Marine Research Findings of the VECTORS Project

This website provides access to the research results of the VECTORS project, which can be used to support marine management decisions, policies and governance as well as future research and investment. VECTORS was a large scale project that brought together more than 200 expert researchers from 16 different countries. It examined the significant changes taking place in European seas, their causes, and the impacts they will have on society.


All of the publications created as part of the VECTORS project are listed below (latest first).
Please use the search box below to find a publication by title, author(s), year or keyword; the most relevant results will be displayed at the top of the page.

  • Galil, B.S. (2013). Going going gone: the loss of a reef building gastropod (Mollusca: Caenogastropoda: Vermetidae) in the southeast Mediterranean Sea. Zoology in the Middle East 59(2), 179-182. doi:10.1080/09397140.2013.810885
    View abstract The gregarous vermetid gastropods modify their environment, both ecologically and physically (Chemello & Silenzi, 2011). Yet the existence of the vermetid protected platforms which mitigate and hinder wave-induced coastal erosion depends on a delicate balance between opposing forces: vermetid accretion deposition and cementation on one hand, bioerosion and marine erosion on the other. Mediterranean vermetid reefs depend for the existence on the actively-built rims constructed by gastropods of the genus Dendropoma Morch, 1861. As early as 1961 Safriel (1966) observed that along the central coast of Israel "although the raised margins of the platforms and terraces are completely coated by Dendropoma shells, the majority of these are empty, the animals having been probably killed by overgrowth of a dense algal population". Living individual s are confined to patches clear of seaweeds, mainly found around burrows inhabited by the crab Pachygrapsus which feeds on the surrounding algae. The majority of living patches are on the exposed rims of the platforms.� With fewer individuals than are need to counterbalance marine and bio-erosion, we may fact decimation of this particular endemic seascape.
  • Galil, B.S., P. Genovesi, H. Ojaveer, G. Quílez-Badia and A. Occhipinti. (2013). Mislabeled: eco-labeling an invasive alien shellfish fishery. Biological Invasions 15(6). doi:10.1007/s10530-013-0460-9
    View abstract The invasive alien Manila clam fishery from Ria Arousa, Spain, was recommended for a Marine Stewardship Council certificate which implies the fishery maintains the integrity of the ecosystem. By certifying an environmentally harmful invasive alien clam fishery, the MSC harms the environment and risks its credibility, at the time when environmental responsibility and social responsibility are increasingly important to policymakers, industry, and consumers, particularly in Europe. We call on the MSC to reassess their evaluation of the Ruditapes philippinarum fishery in Ria Arousa, Spain.
  • Galil, B.S. and R. Gevili. (2013). A moveable feast: Beroe cucumis sensu Mayer, 1912 (Ctenophora; Beroida; Beroidae) preying on Mnemiopsis leidyi A. Agassiz, 1865 (Ctenophora; Lobata; Bolinopsidae) off the Mediterranean coast of Israel. BioInvasions Records 2(3), 191-194. doi:10.3391/bir.2013.2.3.03
    View abstract In the winter months of 2012 and 2013 aggregations of the native comb jelly Beroe cucumis were observed and photographed along the Israeli coast preying on the invasive American comb jelly Mnemiopsis leidyi. It is suggested that native beroid may take part in controlling populations of the invasive ctenophore.
  • Hufnagl, M., M.A. Peck, R.D.M. Nash, T. Pohlmann and A.D. Rijnsdorp. (2013). Changes in potential North Sea spawning grounds of plaice (Pleuronectes platessa L.) based on early life stage connectivity to nursery habitats. Journal of Sea Research 84, 26-39. doi:
    View abstract We explored the hypothesis that spawning ground locations of North Sea plaice reflect the locations of nursery grounds using drift scenarios based on a baroclinic, shallow-water circulation model (HAMSOM). The transport of pelagic eggs and larvae was simulated each year from 1975 to 2006 using in situ forcing, temperature-dependent development and stage-specific behaviour of eggs and larvae. This long-term simulation period also allowed us to explore climate effects. A release position was considered a potential and suitable spawning site if larvae from that area reached coastal nurseries after the onset of metamorphosis. In general, larvae were transported in an anti-clockwise direction and settled in nurseries that were relatively close to the release positions. Spawning locations that were offshore were poorly connected to nursery grounds while those closer to the shore had higher connectivity. Simulated suitable spawning locations broadly agreed with the main centres of egg production (English Channel, Southern Bight, German Bight), except for the known spawning grounds south of Dogger Bank. Over the 31-year simulation period, positive and negative trends in transport success were found for the western and eastern parts of the North Sea, respectively. Changes in the west (Flamborough Head) were mainly due to changes in water circulation patterns whereas those in the east (northern German Bight) were induced by changes in both currents and water temperature. The implications of these findings, and the significant correlation between changes in drift and recruitment, suggest that climate-driven changes in the suitability of nursery grounds will directly affect the distribution and productivity of plaice in the North Sea.
  • Hooper, T.L. and M.C. Austen. (2013). Tidal barrages in the UK: Ecological and social impacts, potential mitigation, and tools to support barrage planning. Renewable and Sustainable Energy Reviews 23, 289-298. doi:
    View abstract The UK Government is committed to ambitious targets to reduce carbon emissions. The large tidal ranges in estuaries on the west coast of the UK make the deployment of tidal barrages an attractive proposition, and repeated feasibility studies have been undertaken. No barrage scheme has yet been taken forward, and one factor contributing to this reluctance to proceed is the significant environmental impacts that could result from the barrage construction and operation. This paper provides a detailed review of the current understanding of the potential ecological and social impacts of tidal barrages, including a case study of La Rance in northern France, and a discussion of strategies for mitigating barrage impacts. The review considers how more comprehensive ecological modelling could reduce uncertainty in predicting the impacts in specific estuaries, and discusses the use of Multi-criteria Analysis and ecosystem valuation as tools for evaluating the disparate costs and benefits of barrages schemes.
  • Gravili, C., C.G. Di Camillo, S. Piraino and F. Boero. (2013). Hydrozoan species richness in the Mediterranean Sea: past and present. Marine Ecology 34(Suppl. 1), 41-62.
    View abstract The Mediterranean hydrozoan fauna (Siphonophora excluded) comprises 400 species; most (68%) occur in the Atlantic Ocean, 20% are endemic to the Mediterranean, 8% are of Indo-Pacific origin, and 4% are non-classifiable. There are 69 nonindigenous (NIS) species in the basin: 44% of these are casual (recorded just one or very few times), 28% established (widely recorded in the basin), 6% invasive (established NIS that are able rapidly or largely to disseminate away from the area of initial introduction, having a noticeable impact on the recipient community), and 22% questionable (of doubtful taxonomic status). Entry through the Suez Canal and range expansion through the Gibraltar Strait, often enhanced by ship traffic, appear to be the main processes for recent species introductions, but uncertainties remain for many NIS. Species additions immediately result in larger local or regional species pools, but the newcomers might impact on populations of native species, altering extinction probabilities. A more reliable evaluation of the species pool can be accomplished by adding new species when they enter the taxonomic record (i.e. the records of any taxon in all types of literature), and by removing species that have not been found for a "reasonable" time (e.g. several decades). Of the 400 non-siphonophoran hydrozoan species known to occur in the Mediterranean Sea, positive records in the last 10 years are available for 156 species (39%), whereas records of the remaining 244 species are older than a decade: 67 species have not been recorded for 41 years, 13 for 31-40 years, 79 for 21-30 years, and 85 for 11-20 years.
  • Galil, B.S., B. Kumar and A. Riyas. (2013). Marivagia stellata Galil and Gershwin, 2010 (Scyphozoa: Rhizostomeae: Cepheidae), found off the coast of Kerala, India. BioInvasions Records 2(4), 317-318. doi:10.3391/bir.2013.2.4.09
    View abstract A specimen of the cepheid scyphozoan Marivagia stellata Galil and Gershwin, 2010, described from the Mediterranean coast of Israel, is here reported from Kerala, on the southwestern coast of India. The present record establishes M. stellata as the fourth scyphozoan species introduced to the Mediterranean through the Suez Canal.
  • Goren, M., N. Stern and B.S. Galil. (2013). Bridging the gap: first record of Mertens' prawn-goby Vanderhorstia mertensi in Israel. Marine Biodiversity Records 6, e63. doi:10.1017/s1755267213000419
    View abstract The Indo-Pacific fish species Vanderhorstia mertensi was found, for the first time, off the Israeli coast in the Mediterranean. This shrimp-associated goby was reported so far only from the Mediterranean coast of Turkey. Its presence verifies the status of V. mertensi as an Erythraean alien.
  • Ferrario, J., D. Savini, A. Lodola, A. Marchini and A. Occhipinti Ambrogi. (2013). Risk of non-indigenous species introduction via international seafood trade: the case of Chioggia fish market. Biologia Marina Mediterranea 20(1), 236-237.
    View abstract Fish and shellfish trade and aquaculture have been increasing over the last few years due to the rising demand for live and processed products. In Italy, the most important area of production and exchange is the Adriatic Sea (, particularly the Lagoon of Venice. The global movement of organisms of economic interest is an important pathway of non-indigenous species (NIS) introduction into aquatic ecosystems by means of intentional release or unintentional dispersal of imported organisms-target species as well as non-target species associated with packaging (Weigle et al., 2005; Minchin, 2007). Within the EU-FP7 Project VECTORS (Vectors of Change in Ocean and Sea Marine Life, Impact on Economic Sectors) a preliminary investigation on species traded at the fish market of Chioggia, the most important fisherman locality near Venice, was carried out, with the aim of exploring the potential role of seafood trade in NIS introduction.
  • Ghabooli, S., T.A. Shiganova, E. Briski, S. Piraino, V. Fuentes, D. Thibault-Botha, D.L. Angel, M.E. Cristescu and H.J. MacIsaac. (2013). Invasion pathway of the ctenophore Mnemiopsis leidyi in the Mediterranean Sea. Plos One 8(11), e81067. doi:10.1371/journal.pone.0081067
    View abstract Gelatinous zooplankton outbreaks have increased globally owing to a number of human-mediated factors, including food web alterations and species introductions. The invasive ctenophore Mnemiopsis leidyi entered the Black Sea in the early 1980s. The invasion was followed by the Azov, Caspian, Baltic and North Seas, and, most recently, the Mediterranean Sea. Previous studies identified two distinct invasion pathways of M. leidyi from its native range in the western Atlantic Ocean to Eurasia. However, the source of newly established populations in the Mediterranean Sea remains unclear. Here we build upon our previous study and investigate sequence variation in both mitochondrial (Cytochrome c Oxidase subunit I) and nuclear (Internal Transcribed Spacer) markers in M. leidyi, encompassing five native and 11 introduced populations, including four from the Mediterranean Sea. Extant genetic diversity in Mediterranean populations (n=8, Na=10) preclude the occurrence of a severe genetic bottleneck or founder effects in the initial colonizing population. Our mitochondrial and nuclear marker surveys revealed two possible pathways of introduction into Mediterranean Sea. In total, 17 haplotypes and 18 alleles were recovered from all surveyed populations. Haplotype and allelic diversity of Mediterranean populations were comparable to populations from which they were likely drawn. The distribution of genetic diversity and pattern of genetic differentiation suggest initial colonization of the Mediterranean from the Black-Azov Seas (pairwise FST=0.001-0.028). However, some haplotypes and alleles from the Mediterranean Sea were not detected from the well-sampled Black Sea, although they were found in Gulf of Mexico populations that were also genetically similar to those in the Mediterranean Sea (pairwise FST=0.010-0.032), raising the possibility of multiple invasion sources. Multiple introductions from a combination of Black Sea and native region sources could be facilitated by intense local and transcontinental shipping activity, respectively.
  • González-Duarte, M.M., C. Megina, S. Piraino and J.L. Cervera. (2013). Hydroid assemblages acress the Atlantic-Mediterranean boundary: is the Strait of Gibraltar a marine ecotone? Marine Ecology 34(Suppl. 1), 33-40.
    View abstract Strong gradients in physico-chemical properties between abutting water masses create prominent transition zones in the marine environment. The Strait of Gibraltar forms the well defined boundary between the Mediterranean and Atlantic, and this paper examines spatial variation of hydroid assemblages in this transition zone. Although several studies highlighted the transitional character of the Strait and defined it as an ecotone, the benthic hydroid assemblages did not show differences between the Gulf of Cadiz and the Alboran Sea. However, there is an asymmetrical influence of the Atlantic waters on the coastal benthic ecosystems of the Alboran Sea, which maintains a more Mediterranean character in the hydroid assemblages of the northern coast, whereas a more Atlantic character was found in the rest of the studied sites. The transition zone between Atlantic and Mediterranean benthic communities could be associated with an Atlantic Influence Zone rather than with the Strait of Gibraltar itself.
  • Daewel, U., S.S. Hjøllo, M. Huret, R. Ji, M. Maar, S. Hiiranen, M. Travers-Trolet, M.A. Peck and K.E. van de Wolfshaar. (2013). Predation control of zooplankton dynamics: a review of observations and models. ICES Journal of Marine Science 71(2), 254-271. doi:10.1093/icesjms/fst125
    View abstract We performed a literature review to examine to what degree the zooplankton dynamics in different regional marine ecosystems across the Atlantic Ocean is driven by predation mortality and howthe latter is addressed in available modelling approaches. In general, we found that predation on zooplankton plays an important role in all the six considered ecosystems, but the impacts are differently strong and occur at different spatial and temporal scales. In ecosystems with extreme environmental conditions (e.g. low temperature, ice cover, large seasonal amplitudes) and low species diversity, the overall impact of top-down processes on zooplankton dynamics is stronger than for ecosystems having moderate environmental conditions and high species diversity. In those ecosystems, predation mortality was found to structure the zooplankton mainly on local spatial and seasonal time scales. Modelling methods used to parameterize zooplankton mortality range from simplified approaches with fixed mortality rates to complex coupled multispecies models. The applicability of a specific method depends on both the observed state of the ecosystem and the spatial and temporal scales considered. Modelling constraints such as parameter uncertainties and computational costs need to be balanced with the ecosystem-specific demand for a consistent, spatial-temporal dynamic implementation of predation mortality on the zooplankton compartment.
  • Coppa, S., G.A. de Lucia, P. Magni, P. Domenici, F. Antognarelli, A. Satta and A. Cucco. (2013). The effect of hydrodynamics on shell orientation and population density of Pinna nobilis in the Gulf of Oristano (Sardinia, Italy). Journal of Sea Research 76, 201-210. doi:
    View abstract Pinna nobilis is the largest endemic bivalve of the Mediterranean Sea, declared protected since 1992. Although hydrodynamic stress induced by waves is known to influence density, size and orientation of P. nobilis, the effect of other hydrological features is unknown. This paper considers a P. nobilis population living within a Posidonia oceanica meadow in the Gulf of Oristano (Sardinia, Italy). We hypothesize that spatial differences in density and orientation of P. nobilis may be related to significant wave height (HS), wave direction (DW), bottom current direction (DBC) and bottom current speed (SBC). A population of P. nobilis was investigated at different sites and its distribution was correlated to hydrodynamics by means of a numerical modeling approach. The spatial distribution was patchy, with a density of 0.06-6.7 ind.100 m-2. A non-uniform distribution of shell orientations (OS) was demonstrated in 4 sites out of 6. DBC and SBC were the main factors affecting OS, while waves had little influence. A SBC of 0.07 m s-1 appears to be the threshold for inducing specimen directionality with shells aligned to the current and the ventral side exposed to the flow. This suggests that feeding strategy is a key factor in determining OS, in addition to drag minimization. We also highlighted the role of adjacent lagoons in supporting high densities as a result of high food availability. These findings demonstrate the usefulness of modeling techniques in explaining the spatial distribution pattern of P. nobilis and in contributing to our knowledge of its ecological traits.
  • David, M., S. Gollasch and M. Pavliha. (2013). Global ballast water management and the "same location" concept: a clear term or a clear issue? Ecological Applications 23(2), 331-338. doi:10.1890/12-0992.1
    View abstract The United Nations recognized the transfer of harmful organisms and pathogens across natural barriers as one of the four greatest pressures to the world's oceans and seas, causing global environmental changes, while also posing a threat to human health, property, and resources. Ballast water transferred by vessels was recognized as a prominent vector of such species and was regulated by the International Convention for the Control and Management of Ship's Ballast Water and Sediments (2004). Permanent exceptions from ballast water management requirements may apply when the uptake and discharge of ballast water occur at the "same location." However, the "same location" concept may be interpreted differently, e.g., a port basin, a port, an anchorage, or a larger area even with more ports inside. Considering that the Convention is nearing the beginning of enforcement, national authorities all around the world will soon be exposed to applications for exceptions. Here we consider possible effects of different interpretations of the "same location" concept. We have considered different possible extensions of the same location through environmental, shipping, and legal aspects. The extension of such areas, and the inclusion of more ports, may compromise the Convention's main purpose. We recommend that "same location" mean the smallest practicable unit, i.e., the same harbor, mooring, or anchorage. An entire smaller port, possibly also including the anchorage, could be considered as same location. For larger ports with a gradient of environmental conditions, "same location" should mean a terminal or a port basin. We further recommend that IMO consider the preparation of a guidance document to include concepts, criteria, and processes outlining how to identify "same location", which limits should be clearly identified.
  • David, M., S. Gollasch and E. Leppakoski. (2013). Risk assessment for exemptions from ballast water management--the Baltic Sea case study. Marine Pollution Bulletin 75(1-2), 205-217. doi:10.1016/j.marpolbul.2013.07.031
    View abstract The International Convention for the Control and Management of Ship's Ballast Water and Sediments sets requirements to prevent organism transfers. Vessels on certain routes can be exempted from such requirements based on risk assessment (RA). As the convention nears its entry into force, the interest in exemptions increases. Such RA should be conducted according to the International Maritime Organization G7 Guidelines. We present a RA study for exemptions applied to intra-Baltic shipping considering different RA methods, i.e., environmental matching, species specific method including target species and species biogeographical aspects. As reliable species data in the ports considered are unavailable and following the precautionary principle, no exemptions should be granted. To ensure data reliability, port baseline surveys and regular monitoring programs should be undertaken during the exemption period as new species found influence the RA result. The RA model prepared is considered as of value to other areas worldwide.
  • Brachvogel, R., L. Meskendahl, J.-P. Herrmann and A. Temming. (2013). Functional responses of juvenile herring and sprat in relation to different prey types. Marine Biology 160, 465-478.
    View abstract The relationship between particulate-feeding rates and prey concentrations (functional response) of juvenile herring and sprat (5-9 cm total length) was investigated in controlled feeding experiments monitored by an underwater camera system. A special tank system was developed allowing the regulation and quantification of low prey concentrations (1-160 L-1). Non-evasive Artemia nauplii was used as prey to estimate the maximum biting rate of both predators. In contrast, Acartia tonsa with a high escape ability was used as a realistic prey type. Herring and sprat showed a type II functional response for both prey types. Nonlinear mixed effects model revealed no significant difference between the functional responses of both predators, except that herring showed significantly higher biting rates than sprat at A. tonsa concentrations below *40 L-1. For both predators feeding rates were significantly higher with Artemia nauplii than with A. tonsa. Video analysis indicated that sprat, unlike herring, is an obligate particulate-feeder.
  • Bulleri, F. (2013). Grazing by sea urchins at the margins of barren patches on Mediterranean rocky reefs. Marine Biology 160(9), 2493-2501. doi:10.1007/s00227-013-2244-2
    View abstract The role played by the urchins, Paracentrotus lividus and Arbacia lixula, in the formation and persistence of barren areas dominated by encrusting coralline macroalgae is yet to be fully elucidated. This study, carried out in the NW Mediterranean (43A degrees 30'N, 10A degrees 20'E) between February 2005 and April 2006, investigated how the loss or density decrease in one or both urchin species influences the recovery of erect macroalgal stands (dominated by filamentous forms) at the margins of barren areas. At a depth of 4-6 m, three barren patches were assigned to each of the following treatments: (1) control (natural densities of A. lixula and P. lividus); (2) 50 % of the natural density of A. lixula and natural density of P. lividus; (3) total removal of A. lixula and natural density of P. lividus; (4) 50 % of the natural density of P. lividus and natural density of A. lixula; (5) total removal of P. lividus and natural density of A. lixula; (6) 50 % of the natural densities of both A. lixula and P. lividus; (7) total removal of both A. lixula and P. lividus. The effects of the herbivore treatments were evaluated either in the presence or the absence of encrusting corallines. The partial or total removal of A. lixula, P. lividus or both favored the proliferation of filamentous macroalgae at the margins of barren patches. The presence of encrusting corallines reduced the development of these macroalgae. The results of this study suggest that a moderate decrease in the density of just one of the two species can decrease the ability of the herbivore assemblage to control the proliferation of filamentous macroalgae at the margins of barren patches. The extent of barren areas appears, therefore, to be regulated by the outcome of density-dependent interactions between the two species of sea urchins.
  • Boero, F., G. Belmonte, R. Bracale, S. Fraschetti, S. Piraino and S. Zampardi. (2013). A salp bloom (Tunicata, Thaliacea) along the Apulian coast and in the Otranto Channel between March-May 2013. F1000Research 2(181). doi:10.12688/f1000research.2-181.v1
    View abstract Between March-May 2013 a massive Salpa maxima bloom was recorded by a citizen science study along the Ionian and Adriatic coast of the Salento peninsula (Italy). Citizen records were substantiated with field inspections along the coast and during an oceanographic campaign in the Otranto Channel. Salps clogged nets, impairing fishing activities along the coast. Swimmers were scared by the gelatinous appearance of the salps, and thought they were jellyfish. At the end of the bloom the dead bodies of the colonies, that were up to 6-7 m long, were accumulated along the coast and stirred by the waves, forming foams along dozens of kilometers of coast. The bloom also occurred at the Tremiti Islands, north of the Gargano Peninsula. The possible impacts of such events on the functioning of pelagic systems are discussed.
  • Bogi, C. and B.S. Galil. (2013). Cylichna villersii, an Erythraean cephalaspideid snail (Mollusca: Gastropoda: Opisthobranchia) in the eastern Mediterranean. Marine Biodiversity Records 6, e92. doi:10.1017/s1755267213000687
    View abstract Cylichna villersii, a minute 'bubble snail' is newly recorded in the Mediterranean Sea from specimens recently collected along the Israeli coastline. The species, described and illustrated, matches the syntype collected in the Red Sea by Savigny. It is the eighth Erythraean alien cephalaspideid species recorded in the Levantine Basin. The recent collection of many living specimens in several continuously sampled locations attests to the speed of its establishment in the south-eastern Levant.
  • Bogi, C. and B.S. Galil. (2013). Monotygma watsoni, an Erythraean alien pyramidellid (Mollusca: Gastropoda: Heterobranchia) newly recorded in the Mediterranean Sea. Marine Biodiversity Records 6, e80. doi:10.1017/s175526721300047x
    View abstract Monotygma watsoni, a minute parasitic pyramidellid gastropod is newly recorded in the Mediterranean Sea from specimens recently collected off the Israeli coastline. The species, described and illustrated, matches the three syntypes of Ceratia watsoni collected in the southern Red Sea. The characteristic protoconch and teleoconch sculpture led us to assign it to Monotygma. It is the fourteenth Erythraean alien pyramidellid species recorded in the Levantine Basin. The large populations of a great number of Erythraean aliens in the Levantine Basin may serve as reservoir hosts for pyramidellids, many of which seem to be parasitic generalists, and may be introduced to native Mediterranean hosts.
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