyal r Bathyal r 115 Caryophyllia ock withCaryophyllia ock withMadracispharensis ock withDesmophyllumdianthus including: ‑ sea, somescleractinian Dendrophyllia ramea ‑ MAP RAC/SPA, Calyx nicaeensis ‑ dimensional Cladocora 73

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Deep-sea oysters formed the most important mollusc aggregations found, with up to 1000 individuals/m2.

Other stony coral species also represented part colonisation by hydrozoans, contributing further of important communities, while some species to the decline of the colonies.122 In Norway, may be new to science, such as the potential S. pallida is considered one of the sea fan Anomocora species found during the expedition, species that must be prioritised for protection, which could be endemic to the area. The under the Monitoring Guidelines for oil drilling.123 ecosystem role of species such as these, about This community type is included in the reference which so little is known, remains to be discovered. list of Mediterranean Marine Habitats proposed Gorgonian/sea fan communities for the selection of sites to be included in the national inventories of places of natural interest for More than 20 species of gorgonian have been their conservation,115 as ‘Bathyal rock with Swiftia described from the Mediterranean Sea.116 This pallida’. expedition marked the first time that sea fans had been discovered in Lebanon. Gorgonians are very uncommon in the Levantine Sea, and here – as with the rest of Swiftia pallida is a gorgonian known as northern the Mediterranean – it is a community type that sea fan, which is distributed throughout the entire must be protected and managed, under the Mediterranean Sea (from the Alboran Sea to Dark Habitats Action Plan.73 Such measures the Levantine Sea)117 and in the Atlantic Ocean may include the identification and protection of (mainly in the eastern part from Norway118 to threatened populations, the designation of MPAs, and the Macaronesian Archipelagos).107 and measures designed to address specific It has also been found in the Western Atlantic, in identified threats. Given the known sensitivity of deep waters off of .119 In general, S. pallida to impacts associated with exposure its distribution appears to be limited by water to oil, the planned exploratory drilling for oil and temperature; in the Mediterranean it is normally gas in Lebanese waters is reason for particular found between 200 m and 700 m depth, while in concern. northern areas it can be found in much shallower water. It can be fixed to rocky bottoms, as well as Sea pen fields on gross detritic beds with cobbles, and tolerates Sea pens play an important role in forming ‘animal some degree of sedimentation.117 forests’ on soft bottoms, thereby providing habitat S. pallida is vulnerable to a range of human for a variety of associated species, including those impacts, as has been shown across various of commercial value. The most widely and densely regions where it occurs. In Scotland, this species distributed sea pen aggregations in Lebanese has been recognised as one of the more vulnerable waters are those of red sea pen (Pennatula rubra) gorgonians to temperature increases,120 and could on soft bottoms between the shelf break and the therefore disappear from part of its current range head of the canyons. In some locations they also of distribution in response to climate change. appeared in mixed aggregations with slender sea pen (Virgularia mirabilis). In the Mediterranean The species has also been shown to be sensitive Sea, P. rubra can reach densities of more than to oil leaks and spills. After the Deepwater Horizon 9000 colonies per square kilometre.124 In Syrian accident in the , different species of waters, communities dominated by P. rubra and sea fans – including S. pallida – exhibited “signs the echinoid Shizaster canaliferus have been of stress response, including excessive mucous identified as the ones with the highest levels of production and retracted polyps” as described biomass (30g/m2) and with a high associated 121 by White et al. (2012). These toxic oil effects species richness.125 Alongside its high ecological can be long-lasting. When areas impacted by value, P. rubra is also considered threatened in oil leakage were revisited several years later, the the Mediterranean; it is Red Listed as Vulnerable most affected sea fans showed high levels of because of inferred population declines of must beconsidered oneofthekeydeep rubra),and were formedbyred seapen(P. largest aggregations of sea pens identified caused byfishingactivities. identified asVMEindicatorsfrom adverseimpacts Mediterranean, toprotect thosespecies the GeneralFisheriesCommissionfor Plan andmeasures beingdevelopedwithin out inlinewithboththeDarkHabitatsAction management of this species should be carried vulnerability to human impacts. Conservation and biodiversity andcommercial fisheries,andits Mediterranean, duetoitsimportanceforboth protection inLebanesewaters,andthewider quadrangularis) is also a priority for pen (F. community typesinLebanon.Thetallsea (VME) indicatorspecies considered to be a vulnerable trawl fisheries. among thespeciesdiscarded asbycatchfrom fishing activities,with red seapenknowntobe declines are dueinlarge part totheimpactsof as Vulnerable intheMediterraneanbyIUCN. Parapenaeus longirostris. and as thecrustaceansNephropsnorvegicus bottoms, and supports commercial species such an importantrole asahabitat builderonsoft down to604m.Thisspeciesisknownplay quadrangularis) wasfoundonmuddybeds In deepestareas, thetall seapen(Funiculina years. approximately 40%over 20 of upto1000individuals per square metre. natural and artificial, reaching concentrations These oysterscansettleon hard substrata,both on hard bottomsintheheadsofcanyons. objects, andwere alsoveryfrequently observed cases overtheshelf,coveringrocks andartificial These aggregations were observedinsome a distributionrangingfrom 60mto250depth. deep in Lebanesewaters were thedense mantles of The mostimportantmolluscaggregations found Oyster aggregations/concretions habitats. Mediterranean ActionPlanforprotecting dark included ashabitatbuildersunderthe All ofthesepennatulaceansare keyspecies reefs andbanksintheMediterranean(e.g.,on They havebeenfoundindense groups forming ‑ sea oyster(Neopycnodontecochlear),with 73,117 IntheLebanesedeep-sea, 126,127,128 132 andhasbeenassessed 129,130,131 Tall sea pen is 53 These 133 ‑ sea 134

from trawlersfishingin Tunisian waters. example, protected accordingly. can havedetrimentaleffects uponthem. occurs, includingbanningdemersalfisheriesthat damage totheseafloorwhere thishabitat implement conservationmeasures toprevent bottoms. Therefore, it is recommended to complexity ofthehard andmixedsediment These biohermssignificantlyincrease the worms, sponges,bryozoans,andcorals. fasciata), as well as sessile organisms, such as Thorogobius ephippiatus,andLappanella fishes (e.g.,Anthiasanthias,Callanthiasruber attenuata, andEchinaster(Echinaster)sepositus), as echinoderms(e.g.,Cidariscidaris,Hacelia species of mobileorganisms were observed, such sponges. Intheseaggregations, avarietyof (primarily oysters alsoformedmantlestogetherwithcorals In theaggregations observed inLebanon,the and variousechinoderms. large bryozoans,other molluscs, polychaetes, in associationwiththeseaggregations include other species.Themostcommonspeciesfound Habitats ActionPlan, as keyhabitat-formingspeciesundertheDark and corals).Assuch,theyshouldbeconsidered organisms maysettle(e.g.,sponges,polychaetes, declines andcreates substrateuponwhichother in siteswhere otherspecieshaveundergone species, whichinsomecasesestablishesitself of theirremains. N. cochlear complexity, whetherasreefs of livingoysters,or to bothincreased biodiversityandhabitat similar role tocoralreefs, inthattheycontribute Oyster aggregations inLebanesewatersplaya on theGorringeBank;Atlanticseamounts; in wideaggregations thatcarpet theseabed(e.g., and intheAzores) Alboran Sea or sub-lethaleffects” upontheoysters. discharges anddredge disposalmayhavelethal community type,giventhat“pollutedwater is alsonecessaryfortheconservationofthis these direct impacts,waterqualitymanagement the Chella bank; hasbeenfoundindiscardsN. cochlear Madracis pharensis),polychaetes and 136 ; andinthisstudy, inLebanon),and 105 138,139 in the Gulf of Lions; 73 andserveassubstratefor andbemanaged is anopportunistic 141 140 135 Beyond 140 in the For 137

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Echinoderm-dominated communities Lebanese canyons studied. Holothurians are known to create large aggregations that play a Various echinoderms are known to form large crucial role for deep-sea ecosystems;142,142,151,152 aggregations on the seabed, providing different sea cucumbers excrete inorganic nitrogen and ecosystem services that are key for the entire phosphorus, enhancing the productivity of benthic ecosystem.142 In Lebanese waters, the largest biota.153 Therefore, they are key elements for the such concentrations of echinoderms observed cycling of different nutrients, and put sources of during the expedition were, on the one hand, food back into circulation and available to other those formed by sea urchins and, on the other species. These include other echinoderms, such hand, those made up of deep‑sea cucumbers. as sea urchins that feed on holothurian fecal With resepect to sea urchins, the main casts,154 reintroducing nutrients into the food web. aggregations were formed by hatpin sea urchin As explained by Purcell et al. (2016)153 impacts (Centrostephanus longispinus) around and in affecting the abundance of holothurians could coralligenous beds, and by long-spine slate pen “decrease sediment health, reduce nutrient sea urchin (Cidaris cidaris) on soft bottoms in recycling and the capacity of ecosystems to buffer deeper areas. against ocean acidification, diminish biodiversity C. longispinus has not typically been described of associated symbionts and reduce the transfer as creating large aggregations.143,144 This species of organic matter from detritus to higher trophic appears to avoid ; therefore, it normally levels”. appears in areas below 30 m depth. Its The roles played by echinoderms in deep‑sea distribution is also restricted by water ecosystems are poorly understood, both in temperatures, as it only resists a limited thermal general, and in Lebanese waters. In a limited range of between approximately 12°C and number of cases, the value and importance of 20°C.145 The species is likely to play a key such habitats has been determined, such as the role in maintaining the health of coralligenous identification of beds as ‘essential fish concretions, as has been documented in other habitat’ for commercial species,155 or of certain Centrostephanus species worldwide.146 In the holothurian species that are clearly very important Mediterranean, this species is included on the for food and biogeochemical cycling. For many Annex II list of endangered or threatened species, other echinoderms, however, little is known. Their and therefore requires legal measures to ensure function and contribution to deep‑sea areas the protection of both it and its habitat.41 deserves further study, and should be taken into The other main aggregating urchin species, account to improve the management of these C. cidaris, is another echinoid species that lives ecosystems in Lebanese waters and elsewhere. far from sunlight. This urchin can be found much Fossil reefs deeper than C. longispinus; in the Mediterranean it is normally found in soft and hard bottom Coral aggregations act as a reef, providing shelter areas between 50 m depth and close to 600 m and substrate for many species, both when the depth,147 although its depth distribution has polyps are still alive, and when only a fossil or been mentioned to extend below 2000 m.148 It subfossil structure remains.156 is the most conspicuous sea urchin in Lebanese Although no living deep-sea coral reefs were waters. This species is considered an important found in the Lebanese canyons surveyed during factor influencing marine species aggregations the present expedition, some hard bottoms and distributions, and an important player as a in deep-sea areas were the result of fossilised bioeroder of deep-sea ecosystems, including corals, worms and other species. These fossil those formed by cold-water corals.149,150 reefs act very similarly to rocky bottoms, allowing The other important echinoderms in deep areas species to settle and proliferate. The intricate are sea cucumbers. Both Mesothuria intestinalis morphology of fossil reefs allows rare species to and Penilpidia ludwigi were very abundant in the settle, increasing new biodiversity levels and giving rise to new biological associations. Hard bottoms, whether bioherms or lithoherms, are scarce on systems are minimised oravoidedaltogether. ‘dark habitats’,toensure thatthreats to these managed underthesame frameworkasother Convention shouldensure thatfossilreefs are and otherContractingParties totheBarcelona importance mustbetakenintoaccount.Lebanon a diversityofspecies.Assuch,their ecological such substrataare veryscarce), andsupporting valuable hard substrateindeep-seaareas (where clearly neverthelessstillactasreefs, byproviding Although fossilreefs are nolongerlivingreefs, they other organismsthatlivewithinthereef structure. fauna ofcrustaceans,echinoderms,fish,and like sp.,worms pallida, scyphozoanslikeNausithoe calveri, soft coralslike the gorgonian Swiftia like species like Farrea bowerbanki, other living corals expedition were foundtohosthexactinellid Lebanese fossilreefs surveyedduringthe scleractinians. forming patches or thickets of fossil or subfossil are commonlyfoundintheMediterraneanSea Madrepora oculataandDesmophyllumdianthus Cold watercoralslikeLopheliapertusa, ecosystems intheworld. the developmentofsomemostproductive marine bottoms,buttheyprovide substratefor stages ofreef building. new coralreefs throughout different successional species, includingnewcoralsthatcangivebirthto framework couldpromote furthersettlementand Lepidorhombus whiffiagonis Caryophyllia (Caryophyllia) (Caryophyllia) dianthusandCaryophyllia D. Vermiliopsis monodiscus,etc.andvagile 157,158,159 Dead corals and coral 160 Jounieh Beirut key observationsincluded: of ecologicalandconservationinterest. Ingeneral, areas surveyed,showing selected mainfeatures them. Table 1provides acomparisonofthefive and communities of interest that occurred within total biodiversity, andintheparticularspecies supported. Nevertheless,theyvariedintermsof of themarinehabitatsandcommunities that they The surveyed areas were broadly similar in terms AREAS SURVEY – – – – – – – – – – The ar also mostcommonintheBeirutarea. stony coralsinthecanyonheadswere The aggregations formedby oystersand and abundantofallthoseobserved. fields thatwere themost widespread abundant inthearea, including seapen Some keycommunitytypeswer Leptasterias sp.). be Atlantic(i.e.,Haceliasuperbaand species thatare typicallyconsidered to Farrea bowerbankii and two starfish from Jounieh,suchastheglasssponge during theexpeditionwere documented importance. aggregations beingofparticular reefs (inthedeepestareas) andsponge nearly 300 speciesidentified. this area. (cf. coral discovered during the expedition starfish). The potential new species of and thepotentialnewspeciesof urchin (Centrostephanus longispinus) aggregation oftheprotected hatpin and echinoderms(includingthelargest abundance ofscleractinians,bryozoans, some keyspeciesofinterest, withahigh Many oftherar wer All ofthekeycommunitytypesdescribed r The highestdiversityofspecieswas ecorded from intheJounieharea, with e present inthearea, withfossil sp.)wasalsofoundin Anomocora ea appeared tobeimportantfor est speciesobserved e highly 59

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–– Jounieh provided the first-ever record Tarablus/Batroun of a gorgonian (Swiftia pallida) from – The Tarablus/Batroun area was Lebanese waters. – characterised by a diversity of key habitat –– Similar to the Beirut area, some key types, and a variety of protected and/or habitat-forming species were highly threatened species. abundant, such as sea pens. ––The occurrence of hard substrata was Sayniq relatively high in the area, partly reflecting the high abundance of habitat-forming –– The area accounted for the second- species. highest number of identified species. ––The area was important for more than –– Among the key community types in the half of the protected and/or threatened area, coralligenous formations, maërl species documented. These species beds, and sponge aggregations were included the threatened and Annex II- particularly well represented. listed tree coral (Dendrophyllia ramea). –– Particular species of interest included the This species was found to occur in large bryozoans (Hornera frondiculata) deep areas and on muddy bottoms, an and rare, new, and protected sponge occurrence that was only recently first species. observed in deep-sea areas near Cyprus.

St. George The three categories of features included in –– The most extensive and diverse Table 1 are summarised in Figure 41, showing the coralligenous and maërl habitats were sites of highest importance for key communities, found in the St. George area. protected and/or threatened species, and species of particular interest. When combined, –– Two chondrichthyan species that are this information indicated a series of sites of classified as Near Threatened (Chimaera highest conservation interest, spread across the monstrosa and Dipturus oxyrhincus) five survey areas (Map 15; see Annex 2 for further were observed only from this area. details). –– St. George was also the only area in which the Annex II-listed species of coral Desmophyllum dianthus was observed, in association with the deep‑sea fossil reefs.

© OCEANA/ Enrique Talledo Figure 41. Relative conservation interest of the study areas, according to: a) key community types; ) protected and/or threatened species; and c) other species of general scientific interest. 61

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Table 1. Main features of interest in each of the study areas.

Beirut Jounieh Sayniq St. George Tarablus/ Batroun Total number of identified taxa 214 299 250 210 214 Key community types Coralligenous formations Coralligenous formations (dead) Fossil reefs Relative abundance Rhodolith/maërl beds (estimated) Echinoderm-dominated communities High Gorgonian/sea fan communities Medium Oyster aggregations/concretions Low

Sea pen fields Sponge aggregations Vermetid reefs Protected/threatened species Chordata Chimaera monstrosa Chordata Dipturus oxyrhincus Chordata Epinephelus marginatus

Chordata Merluccius merluccius Abundance Chordata Tursiops truncatus (number of individuals per Dendrophyllia ramea area) Cnidaria Desmophyllum dianthus ≥10 Cnidaria Funiculina quadrangularis >1 and <10 Cnidaria Pennatula rubra 1 Crustacea Scyllarus arctus Echinodermata Centrostephanus longispinus Pinna rudis Mollusca Tonna galea Porifera Sarcotragus foetidus Porifera Spongia (Spongia) officinalis Species of interest Annelida Sabella spallanzanii Brachyopoda Gryphus vitreus Bryozoans Adeonella pallaisi

Bryozoans Caberea sp. Abundance Bryozoans Hornera frondiculata (number of individuals per Bryozoans Reteporella spp. area) Chordata Etmopterus spinax ≥10 Cnidaria cf. Anomocora sp. >1 and <10 Echinodermata Hacelia superba 1 Echinodermata Leptasterias sp. Echinodermata Luidia sp. Foraminifera Pelosina cf. arborescens Porifera Axinella spp. Porifera Axinella sp. nov (possible) Porifera Calyx nicaeensis Porifera Farrea bowerbankii Sites of highest conservation interest. Map 15. Sitesofhighestconservation 63

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MAIN THREATS Threats observed Marine litter Across all of the areas studied, surveys revealed species were documented: the algae Caulerpa the pervasive threat of marine litter, in all habitat scalpelliformis; the fish species cf. Cheilodipterus types, and at all depths. In some cases, the novemstriatus, Equulites klunzingeri, Fistularia sources of this waste were likely coastal, with commersonii, Nemipterus randalli, Ostorhinchus ocean currents carrying litter offshore, while in fasciatus, Pterois miles, Sargocentron rubrum, other cases it was apparent that large objects Siganus luridus, Stephanolepis diaspros and such as appliances, tyres, and barrels had been Torquigener flavimaculosus; and the mollusc dumped directly at sea. These observations species Fulvia fragilis, Mactra olorina, and Murex reflect the broader and significant challenges forskoehlii. that Lebanon faces regarding the disposal and Non‑native species are of particular concern in management of the country’s solid waste.161 cases where they become invasive, establishing The environmental benefits of improved waste viable populations outside of their natural management policies and practices would be far- ranges and causing changes to the structure reaching, not only limited to terrestrial and coastal and function of local ecosystems, such as by systems, but extending to deep-sea areas such displacing native species, altering habitats, as those surveyed here, and beyond. or modifying foodwebs.165,166 Such changes Fisheries can have significant ecological and economic While direct impacts of fishing activities on consequences, with resulting impacts on both 167 benthic communities were limited (largely due to biodiversity and ecosystem services. Of the the lack of bottom-contact fisheries), discarded alien species observed during the expedition, five fishing nets, lines, and traps were also commonly are considered to have invasive characteristics found, particularly around coralligenous habitat, (Fistularia commersonii, Fulvia fragilis, the shelf edge, and the heads of canyons. Sargocentron rubrum, Siganus luridus, and Although deep‑sea benthic habitats and species Stephanolepis diaspros). Both F. commersonii are currently out of the reach of Lebanese and S. luridus have been identified as species fisheries, they are nevertheless clearly affected by with high impacts on ecosystem services or 167 anthropogenic activities. biodiversity.

Alien species In 2016, 46 scientists from the Mediterranean region signed a manifesto expressing their Five percent of the species that currently inhabit concern about bio-invasions and their effects the Mediterranean Sea are of alien origin.162 Many on the Mediterranean marine ecosystem, and of these species have entered from the Red Sea; calling on parties to the Barcelona Convention more than 60% of the 614 multicellular non- to take immediate action.168 UNEP‑MAP has indigenous species with populations established also expressed its concern about the impact of in the Mediterranean have reached this sea via invasive alien species, and highlighted the need the Suez Canal.163 As a result, the Levantine Sea for Mediterranean countries to take action to is one of the most affected areas in the region. avoid their intentional or unintentional release, and According to the Marine Mediterranean Invasive associated harmful impacts on the ecosystem.169 Alien Species database, which is maintained by Given the scale of the problem, effective solutions UNEP‑MAP RAC/SPA, 215 marine species have will depend on joint actions implemented by been recorded from Lebanon that are either alien Mediterranean countries, including Lebanon, to or cryptogenic (i.e., of unknown origin).164 both prevent the introduction of further exotic This expedition was focused on deep-sea areas, species in the future, and to limit the impacts of while most alien species in the Mediterranean those that have already established themselves in are associated with shallow waters. However, Mediterranean waters. even in the deeper areas surveyed, various alien decreased benthicstructuralcomplexity. increased sedimenterosion, asaresult of in favourofinfaunahasbeenshowntocause In particular, thereduction ofepibenthicfauna associated with changes in sediment erosion. the abundanceofbenthicspecieshavebeen In otherareas, liketheNorthSea,changesin a meansofreducing environmental degradation. considered, alongwithland-basedmeasures, as impacts, orshippingactivities)andmustbe as resuspension ofsediment duetofisheries be useful in the face of erosive impacts (such protective benefitsofthese ecosystemscan marine ecosystem. and increasing thegeneralresilience ofthe thereby decreasing impactsonthesebottoms erosion andresuspension inimportanthabitats, occur. Thiscomplexitycanhelptoreduce well aswhere theremains oftheseorganisms provide living three-dimensional structures, as environment is higher in areas with epifauna that Ecosystem complexityinthedeep-sea Erosion Other threats Mediterranean (over 65 billionbarrels). fields in surrounding areas, like the North African small incomparisonwiththemost important oil 1.7 have beenestimatedatbetweenroughly Oil reserves Mediterranean intheEastern Oil exploration exploration isexpectedtocommencein2019. discoveries, andreserves are notproven. Drilling To datethere havenotbeen anylargecommercial approximately 25-30trillioncubicfeetofgas. ofbarrels ofoiland are estimatedat660 million In Lebanesewaters,hydrocarbon resources cubic feet. trillion of gas, accounting for an estimated 122 the mostimportantpredicted resources are those Mediterranean reserves,Of the estimated Eastern been demonstratedtoreduce erodibility. of spongespiculeswithinsoftsedimenthave ‑ . billionbarrels. Thisamountisrelatively 2.5

For example,thepresence 170 86 The 171,172 173

ecosystems. extensive areas anddestroy ordamagevulnerable in otheraccidentandroutine spillscouldpollute biologically yet.Theimpactofleaks,asproven species andhabitatsinareas notsurveyed to globalclimatechange,canalsoaffect other Oil and gas exploration, apart from contributing rosefish, seabreams, andforkbeards). commercial speciessuch asshrimps,dentex, some stonycoralswere documented, aswell and somepatchesofrocky bottoms(inwhich especially arthropods, fishesandechinoderms), (in which dozens of species were documented, drilling woulddirectly affect bothmuddybottoms the exploratoryarea. Inthese areas, exploratory surveyed, SayniqandBeirut,overlapdirectly with during thisexpedition. Only twoofthe areas these blocksare deeperthan theareas surveyed 16,mostof into tenblocks.AsshownonMap by UNEP in linewiththerelevant ActionPlansdeveloped future threats. Thisplanshouldbedeveloped in Lebanesewaters,thefaceofcurrent and vulnerable ecosystems, habitats,andspecies development ofaplantomanageandprotect provides a scientific foundation for the along the Lebanese coast. This information information aboutdeep-seabenthiccommunities Lebanon Expeditionprovided awealthof The dataobtainedfrom the2016Deep-Sea CONCLUSIONS The exploratoryarea covers 17901m Mediterranean basin. help safeguard thenaturalcorridorofEastern marine protection inLebanesewaters,andto to ensure thecoherence andconnectivityof canyons), habitats,orcommunitytypes,inorder important geological features (e.g.,submarine MPAs. These areas may be designated to protect under theframework of aLebanese network of of thisplanwillbethedeclarationnewMPAs of theBarcelona Convention.Onecriticalelement species listedonAnnex II oftheSPA/BD Protocol and with Lebanon’s legal obligations to protect developed toprotect VMEswithinGFCMfisheries, ‑ MAP RAC/SPA, with measures being 2 , divided 65

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Map 16. Oil and gas exploitation blocks, in relation to survey locations. Acknowledgements expedition, andthecrew oftheSeaPatron. Forces, fortheirsupportandassistanceduringthe these waters. Oceana also thanks the Lebanese Navy as welltheirextensiveknowledgeandexpertiseon CNRS Abed ElRahmanHassounandAliBadreddine from for theirvaluablehelpwithspeciesidentification;Dr. Marie Grenier, Vacelet Dr. Jean Pérez andDr. Thierry JoséTemplado, AlfonsoA.RamosEsplá,Dr. Dr. ÓscarOcaña, authors wouldalsoliketothankDr. and IUCN,fortheirfullsupportcollaboration.The Scientific Research (CNRS Environment, the Lebanese National Council for of thisproject, includingtheLebaneseMinistryof MAVA Fondationpour la Nature, andtoallpartners Oceana isgratefulforthegenerous supportof ‑ L forhavingshared thetimeonboard withus, ‑ L), UNEP © OCEANA/EnriqueTalledo ‑ MAP SPA/RAC 67

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Annex 1 TABLE A1. LIST OF SPECIES IDENTIFIED Species are listed according to survey area, and to sampling method. (#) Species spotted incidentally from/at the deck of the research vessel; (+) Exotic species; (*) Possible new species; (~) Possible dead coralligenous formations.

Tarablus/ Min. depth Max. depth Beirut Jounieh Sayniq St. George ROV Grab Batroun CHLOROPHYTA Caulerpa scalpelliformis + 61.3 82.4 X Chlorophycea indet. 36.0 483.7 XXX X Codium bursa 64.4 64.4 X X Palmophyllum crassum 67.2 67.2 X X Ulva sp. 58.5 58.5 X X Valonia macrophysa 59.9 59.9 X X

Tarablus/ Min. depth Max. depth Beirut Jounieh Sayniq St. George ROV Grab Batroun OCHROPHYTA Dictyota dichotoma 62.7 71.4 X X Lobophora variegata 49.2 71.4 XX X vulgare 62.7 68.6 X X Zanardinia typus 68.6 68.6 X X

Tarablus/ Min. depth Max. depth Beirut Jounieh Sayniq St. George ROV Grab Batroun RHODOPHYTA Amphiroa rigida 48.4 63.6 X X cf. Corallinales indet. 59.9 61.3 X X cf. Corallinales indet. (x) 86.6 255.4 X XXX Corallinales indet. 36.0 122.6 X X XXXX Lithophyllum sp. 63.0 100.4 X X X Lithophyllum incrustans 46.1 80.4 XX Lithothamnion sp. 61.3 71.4 X X Mesophyllum lichenoides 63.6 68.9 X X Mesophyllum sp. 36.0 85.5 XXXXX Peyssonnelia rubra 66.4 66.4 X X Peyssonnelia sp. 68.3 75.1 X X Peyssonneliaceae indet. 36.0 90.9 X X Phymatolithon calcareum 45.6 71.7 X XX Phymatolithon sp. 61.6 61.6 X X Rhodophyta indet. 36.3 82.4 XX X Rhodophyta indet. (dead) 297.0 297.0 X X Rhodymenia sp. 63.6 70.0 X X Spongites fruticulosa 45.6 71.7 XX FORAMINIFERA Adelosina sp. Ammodiscus 3063.0 63.0 Heterostegina depressa+ Foraminifera indet. Komokioidea indet. f ooiie ne.6. 80.7 61.9 Miniacina miniacea cf. Komokioidea indet. Miliolinella subrotunda Pelosina cf. arborescens Pyrgo inornata Pyrgo sp. Quinqueloculina sp. Thurammina papillata PORIFERA PHORONIDA Agelas oroides Axinella sp. Axinella damicornis Antho sp. Amphilectus fucorum Clathrina clathrus Chondrosia reniformis cf. Chelonaplysilla noevus Calyx nicaeensis cf. Biemna variantia Biemna variantia Axinella verrucosa cf. Chondrosia reniformis Axinella sp.* Crambe crambe Corticium candelabrum Cliona rhodensis cf. Crambe crambe i.dphMx et ertJuihSyi St. George Sayniq Jounieh Beirut Max. depth Min. depth i.dphMx et ertJuihSyi St. George Sayniq Jounieh Beirut Max. depth Min. depth St. George Sayniq Jounieh Beirut Max. depth Min. depth 0. 1015.0 906.0 1. 113.9 113.9 2. 124.0 122.6 171.3 171.3 124.0 178.3 122.6 178.3 525.9 525.9 6076.0 76.0 6076.0 76.0 93129XX X 102.9 99.3 37175.2 53.7 68348XXXXXXX X X X X X X 344.8 56.8 3063.0 63.0 37950XXXXXX X X X X X 905.0 53.7 3063.0 63.0 4076.0 64.0 3063.0 63.0 70100XXXXXX X X X X X 180.0 79.9 47.0 79.9 60284XXXXXX X X X X X X 298.4 82.1 96.5 36.0 64.1 36.0 87.5 64.1 348. X X X 85.5 74.5 73.4 83.2 53.7 78.7 82.1 82.1 607. X X 78.7 X 90.0 64.1 36.0 82.4 62.7 64.1 82.4 64133.6 46.4 X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X Tarablus/ Tarablus/ Tarablus/ Batroun Batroun Batroun X X X X X Grab ROV O Grab ROV Grab ROV X X X X X X X X X X X X X X X X X X X X X X X X X X 69

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Demospongiae indet. 46.1 377.1 X X X X X X X Diplastrella bistellata 127.7 127.7 X X Dysidea fragilis 63.0 82.4 X X X Farrea bowerbanki 857.0 857.0 X X Fasciospongia cavernosa 59.9 83.5 X X X X X X X Haliclona cf. (Gellius) angulata 70.9 70.9 X X Haliclona (Reniera) 68.6 78.2 X X mediterranea Haliclona sp. 48.4 88.3 XXXXX Hexactinellida indet. 863.4 863.4 X X Hexadella dedritifera 167.6 167.6 X X cf. Hexadella detritifera 148.5 148.5 XX cf. Hexadella pruvoti 133.6 174.9 XXX Hymedesmia sp. 76.0 76.0 X X Ircinia sp. 49.5 97.6 X X X XX cf. Ircinia sp. 53.7 133.6 X X X Ircinia variabilis 59.9 100.4 X X X Mycale sp. 172.7 172.7 X X Myxilla sp. 207.8 207.8 X X Oscarella lobularis 67.2 68.9 X X Petrosia (Petrosia) ficiformis 36.0 37.4 X X Phorbas fictitius 56.0 64.1 X X X Plakina monolopha 127.7 207.8 X X X Plakortis simplex 173.0 173.0 XX Pleraplysilla spinifera 78.5 100.4 X X X X Porifera indet. 46.4 773.7 X X XXX Prosuberites longispinus 39 76.0 76.0 X X Sarcotragus foetidus 71.2 102.1 X XX Sarcotragus sp. 60.7 88.9 X X XX Sarcotragus spinosulus 46.1 116.1 X X X XXX Scalarispongia scalaris 71.7 78.7 X XX Spongia (Spongia) officinalis 65.0 82.4 XX XX Thenea sp. 525.9 533.5 X X Timea geministellata 90.0 90.0 XX Timea stellata 90.0 90.0 XX Topsentia calabrisellae 39 76.0 76.0 X X Ulosa stuposa 70.6 71.7 X X

Tarablus/ Min. depth Max. depth Beirut Jounieh Sayniq St. George ROV Grab Batroun CNIDARIA cf. Abietinaria abietina 126.0 126.0 X X Actinaria indet. 65.8 674.4 X X X Actiniidae indet. 529.3 529.3 X X 86.6 113.9 X X Alcyonium palmatum Aglaophenia sp. Aglaophenia octodonta indet. Alcyonium sp. cf. Anomocora sp.* calveri (Caryophyllia) Caryophyllia Arachnanthus sp. indet. cf. inornata (Caryophyllia) Caryophyllia cyathus (Caryophyllia) Caryophyllia smithii (Caryophyllia) Caryophyllia sp. (Caryophyllia) Caryophyllia Cerianthus membranaceus Ceriantharia indet. membranaceus cf. Cerianthus Dendrophyllia ramea Cnidaria indet. Cerianthus sp. cf. Edwardsiella sp. Eudendrium sp. Epizoanthus sp. Desmophyllum dianthus Funiculina quadrangularis cf. Hydroidolina sp. f aolvdeidt 3. 438.4 indet. 438.4 cf. Hormathia digitata Hormathia alba cf. Haloclavidae indet. Madracis pharensis cf. Lytocarpia myriophyllum Lytocarpia myriophyllum cf. Hydrozoa indet. Nausithoe sp. Nemertesia antennina cf. Nausithoe sp. s.247244.7 244.7 cf. Pachycerianthus sp. s.803860.3 860.3 Paralcyonium spinulosum Pachycerianthus sp. Nemertesia sp. 3. 7. X X X X X X X 377.1 207.8 138.6 X 207.8 107.1 107.1 7. 1. X 215.4 175.2 3. 7. X X X X X X X X 773.7 X 776.5 433.7 729.2 537.4 729.2 8. 287.1 287.1 5. 7. X 177.7 151.3 2. 2. X X 729.2 127.2 8. 8. X X 885.3 822.0 885.3 822.0 7. 172.4 529.3 172.4 448.3 703.6 703.6 4. 664.0 521.1 441.3 682.8 521.1 682.8 0. 0. X 604.4 506.2 1. 510.7 979.8 510.7 473.3 2. 122.9 122.9 845.7 845.7 9. 7. X X 771.7 192.4 3. 531.0 531.0 7. 192.4 173.2 1372.6 74.0 61.3 60.2 84145XX X 144.5 48.4 16152XX X 175.2 61.6 5797XXXXXX X X X X X 977 55.7 19849.0 61.9 84152XXXXXX X X X X X 175.2 48.4 1191.1 91.1 92736XXXXXXX X X X X X X 703.6 49.2 13219XX X 221.9 X 61.3 126.6 70.9 61121.2 46.1 81192X 119.2 68.1 X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X 71

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Pennatula rubra 65.0 206.1 X X XXX Pennatula sp. 126.0 198.3 XXX Pteroeides griseum 171.5 172.7 XX indet. 175.2 175.2 XX Sertularella sp. 65.2 90.0 X XX Sideractis glacialis 457.3 457.3 X X Swiftia pallida 531.0 606.6 X X Virgularia mirabilis 66.1 131.9 X X X X X

Tarablus/ Min. depth Max. depth Beirut Jounieh Sayniq St. George ROV Grab Batroun ANNELIDA Annelida indet. 254.6 359.7 X X Bonellia viridis 59.9 977.8 X X X X X X Branchiomma bombyx 93.6 93.6 X X Ditrupa arietina 76.0 858.9 X X X XX cf. Ditrupa arietina 846.2 846.2 X X Errantia indet. 344.8 344.8 X X Filograna implexa 60.7 185.6 X X X XX cf. Filograna implexa 63.6 63.6 X X Hermodice carunculata 53.2 167.6 X X X X X X Lanice conchilega 255.4 405.3 XXX cf. Lanice conchilega 295.0 774.5 X XX Myxicola aesthetica 69.5 74.5 XX Myxicola infundibulum 66.4 66.4 X Polychaeta indet. 56.8 976.7 X X X X X X X Protula intestinum 77.3 77.3 X X Protula sp. 110.0 379.4 X X X Protula tubularia 120.4 644.9 X X X cf. Protula tubularia 192.4 491.0 X X X Sabella cf. discifera 172.7 172.7 XX Sabella pavonina 70.6 177.7 X X X Sabella sp. 77.3 77.3 X X Sabella spallanzanii 80.1 82.1 X X Sabellidae indet. 63.6 472.2 X X X X X X X Seraphsidae indet. 65.5 92.2 X X X X Serpula sp. 138.6 138.6 X X Serpula vermicularis 74.5 602.4 X X XXX Serpulidae indet. 60.2 906.0 X X X X X X X Spirorbis sp. 63.8 63.8 X X cf. Vermiliopsis monodiscus 435.6 674.4 X X X Adeonella pallasii cf. Amathia sp. Alcyonidium sp. Adeonella sp. Bryozoa indet. Bryozoa Bugula sp. Caberea ellisii Caberea boryi Cellaria cf. fistulosa Caberea sp. Cellepora sp. Celleporina sp. Crisia sp. Dentiporella sardonica cf. Escharina sp. Exidmonea atlantica Frondipora verrucosa Hornera frondiculata Margaretta cereoides cf. Licorniavieirai Parasmittina sp. Reteporella grimaldii Pentapora fascialis Reteporella sp. Rhynchozoon neapolitanum cf. Reteporella sp. BRACHIOPODA CHELICERATA ARTHROPODA/ Argyrotheca cuneata cf. Pycnogonida Gryphus vitreus Gryphus Brachiopoda indet. Megathiris detruncata Megerlia truncata Novocrania anomala i.dphMx et ertJuihSyi St. George Sayniq Jounieh Beirut Max. depth Min. depth i.dphMx et ertJuihSyi St. George Sayniq Jounieh Beirut Max. depth Min. depth St. George Sayniq Jounieh Beirut Max. depth Min. depth 7. 171.5 171.5 3. 138.6 138.6 0. 101.0 101.0 0. 101.0 101.0 1. 618.7 618.7 6. 975.0 462.3 2. 2. X X 221.9 122.3 52107.1 65.2 30112XXXX X X X 71.7 X 68.1 121.2 71.4 68.1 63.0 30553XXXXXXX X X X X X X 545.3 63.0 687. X X 75.4 56.8 44102.9 113.6 64.4 61.6 3063.0 63.0 6076.0 76.0 68192XXX X X 119.2 56.8 6076.0 76.0 6076.0 76.0 6076.0 76.0 40101.0 64.0 00140XXX X X 124.0 70.0 4076.0 64.0 57176XXX X X 117.6 83.2 55.7 83.2 07126XXXXXXX X X X X X X 122.6 60.7 309. X 97.3 59.9 63.0 59.9 30101.0 63.0 31167.6 73.1 60220.0 76.0 30101.0 63.0 X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X Tarablus/ Tarablus/ Tarablus/ Batroun Batroun Batroun X X X X X X X X X X X X X X X X X X X X X X X X O Grab ROV O Grab ROV Grab ROV X X X X X X X X X X X X X X X X X X X X X X 73

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Schizomavella 63.6 63.6 X X (Schizomavella) cf. mamillata Schizomavella sp. 56.0 69.5 X XXX Schizoporella sp. 74.5 74.5 XX Schizoretepora sp. 76.0 76.0 X X Scrupocellaria cf. scrupea 76.0 76.0 X X Scrupocellaria sp. 101.0 101.0 X X Smittina cervicornis 63.6 67.2 X X Smittina sp. 68.1 120.4 X X X

Tarablus/ Min. depth Max. depth Beirut Jounieh Sayniq St. George ROV Grab Batroun CHAETOGNATHA cf. Sagitta sp. 784.6 784.6 X X

Tarablus/ Min. depth Max. depth Beirut Jounieh Sayniq St. George ROV Grab Batroun NEMERTEA Phoronis australis 979.0 979.0 XX

Tarablus/ Min. depth Max. depth Beirut Jounieh Sayniq St. George ROV Grab Batroun ECHINODERMATA Antedon mediterranea 59.9 107.1 X X X X X X cf. Antedon mediterranea 133.6 133.6 X X Asteroidea indet. (buried) 84.4 84.4 X Astropecten sp. 327.6 327.6 X X Brissopsis lyrifera 76.0 517.0 X X Centrostephanus longispinus 56.0 94.2 X X X X X X X Ceramaster granularis 73.4 148.5 XX Chaetaster longipes 70.6 70.6 X X Cidaris cidaris 62.7 949.7 X X X X X X X Coscinasterias tenuispina 67.5 98.0 X X XX Echinaster (Echinaster) 49.2 133.6 X XXXX sepositus cf. Echinaster (Echinaster) 58.8 139.5 X X sepositus Echinocardium sp. 64.0 64.0 X X Echinocyamus pusillus 101.0 101.0 X X Echinodermata indet. 45.6 979.2 X X X X X X X Echinoidea indet. 65.2 133.6 X X X X Genocidaris maculata 76.0 76.0 X X Gracilechinus acutus 520.3 834.7 X X Gracilechinus cf. elegans 755.7 755.7 X X Hacelia attenuata 65.2 221.9 X X XXX Hacelia sp. 121.2 121.2 X X Hymenodiscus coronata Hacelia superba Mesothuria intestinalis Luidia ciliaris Luidia sp.* Leptasterias sp. Peltaster placenta Ophiuroidea indet. cf. Ophiothrix sp. Ophiothrix fragilis Ophiopsila aranea Mesothuria sp. cf. Ophiopsila aranea Penilpidia ludwigi Stylocidaris affinis Tethyaster subinermis cf. Stylocidaris sp. CRUSTACEA ARTHROPODA/ Amphibalanus amphitrite Anomura indet. Amphipoda indet. Aristeus antennatus Bathynectes maravigna Balanus sp. Decapoda indet. Dardanus arrosor Cirripedia indet. Brachyura indet. Copepoda indet. cf. Bathynectes sp. cf. Bathynectes maravigna Ebalia deshayesi Ebalia granulosa uaaotaaidt 0. 0. X 345.9 606.3 345.9 606.3 Galathea squamifera Euphausiacea indet. Eumalacostraca indet. Ethusa mascarone Homola barbata Macropipus tuberculatus Liocarcinus depurator Inachus sp. i.dphMx et ertJuihSyi St. George Sayniq Jounieh Beirut Max. depth Min. depth 1. 930.3 260.4 617.8 260.4 4. 7. X X X 979.2 544.2 827.7 827.7 7. 177.7 177.7 780.7 650.5 9. ,0. X 1,005.4 791.7 9. 6. X 367.3 431.4 194.0 431.4 0. 2. X X 827.9 302.0 6. 6. X X X 766.1 465.2 8. ,1. X X X X X X X 1,012.4 212.0 289.7 845.7 212.0 845.7 674.4 521.1 674.4 521.1 9. 193.2 193.2 0. 300.1 300.1 3. 230.0 230.0 979. X 71.2 99.8 71.2 69.7 69291X X 269.1 60.2 86.9 60.2 307. X X X X X 77.1 85.8 63.0 59.9 56.0 59.9 61148.5 86.1 2082.0 82.0 05838XXXXXX X X X X X X 833.8 939.2 60.5 63.0 6865.8 56.8 73715X 75.6 781.5 75.6 77.3 5695.6 95.6 4074.0 74.0 009. X 90.0 90.0 607. X X X 76.2 73.7 76.2 56.0 X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X Tarablus/ Batroun X X X X X X X X X X X X O Grab ROV X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X 75

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cf. Macropipus tuberculatus 180.0 180.0 X X Munida sp. 82.1 158.0 XXXX Munidopsis marionis 551.2 800.7 X X X Munidopsis sp. 448.3 937.1 X XXX Mysidacea indet. 291.1 979.5 X X X X X X cf. indet. 691.3 695.2 X X Neomaja goltziana 418.8 752.8 X XXX cf. Pagurus alatus 850.4 850.4 X X Pagurus alatus 284.3 979.8 XXX X 86.6 113.9 X X cf. Pagurus sp. 396.8 396.8 X X Parapenaeus longirostris 286.9 799.6 X X X X X X cf. Parapenaeus longirostris 305.1 562.2 X X X Periclimenes sp. 59.9 604.4 X XX Plesionika antigai 354.9 373.2 X X Plesionika edwardsii 256.2 618.7 X X X X X X Plesionika gigliolii 318.9 598.5 X X X X X X Plesionika heterocarpus 227.8 535.2 X X X X X X cf. Plesionika heterocarpus 366.7 366.7 X X Plesionika narval 119.5 300.1 X X XXX Plesionika sp. 239.3 599.9 XXX X Scyllarus arctus 287.1 287.1 XX Eusergestes arcticus 372.1 502.3 X X Spinolambrus macrochelos 230.0 526.2 XXX X Stenopus spinosus 287.1 287.1 XX

Tarablus/ Min. depth Max. depth Beirut Jounieh Sayniq St. George ROV Grab Batroun MOLLUSCA alba 74.0 906.0 XX X Abra longicallus 74.0 1015.0 X X X X X X Abra nitida 64.0 101.0 X X Abra prismatica 74.0 76.0 XX X Abra sp. 74.0 74.0 XX X Acanthocardia paucicostata 82.0 82.0 X X Aequipecten opercularis 64.0 101.0 X X X X X Anadara corbuloides 74.0 74.0 X X Anadara gibbosa 74.0 74.0 X X Anadara cf. inaequivalvis + 74.0 74.0 X X Anadara cf. transversa + 74.0 74.0 X X Anomia ephippium 90.0 175.2 X XX Antalis dentalis 74.0 1015.0 XXX X Antalis inaequicostata 64.0 101.0 X X X pespelecani 74.0 74.0 X X noae Aporrhais serresianus Argonauta argo Asperarca nodulosa fusca Astarte sp. Astarte sulcata barbata Basisulcata lepida cf. Berthella sp. Berthella cf. stellata Bittium sp. rugosa Bolinus brandaris indet. cf. riisei Cardiidae indet. Calliostoma sp. Cardiomya costellata Centrocardita aculeata Cassidae indet. ehlsie ne.300300.0 300.0 Cephalaspidea indet. Cerithidea indet. Cephalopoda indet. Cerithidium submammillatum Cerithiopsidae indet. sp. +6. 64.0 64.0 cf. nodulosum+ Corbula gibba 101.0 Copulabyssia corrugata Coccopigya viminensis 101.0 Clio pyramidata corallinus Circenita callipyga+ cf. Creseis virgula Creseis sp. Ctena decussata Cuspidaria cuspidata Cuspidaria rostrata eaoiomsidt 9. 1,018.6 799.5 Decapodiformes indet. indet. Crassopleura maravignae 3. 634.2 634.2 6. 460.4 460.4 1. 111.1 113.6 111.1 113.6 4. 444.6 444.6 0. 4. X 647.2 403.6 8. 688.0 688.0 688.0 688.0 0. 906.0 906.0 0. 101.0 101.0 0. 906.0 906.0 14570X 74.0 517.0 74.0 91.4 40101.0 64.0 4074.0 74.0 6076.0 76.0 4076.0 74.0 4076.0 64.0 30265.0 63.0 4074.0 74.0 64171XXX X X 107.1 66.4 40220XXX X X 76.0 212.0 906 76.0 74.0 72.8 40173.0 906.0 74.0 76.0 4076.0 74.0 127. X 101.0 71.2 63.0 71.2 30906.0 63.0 4064.0 64.0 3076.0 63.0 4074.0 74.0 30542XXXX X X 101.0 X 64.0 544.2 63.0 6076.0 76.0 6076.0 76.0 40220.0 74.0 4074.0 74.0 6 668 668 X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X 77

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Digitaria digitaria 63.0 63.0 X X Dosinia lupinus 906.0 906.0 X X cf. Dosinia lupinus 76.0 76.0 X X Emarginula huzardi 63.0 76.0 X X X Ennucula corbuloides 1015.0 1015.0 X X Ennucula tenuis 74.0 74.0 X X Entalina tetragona 911.0 911.0 X X Euspira catena 74.0 76.0 XX X Flabellina sp. 138.6 138.6 X X Flexopecten hyalinus 76.0 82.0 X X Fulvia fragilis + 74.0 76.0 XX X Fusinus sp. 66.4 66.4 X X Galeodea sp. 531.8 976.7 X XX indet. 56.0 833.8 X X X XXX Glans trapezia 76.0 76.0 X X Goodallia triangularis 74.0 74.0 X X Hadriania craticulata 76.0 76.0 X X Haedropleura flexicosta 76.0 76.0 X X Haminoeidae indet. 906.0 906.0 X X Homalopoma sanguineum 63.0 63.0 X X Japonactaeon pusillus 76.0 76.0 X X Jujubinus sp. 552.1 552.1 XX Karnekampia sulcata 76.0 76.0 X X Kurtziela serga 906.0 906.0 X X Lepetellidae indet. 76.0 76.0 X X cf. Licornia vieirai 101.0 101.0 X X Lima lima 76.0 76.0 X X Limaria tuberculata 74.0 74.0 X X Limatula gwyni 74.0 74.0 X X cf. Loligo vulgaris 646.3 646.3 X X Loligo vulgaris 90.8 90.8 X X Macomangunus tenuis 76.0 76.0 X X Mactra olorina + 101.0 101.0 X X Mactra sp. 76.0 76.0 X X Malleus sp. 76.0 76.0 X X Melanella sp. 64.0 64.0 X X Metaxia metaxa 63.0 63.0 X X Mimachlamys varia 74.0 906.0 XX X Mitra sp. 101.0 101.0 X X Mitrella coccinea 76.0 76.0 X X Mitrella minor 906.0 906.0 X X Murex forskoehlii + 70.0 70.0 X XX indet. 289.9 289.9 XX Nassarius sp. 64.0 906.0 XXXX X Naticarius stercusmuscarum indet. Nucula hanleyi Neopycnodonte cochlear Nucula nitidosa Nucula nucleus Nucula sp. Nucula sulcata Nuculidae indet. Onychoteuthis banksii# Ommastrephes sp. Ocinebrina sp. Nudibranchia indet. Palliolum sp. Pagodula echinata Ommastrephes bartramii Octopus vulgaris Papillicardium papillosum Parvicardium exiguum Parvicardium Parvicardium minimum Parvicardium Parvicardium scabrum Parvicardium sp. Parvicardium Pecten jacobaeus Pectinidae indet. Petalopoma elisabettae Pinna rudis Pyramidellidae indet. Pteroctopus tetracirrhus Pseudamussium peslutrae Pododesmus patelliformis Roxania sp. Retusidae indet. Saccella commutata cf. Rossia macrosoma Rossia macrosoma Rissoidae indet. Ringicula sp. Sepia orbignyana Sepia officinalis Semisaccis granulata Scaphopoda indet. Sepiidae indet. Siliquariidae indet. Similipecten similis 7. 533.8 279.3 0. 300.0 X 300.0 820.1 877.2 820.1 877.2 6. 260.4 161.0 260.4 161.0 0. 906.0 396.8 906.0 666.8 396.8 666.8 906.0 906.0 906.0 906.0 7. 358.3 172.7 170.4 170.4 2. 1. X X 811.1 521.1 4074.0 74.0 57183XXXX X X X 74.0 178.3 74.0 55.7 401,015.0 74.0 401,015.0 74.0 2082.0 82.0 40906.0 64.0 4074.0 74.0 3063.0 82.7 63.0 82.7 55380XXX 64.0 X 64.0 X 358.0 65.5 30101.0 63.0 4076.0 74.0 4076.0 64.0 4074.0 74.0 4074.0 74.0 4074.0 74.0 4076.0 74.0 6076.0 76.0 8861.3 58.8 6076.0 X 76.0 192.9 90.0 401015 64.0 83.5 75.1 789. X X X X 91.4 X 67.8 879.4 76.0 4074.0 74.0 4076.0 74.0 X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X 79

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Spondylus gussonii 756.5 756.5 XX cf. Spondylus gussoni 477.0 477.0 X X Striarca lactea 74.0 76.0 XX X Taranis moerchii 668.0 668.0 X X Tellinidae indet. 465.7 465.7 X X cf. Tenagodus sp. 76.0 76.0 X X Teuthida indet. 458.7 488.2 X X Thracia convexa 76.0 76.0 X X Thyasira cf. subovata 101.0 101.0 X X Thyasira flexuosa 668.0 668.0 X X Thyasira sp. 64.0 64.0 X X Thylaeodus semisurrectus 63.0 63.0 X X Timoclea ovata 64.0 161.0 XX X X Tonna galea 239.3 239.3 X X Tritia cf. heynemanni 302.0 302.0 X X Trochidae indet. 674.9 701.7 X X Turritella turbona 63.0 76.0 XXX X Tylodina perversa 76.0 76.0 X X Veneridae indet. 76.0 76.0 X X Venus verrucosa 101.0 906.0 X X X Vermetidae indet. 59.9 74.0 X X X XX Volvarina mitrella 76.0 76.0 X X

Tarablus/ Min. depth Max. depth Beirut Jounieh Sayniq St. George ROV Grab Batroun CHORDATA Anguilliformes indet. 794.8 794.8 X X Anthias anthias 70.6 293.6 X X X X X X Argentina sphyraena 304.6 581.3 X X X X X X Arnoglossus laterna 179.7 216.5 X X Arnoglossus rueppelii 226.1 366.7 X X X X X X Arnoglossus sp. 458.7 458.7 X X Aulopus filamentosus 193.5 193.5 X X 780.7 931.4 X X X XX Benthocometes robustus 531.8 531.8 X X Callanthias ruber 132.7 221.9 XXX Capros aper 238.5 418.5 X X X Carangidae indet. 82.4 82.4 X X Chauliodus sloani 590.3 892.4 XXX X cf. Cheilodipterus X novemstriatus + Chelidonichthys cuculus 87.2 87.2 X X Chelidonichthys lastoviza 78.5 180.0 X X XX Chimaera monstrosa 881.5 881.5 X X Chlorophthalmus agassizi Coelorinchus caelorhincus Citharus linguatula Coris julis Coelorinchus sp. Conger conger Dentex macrophthalmus Dalophis imberbis Cottoidei indet. cf. mediterraneus Coryphaenoides Electrona risso Dipturus oxyrinchus Epigonus constanciae +5. 56.8 56.8 Fistularia commersonii+ Facciolella oxyrhyncha +7. 12X Gobius geniporus Gobius gasteveni 71.2 Gobiidae indet. 71.2 Etmopterus spinax Equulites klunzingeri+ Epinephelus marginatus Helicolenus dactylopterus Gobius vittatus Gobius sp. Gobius niger Hoplostethus mediterraneus Hymenocephalus italicus s.451491.0 415.1 Hymenocephalus sp. Lampanyctus crocodilus Lappanella fasciata Lepidorhombus whiffiagonis Lepidorhombus boscii Lepidopus caudatus Lesueurigobius sp. Lesueurigobius friesii Lepidotrigla cavillone Lophius budegassa Lobotes surinamensis# Lesueurigobius suerii Macroramphosus scolopax Mullus surmuletus Merluccius merluccius indet. Muraena helena 0471014.7 1014.7 8. 8. X X X X X X 887.0 187.6 6. 8. X X X X X X 788.0 267.7 6. 164.5 930 164.5 418.8 5. 6. X X X X X X 460.7 869.6 151.3 471.9 869.6 471.9 5. 359.7 464.4 255.4 421.8 464.4 421.8 9. 9. X X 799.3 952.8 799.3 952.8 7. 4. X X X X X X 842.6 X 177.7 110.0 110.0 6. 9. X X X X X X 991.6 363.1 5. 7. X X X X X X 979.5 358.8 7. 3. X X 833.0 477.2 2. 7. X X 174.9 124.0 8. 2. X X X X 424.7 X 582.7 285.2 747.5 582.7 432.5 7. 474.2 474.2 0. 3. X X X X X X 332.7 107.1 7. 173.0 388.4 173.0 925.8 294.2 363.1 81233XX X 223.3 68.1 6182.4 46.1 12180XX 71.7 X 82.7 58.8 128.0 82.7 71.2 82.1 82.1 717. X X 77.3 81.9 77.1 61.9 26303XXX X X 320.9 X 340.3 60.7 114.2 72.6 89.4 00219X 241.9 90.0 6446.4 46.4 X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X 81

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Mycteroperca rubra 81.8 81.8 X X Myctophidae indet. 420.2 881.5 XXXXX Nemichthys scolopaceus 672.1 672.1 XX Nemipterus randalli + 68.6 100.7 X X Nettastoma melanurum 487.4 1045.3 X X X X X X Nettastomatidae indet. 682.0 881.5 X X X X Nezumia aequalis 536.6 1,005.4 X X X XX Nezumia sclerorhynchus 536.3 898.0 X X X X Nezumia sp. 538.0 948.0 X X X XX Ophisurus serpens 192.1 192.1 X X Ostorhinchus fasciatus + 65.5 89.4 X X X Pagellus bogaraveo 177.7 504.8 XXXX Pagellus erythrinus 69.2 92.0 X X X Pagellus sp. 65.5 65.5 X X Parablennius pilicornis 48.4 48.4 X X Parablennius rouxi 81.0 81.8 X X Peristedion cataphractum 334.1 375.2 X X X Phycis blennoides 645.1 942.7 XXXXX Phycis phycis 148.5 192.9 X XX Pisces indet. 63.3 979.0 X X X X X X Pleuronectiformes indet. 80.1 375.2 X X X X Pterois miles + 49.5 85.5 X XX Sargocentron rubrum + 36.0 47.0 XXX Scorpaena elongata 127.7 441.8 X X X X X X Scorpaena notata 61.9 344.8 X X X X Scorpaena porcus 49.5 119.8 X XX Scorpaena scrofa 62.4 419.0 X X XXX Scorpaena sp. 80.7 88.0 X X X Scorpaeniformes indet. 435.3 435.3 X X Seriola dumerili 62.7 133.6 XX XX indet. 113.6 113.6 X X Serranus cabrilla 46.1 171.5 X X X X X X Serranus hepatus 61.6 178.3 X X X X X X Serranus scriba 61.6 61.6 X X Serranus sp. 90.0 90.0 X X Siganus luridus + 36.0 49.5 XXX indet. 198.3 198.3 X X Stephanolepis diaspros + 63.6 63.6 X X Stomias boa 399.3 861.4 XXXXX Stomiidae indet. 448.3 448.3 X X Stomiiforme indet. 696.0 696.0 X X Symphurus ligulatus 778.7 849.9 X X X X Symphurus nigrescens 122.6 226.1 X X Symphurus sp. 408.1 408.1 X X Synchiropus phaeton 304.6 474.6 X X X X X X CHORDATA/TUNICATA posbaci ne.6. 20X 72.0 69.2 indet. Aplidium sp. Syngnathus acus conchilega Ascidia cf. mentula Styelinae indet. Thorogobius ephippiatus Tunicata indet. Styela sp. Styela cf. canopus Rhopalaea sp. Polysyncraton lacazei Synodus synodus Synodus saurus Cystodytes dellechiajei Cystodytes sp. sp. Botrylloides indet. cf. Botryllinae cf. Ascidiidae indet. cf. aspersa indet. Ascidia sp. Rhopalaea cf. hartmeyeri Pyuridae indet. Pyura dura cf. Pycnoclavella sp. Pycnoclavella sp. Pycnoclavella cf. communis cf. Polyzoinae indet. cf. Polysyncraton sp. X 110.2 63.0 Torquigener flavimaculatus+ Didemnum sp. Didemnum fulgens violacea Tursiops truncatus# Tripterygion melanurum Halocynthia papillosa cf. cyrnusense Pseudodistoma Microcosmus cf. vulgaris 38 38 i.dphMx et ertJuihSyi St. George Sayniq Jounieh Beirut Max. depth Min. depth 5. 157.8 157.8 1. 1. X 119.2 119.2 X 266.9 266.9 0. 0. X 107.1 107.1 056. X 60.5 60.5 5271.7 65.2 65.5 65.5 029. X X 93.4 60.2 09121X 102.1 70.9 6084.1 56.0 X X 341.4 94.2 6466.4 X X 66.4 107.0 78.5 78.0 68.1 71.7 71.7 X 69.2 69.2 758. X 82.0 X 57.7 X 39.1 X 67.5 X 63.0 56.0 X 83.5 38.8 X X 63.0 119.2 78.5 551.2 95.6 69.2 56.0 77.1 306. X 64.1 X 53.2 X 71.4 63.0 71.0 38.8 81.8 39.1 83.5 69.0 81.8 83.5 32161XXXXXX X X X X X X X X X X X 116.1 X 83.5 53.2 126.3 70.6 80.1 0070.0 70.0 06131XXXXXX X X X X X 163.1 70.6 926. X 69.2 69.2 X X X X X X X X X X X X X X X X X X Tarablus/ Batroun X X X X X X X X X X X X X X X X O Grab ROV X X X X X X X X X X X X X X X X X X X X X X X X X X X 83

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TABLE A2. LIST OF THREATENED SPECIES IDENTIFIED

Species are listed that are either included on Annex II or III of the SPA/BD Protocol of the Barcelona Convention, and/or have been assessed by IUCN as threatened (i.e., Vulnerable (VU), Endangered (EN), or Critically Endangered (CR)) or Near Threatened (NT) in the Mediterranean Sea.

Listed on Listed on IUCN Red List Phylum Species Annex II (SPA/ Annex III (SPA/ Threat Status in Reference BD Protocol) BD Protocol) the Mediterranean Arthropoda/Crustacea Scyllarus arctus X 43 Chordata Chimaera monstrosa NT 26 Chordata Dipturus oxyrhincus NT 26 Chordata Epinephelus marginatus X EN 26, 43 Chordata Merluccius merluccius VU 26 Chordata Tursiops truncatus X VU 26, 41 Cnidaria Dendrophyllia ramea X VU 26, 53 Cnidaria Desmophyllum dianthus X EN 26, 53 Cnidaria Funiculina quadrangularis VU 26, 53 Cnidaria Pennatula rubra VU 26, 53 Echinodermata Centrostephanus longispinus X 41 Mollusca Pinna rudis X 41 Mollusca Tonna galea X 41 Porifera Sarcotragus foetidus X 41 Porifera Spongia (Spongia) officinalis X 43

FIGURE A1. NUMBER OF IDENTIFIED TAXA, ACCORDING TO BROAD TAXONOMIC GROUP

200

180

160

140

120

100

80

60 Number Identi ed of Taxa

40

20

0

Annelida Bryozoa Cnidaria Plantae Porifera Chordata MolluscaNemertea Phoronida Unknown Brachiopoda ChaetognataChlorophyta Foraminifera Ochrophyta Rhodophyta Echinodermata Chordata/Tunicata Arthropoda/Crustacea Arthropoda/Chelicerata

Taxonomic Group Table 1), asfollows: Spatially of surveyareas Methodologyforassesingrelative conservationinterest Annex 2. for thekeycommunities,pr Annex 2 4) 1) 3) 2)

yielding amaximumvalueof9points.(See Figure 41c). Thesetwoscores 1 point). were 1 individual: multiplied, 2 points; <10 individuals: >1 and abundance, according to the same categories shown in Table 1 (≥10 individuals: 3 points; other speciesofgeneralscientificinterest (1 point). Theywere alsoscored fortheir relative species recorded from Lebanon(3 points), rare speciesfor theMediterranean(2 points), or added together, yieldingthefinalindexofconservationinterest, showninMap 15. Each ofthethr Other speciesofparticularinter (See Figure 41b). >1 and <10 individuals: 2 points; 1 individual: 1 point). The maximum value was 18 points. abundance, according to the same categories shown in Table 1 (≥10 individuals: 3 points; andmultipliedbyathirdvalue ofupto6 points), score, whichwasbasedontheirrelative Thesetwoscores were addedtogether(fora 2 points). orAnnex III: 3 points; (Annex II: oftheSPA/BD orAnnex III or listedunderAnnex II Protocol oftheBarcelona Convention and/ NearThreatened: 1 point) Vulnerable: 2 points; Red Listed(Endangered: 3 points; Pr (See Figure 41a). if notincluded).Thesetwoscores were multiplied,yieldingamaximumvalueof6 points. other calcareousbio ofthecoralligenousand the MediterraneanSeaorActionPlanforconservation underwater caves andcanyons,aphotichardbeds and chemo ofhabitatsandspeciesassociatedwithseamounts, Action Planfortheconservation were alsoscored according towhethertheywere includedintheUNEP-MAPRAC/SPA They and low:1 point). medium: 2 points; categories shown inTable 1 (high: 3 points; Key communitieswer ‑ otected and/orthreatened specieswere alsoscored according towhethertheywere referenced datafrom theunderwatersurveys,ataresolution of500mxm,were combined ee indiceswasrescaled tofallbetween 0 and 1. The three indiceswere then ‑ otected and/orthreatened species,andparticularspeciesofinterest (see (2 points if included; 1 point ifincluded;1 point concretions intheMediterraneanSea(2 points e scored fortheirrelative abundance,according tothesame est were scored according towhethertheywere new ‑ synthetic phenomena in © OCEANA/EnriqueTalledo 85

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Annex 3 References edj, G.,Laubier, L.1985.ThedeepMediterraneanbenthos.In:Moraitou

24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 1 9 8 7 6 5 4 3 2

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