Content uploaded by Stephan Gollasch
Author content
All content in this area was uploaded by Stephan Gollasch on Jun 21, 2017
Content may be subject to copyright.
Life in ballast tanks 217
LIFE IN BALLAST TANKS
STEPHAN GOLLASCH1*, ELSPETH MACDONALD2,
SARA BELSON3, HELGE BOTNEN4, JENS T. CHRISTENSEN5,
JOHN P. HAMER6, GUY HOUVENAGHEL7, ANDERS JELMERT8,
IAN LUCAS6, DANIEL MASSON9, TRACY MCCOLLIN10,
SERGEJ OLENIN11, AGNETA PERSSON12, INGER WALLENTINUS12
LAMBERTUS P. M. J. WETSTEYN13 & THOMAS WITTLING14
1GoConsult, Hamburg, Germany
2Food Standards Agency, Aberdeen, Scotland
3Maritime Research Centre, Southampton Institute, Southampton, UK
4UNIFOB, Section of Applied Environmental Research, High Technology
Centre, Norway
5Dept. of Marine Ecology, Institute of Biological Sciences, University of
Aarhus, Aarhus C, Denmark
6University of Wales, Bangor, School of Ocean Sciences, Menai Bridge,
Anglesey, UK
7Université Libre de Bruxelles, Brussels, Belgium
8Institute of Marine Research, Austevoll Aquaculture Research Station,
Storebø, Norway
9IFREMER, Station de La Trembalde, France
10FRS Marine Laboratory, Aberdeen, United Kingdom
11Coastal Research and Planning Institute, Klaipeda University,
Lithuania
12Department of Marine Botany, Göteborg University, Göteborg, Sweden
13National Institute for Coastal and Marine Management, Middelburg,
The Netherlands
14Institut für Hydrobiologie und Fischereiwissenschaft, Universität
Hamburg, Hamburg, Germany
*Corresponding author SGollasch@aol.com
Abstract
The abundance and diversity of species in ballast water, a recognised vector for the accidental
introduction of nonindigenous organisms, has been examined through many studies around the
world over the last 25 years. The results of European research activities are summarised in this
contribution by outlining the objectives of some of these studies, and by focusing on the diversity
of taxa determined from ballast water and tank sediment samples. In total 1508 samples (1219
ballast water, 289 tank sediment) were collected on 550 ships. A total of 990 taxa were identified
during the 14 European shipping studies. The diversity of species found included bacteria, fungi,
protozoans, algae, invertebrates of different life stages including resting stages, and fishes with a
body length up to 15 cm. Crustacean, molluscan and polychaete invertebrates and algae form the
majority of species found.
1 Introduction
The introduction of nonindigenous organisms to new areas has resulted in populations
of many species being established outside their native ranges, with, in some cases, po-
S. Gollasch et al.
218
tentially deleterious effects on native species, the receiving habitats and their economies
(Hedgpeth 1993). Whilst some species are introduced deliberately to new areas for
aquaculture purposes, shipping is often regarded as one of the main vectors for the acci-
dental transfer of aquatic organisms. Organisms may be transferred by ships as hull
fouling and also inside ballast tanks. Since the introduction of steel hulled vessels in the
late 19th century, water is the preferred ballast material used by ships to stabilise and
trim the vessel and to submerge the propeller when ships are not fully loaded. The
global nature of the shipping industry makes it inevitable that many ships must load
ballast water in one area and discharge it in another. It is most likely that the modern
shipping industry, with faster ships transporting large quantities of ballast between geo-
graphically separate areas, has an increasing potential to transport nonindigenous spe-
cies to new areas. It has been estimated that major cargo vessels annually transport
nearly 10 billion tonnes of ballast water world-wide, indicating the global dimension of
the problem. On average, 3,000 – 4,000 species (Carlton and Geller 1993; Gollasch
1996) have been estimated to be transported between continents by ships each day.
Organisms discharged with ballast water in coastal waters and ports of call beyond their
native range may have the potential to impact native species and ecosystem functions,
fishing and aquaculture industries, as well as public health. However, the introduced
organism must first survive the ballasting and subsequent deballasting processes in
addition to the period in transit inside the ballast tanks (Carlton 1985). Conditions upon
discharge must also be suitable for survival, and the organism must then find sufficient
food and resources, suitable habitat, and in some cases, other individuals with which it
can reproduce before a population can become established (Gollasch 1996). The inocu-
lation stage is simply one step in the complex process of invasion biology (Carlton
1985).
The first account attributing the introduction of an aquatic organism to ballast water
transport was published by Ostenfeld (1908) after a mass occurrence of the Asian phy-
toplankton algae Odontella (Biddulphia) sinensis in the North Sea in 1903. The first
documented studies to sample ships' ballast water were carried out 70 years later by
Medcof (1975) followed by e.g. Hallegraeff and Bolch (1991), Locke et al. 1991, Lucas
et al. (1999). This contribution summarizes the methods used and the diversity of taxa
transported in ballast water and tank sediments sampled in completed and ongoing
European shipping studies.
2 Material and methods
The European research considered here consists of practical studies in which ballast
water and, in some cases, sediments, were sampled from ships' ballast tanks (Table 1).
Ballast water samples were taken using a variety of nets, hoses and pumps operated via
tank openings (manholes), pumps operated via sounding pipes or air vents connecting
the ballast tanks to the ships´ surface and by extracting water at the ships´ ballast pump
(in line sampling). The net haul depth varied according to tank design from 1.5 m to >
15 m. Sterile samples were collected by immersing sterile 1L bottles from opened man-
holes and by collecting sediment in sterile cut-off syringes (Table 2). Ballast water
sampling methods vary greatly world-wide, and the particular method used will often
depend upon the main taxa that particular researchers are interested in, and the practical
and logistical requirements of the ships to be sampled. This variety in methods has been
Life in ballast tanks 219
addressed by a European research initiative entitled “Concerted Action Introductions
with Ships” and is reported elsewhere (Rosenthal et al. 2000).
Table 1. Number of ships sampled, number of samples and duration of shipping studies
carried out in European countries.
Shipping study Number of ships
sampled
Number of sam-
ples taken
Ballast Tank
water sediment
Duration of
study
Type of
sampling
Belgium 5 26 6 1995-1998 in ports
en-route
Denmark 1 8 ongoing, start
2000
en-route
England 20 49 1998 – 2000 in ports
England &
Wales
112 114 102 1996 – 1999 in ports
France 0 0 0 ongoing, start
2000
in ports
Germany 1 189 131 71 1992 – 1996 in ports
Germany 2 9 1 12 1998 – 1999 in ports
Lithuania 11 22 1999 – 2000 in ports
Netherlands 17 21 2 ongoing, start
1999
in ports
Norway 1 45 1996 – 1997 in ports
Norway 2 6 8 4 1998 – 1999 in ports
Scotland 127 134 92 1994-1997 in ports
Sweden 3 1996 in ports
EUCA 5 705 1998 – 1999 en-route
Total 550 1219 289
Vessels sampled ranged from small cargo vessels of < 1,000 deadweight tonnes (dwt) to
very large crude carriers (VLCCs) of > 300,000 dwt. The ballast water sampled was
from more than 200 different origins world-wide, but predominately from the northern
hemisphere. The principal objective of all studies was to identify the variety of species
transported with ships ballast water.
In most case studies, ballast water samples were preserved by the addition of formalde-
hyde or glutaraldehyde or other fixative (e.g., Lugol´s iodine). In some studies unpre-
served samples were taken for culturing purposes and flow-cytometrical analysis. The
organisms were identified to the lowest taxonomic level possible using stereomicro-
scopes, microscopes, selective culture media, and in one case study, DNA-based tools
(Polymerase Chain Reaction followed by gene sequencing). Wherever possible, the
individuals were determined to species level. Due to unfavourable conditions (see be-
low) some individuals were not determined to the species level, but are included as
"cf.". In other cases where a species identification was not made, only the genus was
determined and this is noted by "sp.". Specimens that could not be ascribed to a genus
level were assigned to the lowest appropriate taxonomic unit (Family, Order etc.) with
the addition "indet.". Reasons why some organisms could not be identified to the spe-
cies level included the following. Some resting stages (phyto- and zooplankton) and
invertebrate larvae can only be determined to species level by rearing the specimens to
motile, juvenile or adult stages. As a majority of the samples were fixed, such cultiva-
S. Gollasch et al.
220
tion was not possible. Second, cultivation of unknown specimens is difficult, as the
optimal culture conditions are generally unknown, and in many cases can only be at-
tempted by trial and error. This is time consuming and costly, with no guarantee of
success. Third, it can be very difficult to culture all species in a sample considering that
many are present at low densities. Fourth, in some samples only one individual was
found. For some species (e.g. copepods) it is essential to investigate both sexes or adults
to confirm the identification. Fifth, organisms were frequently damaged or in poor con-
dition thus rendering difficult culturing the species. Sixth, it is very difficult to identify
specimens which may have originated from a wide variety of locations as taxonomic
keys might not adequately represent the range in variation of the species. Finally, for the
microbial community, in some cases the percentage of bacteria that could be cultured
was low compared to total numbers, and in some cases when molecular methods were
used, identification was hampered by the incomplete library of genomic sequences for
marine bacteria.
Bray-Curtis similarity analysis. Several shipping studies considered here sampled the
ballast water for zooplankton species. Results from these studies were selected for a
Bray-Curtis similarity analysis as e.g., surveys where the main focus was to collect
bacteria should not be compared with a survey focussing sediment dwelling species,
fish or zooplankton. Comparing studies that targeted different types of organisms would
likely result a very low similarity. Species found in sediment samples were omitted
unless they were found both in sediment and water samples.
A short description of each practical study follows below. While this list attempts to
summarise all the European studies, it should not be assumed to be fully comprehensive
(Table 1).
Belgium, Universite Libre de Bruxelles (total number of taxa identified: 28). The
investigation entitled "Study of the Potential Role of Transportation of Ships Ballast
Water on the Geographical Extension of Blooms of Toxic Algae" was carried out at the
Université Libre de Bruxelles. The main results of the study were that risks do exist
concerning the introduction of nonindigenous toxin-producing phytoplankton species
into European waters with ballast water or sediment discharges. They recommended the
implementation of ballast water management guidelines at an international level (Van-
den Boeck 1995).
Denmark, University of Aarhus, Institute of Biological Sciences, Dept. of Marine
Ecology (total number of taxa identified: 4). A joint project between Lithuania and
Denmark has recently started to regularly sample the Ro-Ro Ferry "Urd" (Scandlines),
running between Aarhus/Aabenraa (Denmark) and Klaipeda (Lithuania). The main
objective of the study is to assess the survival of planktonic organisms en-route. It is
planned to regularly sample the ship before departure in the port of Aarhus and after
arrival in the port of Klaipeda (and vice versa). Additionally, it is planned to carry out
port profiles of the Danish harbours Aarhus and Fredericia, and to sample ships that
arrive in these ports from outside the Baltic Sea (J. Christensen, University of Aarhus,
Denmark, pers. comm.).
Life in ballast tanks 221
Table 2. Ballast tank sampling methods used by each European shipping study.
Shipping
study
Methods: Ballast water samples Methods: Tank sediment samples
Belgium In line, sampling at ships’ ballast pump
Pump, used to collect water samples using a hose via
air vents
Trowel, scoop and bottle to collect
sediment from the bottom of emp-
tied and ventilated tanks
Denmark Water sampler, lowered down through tank opening. On
deck the sample volume was concentrated by using a 60
µm plankton net (zooplankton). Phytoplankton samples
were not concentrated.
England Bottle, repeatedly lowered down through tank opening to
below the water surface
In line, sampling at ships’ ballast pump
England &
Wales
Hose, 2.5 cm internal diameter, weight on the end,
lowered through manholes to collect integrated water
samples
Hand-hauled plankton net, 53 µm mesh 30 cm diameter,
75 cm length lowered through manholes
Pump, used to collect water samples using a hose via
manholes or sounding pipes.
In line, sampling at ships’ ballast pump
Pump, used to pump sediment
slurry using a hose via manholes or
sounding pipes
Trowel, scoop and bottle to collect
sediment from the bottom of emp-
tied and ventilated tanks
France Bottle, repeatedly lowered down through tank opening to
below the water surface
Germany 1 Hand-hauled plankton net, cone shaped opening, cone
diameter 9,7, net diameter 25 cm, length 95 cm, mesh
size 55µm (zooplankton) and 10µm (phytoplankton),
Pump, small hand pump operated via sounding pipes,
hose diameter 14 mm, flow rate 1l/min.
In line sampling at the ship´s ballast pump
Small trawl, width 30 cm, mesh
size 250 µm,
Trowel, scoop and bottle to collect
sediment from the bottom of emp-
tied and ventilated tanks
Germany 2 Hand-hauled plankton net (net diameter 25 cm, length
95 cm, mesh size 55µm )
Trowel, scoop and bottle to collect
sediment from the bottom of emp-
tied and ventilated tanks
Lithuania Hand-hauled plankton net, cone shaped opening cone
diameter 9,7, net diameter 25 cm, length 95 cm, mesh
size 55µm (zooplankton) and 10µm (phytoplankton)
Nether-
lands
Hand-hauled plankton net, net diameter 25 cm, length
50 cm, mesh size 20 µm (phytoplankton) and 55 µm
(zooplankton),
In line sampling at the ship´s ballast pump,
Pump, hand pump (Whale bilge pump) operated via
sounding pipes, inner hose diameter 13 mm (outer 19
mm), flow rate ca. 30 l/min.
Trowel, scoop, to collect sediment
from the bottom of emptied and
ventilated tanks
Norway 1 Hand-hauled plankton net, opening diameter 20 cm,
length 100 cm, mesh size 20-30 µm (zooplankton)
Ruttner water sampler, 1.5 l (phytoplankton)
Norway 2 Sterile bottle lowered down through manholes Sterile “decapitated” syringe col-
lecting sediment plug in sediment
at bottom of emptied and ventilated
tanks
Scotland Vertical hose (2.5 cm internal diameter through manhole
or 1.25 cm hose through sounding pipe) for phytoplank-
ton.
Hand-hauled plankton net (0.5 m diameter ring net, 68
µm mesh size) or by filtering a known volume of water
through a 68 µm mesh.
In line sampling at the ship´s ballast pump
Pumping sediment slurry from
bottom of tank by hand-operated
mono-pump or battery operated
bilge pump.
Sweden Pump, small hand pump operated via manholes
Bucket lowered down below the water surface through
manhole
Trowel, scoop and bottle to collect
sediment from the bottom of emp-
tied and ventilated tanks
EUCA Hand-hauled plankton net, cone shaped opening, cone
diameter 9.7, net diameter 25 cm, length 95 cm, mesh
size 55µm
S. Gollasch et al.
222
England, Maritime Research Centre, Southampton Institute, United Kingdom
(total number of taxa identified: 68). A research studentship assessed the transport of
phytoplankton in the ballast water of vessels using the Port of Southampton. The project
aimed to establish whether ballast water was a potential source of exotic phytoplankton
species to the Southampton Water estuary, and also whether the number of cells and
taxa could be correlated to factors including the area of origin, age and selected physical
and chemical properties of the water. All of the samples collected originated within the
northern hemisphere, and two-thirds were from European waters (S. Belson, South-
ampton Institute, United Kingdom, pers. comm.).
England & Wales, School of Ocean Science, University of Wales, Bangor (total
number of taxa identified: 252). Between 1996 and 1999, samples were taken from
ships arriving at English and Welsh ports. The aim of the study was to investigate the
range of organisms present in ships` ballast tanks and assess variation in ballast water
organisms in relation to geographic origin and season (McCollin et al. 1999). A further
objective was to assess the potential risk of the introduction of organisms in ballast
water and sediment to English and Welsh waters. Ships were sampled at 20 ports
throughout England and Wales. Some culturing was carried out to investigate the vi-
ability of organisms (Hamer et al. 1999). The majority of ballast water sampled origi-
nated from ports in the northern hemisphere (McCollin et al. 1999).
France, IFREMER, Station de La Tremblade (no taxonomic results available yet). A
study started recently in La Tremblade, involving sampling of ballast tanks to identify
the microbial component of species found (D. Masson, IFREMER, France, pers.
comm.).
Germany 1, Institute for Marine Research, Kiel, and University of Hamburg (total
number of taxa identified: 502). The first European shipping study was undertaken in
Germany (Gollasch 1996), the purpose of which was to conduct a thorough taxonomic
assessment of planktonic and benthic organisms found in ballast water tanks with addi-
tional samples taken from ship hulls (Lenz et al. 2000). The vessels investigated were
selected according to type of vessel (e.g., container ships and bulk carrier) and sea areas
covered by their voyages. The majority of samples originated from tropical and
warm-temperate regions (Gollasch 1996; Lenz et al. 2000).
Germany 2, Institute for Marine Research, Kiel, and University of Hamburg (total
number of taxa identified: 19). A follow up study was carried out from 1998-1999. The
key objective was to sample ballast tank sediments (T. Wittling, Universität Hamburg,
Germany, pers. comm.).
Lithuania, University of Klaipeda (total number of taxa identified: 90). A study
started recently in Klaipeda, involving sampling ballast water and examining hull foul-
ing on ships in dry docks (S. Olenin, University of Klaipeda, Lithuania, pers. comm.).
The Netherlands, National Institute for Coastal and Marine Management, Middel-
burg and AquaSense/Tripos, Amsterdam (total number of taxa identified: 88). Dur-
ing a pilot project in the years 1995 and 1996, in cooperation with the Smithsonian
Environmental Research Center (USA), phytoplankton and zooplankton samples were
taken in cargo holds of bulk-carriers travelling between the ports of Maryland and Rot-
Life in ballast tanks 223
terdam. One key objective was to assess the viability of organisms after an oceanic
voyage (Tripos 1997). In 1998 questionnaires were sent to incoming ships and literature
reviews were carried out to estimate the amount and origin of ballast water carried to
and from The Netherlands and to assess the risks of the introduction of nonindigenous
species into Dutch coastal waters (AquaSense 1998a, b). Since 1999 samples have been
collected on board ships arriving in the ports of Rotterdam, Vlissingen-Oost and Am-
sterdam. The objectives include documentation of the diversity of plankton organisms
and their survival potential in Dutch waters. Culture experiments were carried out at
10°C and 20°C and at salinities 5 PSU, 15 PSU and 32 PSU and also in port water and
in GF/F filtered port water. Finally, number, size and pigment distributions of particles
in live ballast water samples are analyzed by flow-cytometry; this method also provides
information on autotrophic viability (chl-a fluorescence) of plankton upon arrival as
function of voyage duration.
Norway 1, UNIFOB, Section of Applied Environmental Research, Bergen (total
number of taxa identified: 181). In 1996, a project entitled the Sture Project was
launched. The aim of the study was to investigate the potential of unintentional intro-
ductions of nonindigenous aquatic species to Sture via ships’ ballast. All ships sampled
arrived from ports outside Norway (Botnen et al. 2000).
Norway 2, IMR, Bergen and University of Bergen (total number of taxa identified:
3). Between July 1998 and April 1999, ballast water and sediment samples were col-
lected from 6 ships entering the harbour of Mongstad (one ship redirected to Sture) in
Norway to identify the microbial component of species in ballast tanks. The faecal bac-
terium E. coli was found in one vessel out of six, while the cholera bacterium Vibrio
cholerae was not detected in the two ships analysed for its presence (A. Jelmert, Insti-
tute of Marine Research, Norway, pers. comm.).
Scotland, FRS Marine Laboratory, Aberdeen (total number of taxa identified: 327).
Between May 1994 and December 1996, ballast water was sampled from vessels arriv-
ing in Scottish ports. The aim of the study was to investigate planktonic organisms in
ballast water and sediment, with special emphasis on harmful and potentially toxic
phytoplankton (Macdonald 1998; Macdonald and Davidson 1998).
Sweden, Department of Marine Botany, Göteborg University (total number of taxa
identified: 41). A pilot project entitled "Risks associated with introduction of nonindi-
genous organisms to Swedish waters by water/sediment in the ballast tanks of ships”
was carried out in 1996. The project also included a regional survey of dinoflagellates
(Persson 2000) along the province of Bohuslän on the Swedish west coast as a baseline
study. The study focused on phytoplankton and on culturing sampled raw material.
Concentrated (by filtration, 10µm) ballast water samples and untreated sediment sam-
ples were incubated in a culturing chamber (16°C, 80 µEm-2 • s-1, 12:12 h light:dark) in
enriched (with f/10 nutrients) deep sea water from the Swedish west coast. This treat-
ment was performed to compare initial findings in the ship samples with cultures estab-
lished from resting stages undetected in the untreated samples (Persson 2000; Persson et
al. 2000).
European Union Concerted Action (total number of taxa identified: 67). This recently
completed European wide Concerted Action (EUCA) included five Ocean Going Work-
S. Gollasch et al.
224
shops (OGW) that were undertaken to assess the survival of zooplankton organisms in
ballast water en-route. The OGW were undertaken in European waters and during inter-
oceanic voyages. In total, 705 samples were collected during more than 100 days at sea
(Gollasch et al. 2000b, Olenin et al. 2000, Rosenthal et al. 2000).
3 Results
During the 14 European ship sampling studies considered here, a total of 1508 samples
(1219 ballast water, 289 tank sediment) were collected on 550 ships (Table 1). The total
number of taxa identified during all completed shipping studies varied between 3 and
502 per study (Fig. 1) and the number of taxa identified overall was 990 (Table A1,
available at http://www.ku.lt/nemo/EuroAquaInvaders.htm). The diversity of species
found included e.g. bacteria, fungi, protozoans (summarized as "other taxa"), algae,
invertebrates of different life stages including resting stages, and fishes with a body
length up to 15 cm (Table A1). More species were identified in those studies which
sampled more vessels (Fig. 1).
The most frequently collected taxa identified by eight shipping studies were diatoms,
harpacticoid copepods, rotifers (but all were not identified to species level) and the
diatom Skeletonema costatum (Table A1). The second most frequent identified taxa in
seven studies were Ditylum brightwellii, Chaetoceros sp., Navicula sp., Thalassionema
nitzschioides, Thalassiosira sp. (Bacillariophyceae (Diatoms)), and Temora longicornis
(Copepoda, Calanoida), calanoid copepods, larvae of Gastropoda, Bivalvia and Poly-
chaeta (Table A1).
3.1 FLORA
Most of the 497 taxa found in the sampled ballast tanks were determined to species
level (66%) (Fig. 2, Table A1)). In total, taxa of 13 higher taxonomic groups (i.e., phyla
and orders) were identified. Diatoms (250 taxa) and dinoflagellates (126 taxa) were the
predominant groups found. Taxa of all other 11 higher taxonomic groups were found in
frequencies less than 70 (Table 3). Rarely found, with five or fewer records were the
taxonomic groups Cryptophyceae, Prymnesiophyceae, Xanthophyceae, Ulotrichales,
Rhodophyta and filamentous green alga. Resting stages of Xanthophycaea (1 taxa) and
dinoflagellates (64 taxa) were found rarely (Table A1).
3.2 FAUNA
Fifty-five percent of the 425 taxa were determined to species level (Fig. 2, Table A1). In
total, taxa of 18 higher taxonomic groups were determined. Crustacea (240 taxa) and
Mollusca (73 taxa) were dominant faunal members of the ballast tank community. Taxa
of all other 16 higher taxonomic groups were found in frequencies less than 20. Rarely
found, with five or fewer records were the taxonomic groups Ctenophora, Turbellaria,
Nemertea, Oligochaeta, Pantopoda, Insecta, Arachnida, Chaetognatha and Echinoder-
mata (Table 4).
In the most commonly encountered group (Crustacea), Calanoida (54 taxa), Harpacti-
coida (33 taxa) and Cladocera (28 taxa) were dominant (Table 4). In the second fre-
quently identified group (Mollusca), the Bivalvia clearly prevailed with 42 taxa, fol-
Life in ballast tanks 225
lowed by Gastropoda with 30 taxa. Resting stages of seven cladoceran taxa were found
(Table A1).
The similarity between studies is low (10% -33%), which indicates that there are only
few commonly found species within the surveys. The similarity between e.g., the Scot-
tish and English survey is about 25%. The samples from the Lithuanian and German
study 1 have the highest similarity with 33% indicating the high number of species
commonly collected during both studies. The English, Scottish, Dutch and German 2
surveys form one group which has low similarity (20% -25%), but show even a lower
similarity to the other studies (10%) (Fig. 3).
Figure 1. Number of identified taxa according to number of sampled ships (all shipping
studies).
3.3 OTHER TAXA (BACTERIA, FUNGI AND PROTOZOANS)
Most of the 68 taxa found were not determined to species level (22.1%) (Fig. 2, Table
A1). In total taxa of ten higher taxonomic groups were determined. Foraminifera (19
taxa) and Ciliata (16 taxa) were dominant in this category. Taxa of all other eight higher
taxonomic groups were found in frequencies less than ten. Rarely found or less fre-
quently studied, with five or fewer records, were the taxonomic groups Bacteria, Radi-
oloaria, Heliozoa and Fungi (Table 5). Resting stages of one protozoan taxon (Ciliata)
were found (Table A1).
4 Discussion
The purpose of this paper is to document the diversity of taxa transported in ballast
tanks sampled in European shipping studies. The enormous amount of data generated by
these studies is too vast to be considered in great detail here. It was not attempted to
identify native and nonindigenous species, but rather to look at the diversity of species
0
100
200
300
400
500
600
0 50 100 150 200
Number of sampled ships
Number of identified taxa
S. Gollasch et al.
226
(b)
187 236
2
0
100
200
300
400
Number of taxa
(c)
53 15 0
0
100
200
300
400
Higher taxa,
indet.
Species Species cf.
(a)
142
328
27
0
100
200
300
400
transported in ships ballast tanks. Within Europe, it is often not clear for many species
whether they are indeed native or not (i.e., they are cryptogenic (Carlton 1996)), par-
ticularly for the smaller organisms (e.g., bacteria, algae and protozoa) as they are gener-
ally not so well studied or described.
Figure 2. Species, including species with unconfirmed identification (species cf.) of the
flora (a) (total number of taxa 497), fauna (b) (total number of taxa 425), and bacteria,
protozoa and fungi, summarised as "other taxa" (c) identified during all European ship-
ping studies. Specimens that could not be ascribed to a genus level, were assigned to
the lowest appropriate taxonomic unit (Family, Order etc.) summarised here as "higher
taxa" with the addition "indet.".
In total, the 990 species found during the shipping studies considered here belong to a
highly diverse range of taxonomic groups including bacteria, protozoa, phytoplankton,
zooplankton and zoobenthos. The largest specimens found inside a ballast tank were
Petromyzon marina (Cyclostomata, Pisces) of 15 cm length. However, the total number
of taxa identified may be assumed to be an underestimate. One reason is that some
studies targeted specific taxonomic groups. Furthermore, individuals listed as "sp. "
might not be the same in different studies, due to the range of taxonomic expertise
available in different institutes. It is likely that more species are hidden in the category
"indet." although some taxa may also be the same as others have identified to genus or
species level. Even more species are probably not identified because many taxonomists
have their specialities within certain taxonomic groups. It is also most likely that many
more organisms occurred in the sediments which were often not possible to sample and
Life in ballast tanks 227
it also is more difficult to find microscopic organisms in the sediments than in the wa-
ter.
Table 3. Flora: Number of taxa identified during European shipping studies according to
higher taxonomic unit.
Taxa Number %
Cyanophyceae 24 4.8
Chrysophyceae 6 1.2
Dictyochophycaea 6 1.2
Prymnesiophyceae 5 1.0
Bacillariophyceae (Diatoms) 250 50.3
Xanthophyceae 2 0.4
Dinophyceae (Dinoflagellates) 126 25.4
Cryptophyceae 5 1.0
Prasinophyceae 6 1.2
Chlorophyceae 63 12.7
Ulotrichales 1 0.2
Rhodophyta 1 0.2
Filamentous Clorophyta 2 0.4
Total 497 100.0
Sampling ships is different from sampling the environment, especially considering that
sampling access is not always straightforward, and is highly variable between ships.
One needs to deal with several obstacles, such as cargo overlaying the manhole pre-
venting direct access to the ballast water, and support frames installed to stabilise the
ballast tanks that make it difficult to access to the very bottom of the ballast tank. Sam-
pling equipment can also become stuck inside the tanks (Sutton et al. 1998; Rosenthal et
al. 2000).
Due to the problems encountered when sampling ballast water (Sutton et al. 1998;
Rosenthal et al. 2000), results can be regarded as a minimum qualitative and quantita-
tive assessment of the biological content of ballast water. This is apparent when consid-
ering the increase in identified species when ballast water already sampled with nets is
additionally sampled with traps or the samples cultured in the laboratory. Further, some
shipping studies focus ballast water and do not sample the ballast tank sediments or vice
versa. By selecting the type of samples (i.e., ballast water or tank sediments) some spe-
cies present in the ballast tanks were not collected. The application of a wider range of
sampling methods would possibly increase the range of species found in ballast water,
but most shipping studies are limited by practical constraints and time available for
sampling. The choice of the methods will further depend upon a number of operational
procedures, on ship design, location of and access to the ballast tanks and also the
amount of water in these tanks. The objectives of sampling (e.g. qualitative or quantita-
tive samples, target organisms or all taxa) are other criteria for method selection. The
preferred access to ballast tanks for sampling is via opened manholes. This would nor-
mally result in the use of short nets which are more easily manipulated and can be oper-
ated in ballast tanks which often restrict the depth of sampling tows.
Cone-shaped nets may be regarded as a suitable way of easily and efficiently sampling a
ballast tank. The main reason for the high sampling efficiency is the particular net con-
figuration that increases the filtration rate by limiting the overflow of water from the net
S. Gollasch et al.
228
caused by mesh resistance. It is recommended that the cod end of a net should be made
of a cup with filtration panels on its side and a tap at the base of the cup. If the cod end
is metallic no additional weighting is required to sink the net and this will reduce the
risk of entanglement in structures in the ships ballast tanks. For phytoplankton sampling
nets, it is recommended that relatively small mesh-sizes (e.g., 10 µm) be used. Larger
mesh sizes will exclude smaller species and may result in lower species richness esti-
mates however, fine mesh nets may clog quickly if organisms are very abundant, so a
degree of compromise may be required. In zooplankton studies, nets with mesh size of
55 µm will capture the youngest stages of Mollusca and Crustacea as well as many of
the other taxonomic groups commonly found in ballast water.
Table 4. Fauna: Number of taxa identified during European shipping studies according to
higher taxonomic unit. Crustacea broken down in taxonomic groups.
Taxa Number %
Cnidaria 11 2.6
Ctenophora 2 0.5
Tentaculata 7 1.6
Turbellaria 2 0.5
Rotatoria 16 3.8
Nemertea 1 0.2
Nematoda 16 3.8
Mollusca 73 17.2
Polychaeta 16 3.8
Oligochaeta 3 0.7
Pantopoda 1 0.2
Crustacea 240 56.5
Cladocera 28
Ostracoda 12
Copepoda 1
Calanoida 54
Cyclopoida 21
Harpacticoida 33
Poecilostomatoida 16
Siphonostomatoida 1
Cirripedia 1
Balanomorpha 21
Lepadomorpha 1
Mysidacea 11
Isopoda 6
Amphipoda 14
Cumacea 1
Euphausiacea 3
Decapoda 15
Insecta 4 0,9
Arachnida 1 0.2
Chaetognatha 5 1.2
Echinodermata 2 0.5
Tunicata 6 1.4
Pisces 19 4.5
Total 425 100.0
Cone-shaped nets were the most effective in terms of number of specimens and number
of taxa collected and therefore such nets are recommended in future ballast water sur-
veys. However, this net will not capture all taxa and its exclusive use would lead to an
Life in ballast tanks 229
underestimate of species diversity. The use of pumps, operated via sounding pipes,
exhibited similar effectiveness to the cone-shaped net. However, some pumps are un-
able to lift water from more than 8 meters depth, consequently ballast tanks with low
water levels or in deep location within the ship are unlikely to be sampled at all
(Rosenthal et al. 2000).
Figure 3. Bray-Curtis similarity analysis based on studies that collected fauna in ballast
water. Species found in sediment samples are omitted unless they were found both in
sediment and water samples (Abbreviations see Table A1).
Very few taxa were found in more than four studies and the high number of taxa found
during one or two studies alone may indicate the unique character of different shipping
studies (i.e. according to the inclusion of both sediment and ballast water samples).
However, it should be taken into account that some studies have sampled more than 100
ships and some less than ten, and this is likely to have had an impact on the range of
taxa found. Eventually this may result in more species being identified in those studies
BRAY-CURTIS SIMILARITY
100
90
80
70
60
50
40
30
20
10 En
g
l
Sco
N
l
Ger 2
Eng&Wa
Swe
N
or 1
Lit
Ger 1
EUCA
Be
Dk
S. Gollasch et al.
230
which sampled more vessels, as more samples will have been examined. Furthermore,
the diversity of ballast water origins and the seasonal spread of the times when the bal-
last was loaded contributes to the specific character of each shipping study. Addition-
ally, some shipping studies focused on a range of target groups from both ballast water
and sediment, whilst others were more restricted in their scope. The apparent differ-
ences in taxa encountered when employing different sampling methods is the major
reason for not including quantitative data on the various taxa in this review, and the
comparison of the studies concentrated on qualitative results.
4.1 CONCLUSIONS AND RECOMMENDATIONS
While a wide range of taxa have been identified from preserved ballast tank samples,
this gives no firm conclusions whether specimens were alive, healthy, in poor condition
or dead at the time of the investigation. However, vital stains can be used to identify
organisms that were alive at the time of collection, even if samples cannot be immedi-
ately examined in the laboratory. However, it is recommended that while fixed samples
give valuable results, it would also add to our knowledge to examine living material and
to undertake culturing experiments in order to document taxa that grow in cultures, as
was done in some of the studies considered here. This approach also has limitations,
because if taxa do not develop in cultures, it is not always known whether this was due
to the poor condition of the organisms at the time of culturing, or due to inappropriate
culture conditions.
It is recommended that several sampling methods should be employed to sample the full
range of organisms in ballast tanks.
Table 5. Bacteria, protozoans and fungi, summarised as "other taxa" identified during all
European shipping studies according to higher taxonomic unit.
Taxa Number %
Bacteria 5 7.4
Euglenophyceae 6 8.8
Zoomastigophora 8 11.8
Rhizopoda 9 13.2
Radiolaria 1 1.5
Foraminifera 19 27.9
Heliozoa 1 1.5
Ciliata 16 23.5
Fungi 1 1.5
incertae sedis taxa 2 2.9
Total 68 100.0
The diverse range of nonindigenous species introduced to many regions around the
world - by a range of vectors - and the diversity of organisms transported with ships´
ballast, illustrates the need to develop treatment options in order to minimise the risks of
species introductions in ballast. While retention of ballast water on board for prolonged
periods may reduce the risks of ships transporting viable aquatic organisms between
different areas, it is unlikely that it will completely eliminate these risks. Effective, safe,
economically viable and environmentally sound treatment methods and/or management
options for ballast water are necessary to minimise the risks associated with ballast
water releases. The presence of organisms in ballast tanks that may potentially affect
Life in ballast tanks 231
public health is also of particular concern, and is further justification for research efforts
into effective treatment of ballast water to minimise the unwanted transport of these
organisms on a global scale.
This study has demonstrated that virtually all forms of life are found in ballast water,
and concerted efforts must be made to reduce further invasion.
Acknowledgements
The authors gratefully acknowledge the cooperation of the shipping industry in all these
studies. Funding was provided by: Maritime Research Centre of Southampton Institute,
UK; Ministry of Agriculture, Fisheries and Food, UK; German Environment Protection
Agency; North Sea Directorate of the Ministry of Transport, Public Works and Water
Management, The Netherlands; Norsk Hydro, Norway; Research Council of Norway
(132453/230); Russian State Hydrometeorological University (St. Petersburg); Scottish
Office Agriculture, Environment and Fisheries Department, UK Marine Safety Agency
and Scottish Natural Heritage; Swedish Environmental Protection Agency and WWF
Sweden; European Union, MAST-Programme and UNESCO Intergovernmental
Oceanographic Commission (IOC). We are grateful to all student researchers and col-
leagues that assisted within these studies, who are too numerous to mention individu-
ally. We express our thanks to more than 200 taxonomists world-wide for the support in
species identification.
Tab. A2
LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP
Bacteria
Escherichia coli v B, S 1
Cytophagales gs B 1
a-protecteobacteria gs B 1
filamentous, indet. v B, S 1
solitary, indet. v B, S 1
incertae sedis taxa
Anchrymonas sp. v B, S 1
Metromonas sp. v* B 1
Euglenophyceae, indet. vB x B v*B 3
Anisonema sp. v B 1
Cercomonas sp. v B 1
Euglena sp. v B v B v B x B 4
Phacus sp. vB 1
Rhynchobodo sp. v B 1
Kinetoplastida
Bodo sp. v B v* B, S 2
Rhyncomonas sp. v* B, S 1
Choanoflagellata, indet. vB 1
Codonosiga sp. v B 1
Coccolithophorales, indet. v B 1
Silicoflagellates, indet. v B 1
Zoomastigophora
Rhynchomonas nasuta vB 1
Telonema subtilis vB 1
Rhizopoda
Hartmanella sp. v B 1
Labyrinthula sp. v B 1
Mayorella sp. v B 1
Nuclearia sp. v B 1
Paulinella ovalis vB 1
Platyamoeba sp. v B 1
Vahlkampfia sp. v B 1
Vannella arabica vB 1
Vannella sp. v B 1
Radiolaria, indet. vB 1
Foraminifera, indet. v B x B v B v B v B 5
Ammonia beccarii vS 1
Ammonia sp. v B 1
Brizalina variabilis vS 1
Discorbis sp. v S 1
Elphidium articulatum vS 1
Elphidium crispum vS 1
Elphidium excavatum vS 1
Elphidium sp. v B,S 1
Globigerinidae, indet. v S 1
Globorotalidae, indet. v S 1
Globorotalia tumida vS 1
Nonion sp. v S 1
Planorbulina mediterranensis vS 1
Protelphidium sp. v B,S 1
Rosalina bradyi vS 1
Rosalina sp. v B,S 1
Rotalia sp. v S 1
Spiroloculina communis vS 1
Heliozoa
Acanthocystis sp. v B 1
Ciliata, indet. v B v B v B x B v B, S 5
Aspidisca sp. v B 1
Colpidium sp. v, r B 1
Condylostoma magnum vB 1
Diophrys sp. v B 1
Enchelys sp. v B 1
Euplotes sp. v B v B, S 2
Holosticha sp. v B v B, S 2
Keronopsis sp. v B 1
Oxytricha sp. v B 1
Pleuronema sp. v B 1
Strombidium sp. vB 1
Tintinnidae xB vBvB 3
Uronema sp. v B v B 2
Urostyla sp. v B 1
Vorticella sp. v B v B 2
Fungi vB 1
Flora
Cyanophyceae, indet vB vB 2
Anabaena lemmermannii vB 1
Anabaena scheremetievii vB 1
Anabaena spiroides vB 1
Anabaena sp. vB v B 2
Aphanizomenon flos-aquae vB 1
Coelosphaerium kuetzingianum vB 1
Coelosphaerium minutissimum vB 1
Chroococcus minutus vB 1
Chroococcus sp. vB 1
Total
findings
Eng&WaBe Dk Ger 1 Ger 2 Lit Nl Nor 1 Nor 2 Sco Swe EUCA*
Shipping study
Taxon Engl
LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP Total
findings
Eng&WaBe Dk Ger 1 Ger 2 Lit Nl Nor 1 Nor 2 Sco Swe EUCA*
Shipping study
Taxon Engl
Limnothrix planctonica vB 1
Nodularia spumigena vB 1
Merismopedia glauca xB 1
Merismopedia punctata vB 1
Merismopedia sp. vB 1
Mycroscystis sp. vB 1
Oscillatoria sp. v B 1
Phormidium fovealarum vB 1
Phormidium tenue vB 1
Phormidium sp. v B 1
Planktothrix agardhii vBvB 2
Pseudanabaena limnetica vB 1
Snowella septentrionalis vB 1
Snowella lacustris vB 1
Chrysophyceae, indet. vB vB 2
Calycomonas sp. vB 1
Dinobryon sp. vB 1
Mallomonas sp. v B v B 2
Meringosphaera mediterranea vB 1
Uroglena sp. vB 1
Dictyochophycaea
Apedinella spinifera vB 1
Dictyocha fibula vB x B 2
Dictyocha sp. vB 1
Dictyocha speculum xB 1
cf. Parapedinella sp. vB 1
Pseudopedinella pyriforme vB 1
Prymnesiophyceae, indet. vB 1
Chrysochromulina sp. vB 1
Phaeocystis globosa vB 1
Phaeocystis pouchetii xB 1
Phaeocystis sp. v B vB vB 3
Bacillariophyceae (Diatoms), indet. vB vBvBvB xBv*B vBvB 8
Achnanthes sp. vB v*B vB 3
Actinocyclus octonarius vB 1
Actinocyclus sp. (cf.) vB 1
Actinoptychus senarius v B, S v B v B x B v B 5
Actinoptychus splendens xB 1
Actinoptychus sp. vB 1
Amphora costata vB 1
Amphora sp. v B v B x B v B 4
Anaulus sp. v B 1
Asterionella formosa vB vBvB xB 4
Asterionellopsis sp. v B, S x B v B 3
Asterionellopsis glacialis vB vB x B vB 4
Asterionellopsis kariana vB vBv B x B 4
Asteromphalus sp. v B 1
Attheya septentrionalis vB 1
Aulacodiscus argus xB 1
Aulacodiscus kittoni xB 1
Aulacoseira granulata vB x B v B 3
Aulacoseira sp. vB vB 2
Auliscus sculptus xB 1
Auliscus sp. v S 1
Azpeitia sp. xB 1
Bacillaria paxillifera xB vBvB 3
Bacillaria sp. vB 1
Bacteriastrum sp. v S x B 2
Bellerochea horologicales vB 1
Bellerochea malleus (forma malleus) xB vB 2
Biddulphia alternans vB xB vB 3
Biddulphia biddulphiana xB 1
Biddulphia pulchella vB 1
Biddulphia reticulum vB 1
Biddulphia sp. v B vB 2
Brockmanniella brockmannii vB x B 2
Caloneis liber vB 1
Caloneis sp. v B 1
Cerataulus cf. smithii vS xB 2
Cerataulina dentata vB 1
Cerataulina pelagica vB x B vB 3
Chaetoceros affinis vB x B v B 3
Chaetoceros cf. baculites v* B 1
Chaetoceros brevis vB 1
Chaetoceros compressus vB vB vB 3
Chaetoceros constrictus vB 1
Chaetoceros costatus vB 1
Chaetoceros curvisetus v B vB vB 3
Chaetoceros danicus vBvB vBvB 4
Chaetoceros cf. debilis v B v B v B v* B, S 4
Chaetoceros decipiens vB vB 2
Chaetoceros densus vB 1
Chaetoceros diadema v B x B v* B, S v B 4
Chaetoceros didymum vB x B 2
Chaetoceros eibenii vB 1
Chaetoceros laciniosus vB 1
Chaetoceros lauderi vB 1
LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP Total
findings
Eng&WaBe Dk Ger 1 Ger 2 Lit Nl Nor 1 Nor 2 Sco Swe EUCA*
Shipping study
Taxon Engl
Chaetoceros minima vB 1
Chaetoceros protuberans vB 1
Chaetoceros pseudocrinitum xB 1
Chaetoceros similis vB v B 2
Chaetoceros simplex xB 1
Chaetoceros cf. socialis vBv B v*B v B 4
Chaetoceros subtilis v B x B v* B, S 3
Chaetoceros sp. v B v B v B x B v* S v B v B 7
Chaetoceros wighamii vB v B 2
Climacosphenia loniligera vB 1
Cocconeis sp. v B x BvB vB 4
Coscinodiscus centralis xB 1
Coscinodiscus exentricum vB 1
Coscinodiscus granii xB 1
Coscinodiscus lineatus xB vB 2
Coscinodiscus radiatus vB xB vB 3
Coscinodiscus stellaris vB 1
Coscinodiscus sp. v B, S v B x B v B v B v B 6
Coscinodiscus thorii vB 1
Cymbella sp. vB 1
Cyclostephanos sp. xB 1
Cyclotella caspia xB 1
Cyclotella menegheama vB x B 2
Cyclotella meneghiniana xB 1
Cyclotella striata vB 1
Cyclotella sp. v B vBvB x B vB 5
Cylindrotheca closterium vB x B vB 3
Cylindrotheca sp. vB 1
Cymatosira lorenziana xB 1
Dactyliosolen blavyanus vB 1
Dactyliosolen fragillissimus v B vB vB x B vB 5
Dactyliosolen mediterraneus xB 1
Dactyliosolen phuketensis vB 1
Dactyliosolen sp. v B x B 2
Detonula confervacea vB vB vB 3
Detonula pumila vBv B x B 3
Detonula plumula xB 1
Detonula schroederi xB 1
Diatoma elongatum vB 1
Diatoma sp. v B vB x B vB 4
Dimeregramma sp. v B 1
Dimeregrammopsis sp. v B 1
Diploneis bombus vB 1
Diploneis crabro vB 1
Diploneis lineata vB 1
Diploneis smithii vB 1
Diploneis sp. v B x B v B v B 4
Ditylum brightwellii v B, S v B v B x B v* B v B v B 7
Ditylum sp. v B 1
Donkinia lata vB 1
Eucampia zodiacus vB x B vB 3
Eucampia sp. v B v B 2
Eunotogramma dubium vB 1
Fragilaria crotonensis vB 1
Fragilaria sp. v B, S x B v B 3
Fragilariopsis cylindrus xB 1
Frustulia sp. v B 1
Gomphonema sp. vB 1
Grammatophora marina xB 1
Grammatophora sp. xB 1
Guinardia cylindrus vB 1
Guinardia delicatula vBvBvB x B vB 5
Guinardia flaccida vB vB x B vB 4
Guinardia striata vB x B vB 3
Guinardia sp. v B x B 2
Gyrosigma (cf.) sp. v B vB 2
Heliotheca sp. v B 1
Hemiaulus sinensis xB 1
Hemiaulus sp. v B 1
Hemidiscus cuneiformis vB 1
Hyalodiscus scoticus vB 1
Hyalodiscus sp. vB 1
Lauderia annulata vB x B vB 3
Lauderia sp. v B x B v B 3
Lennoxia sp. (cf.) vB 1
Leptocylindrus danicus vB vB x B vBvB 5
Leptocylindrus mediterraneus xB 1
Leptocylindrus minimus vB x B v B vB 4
Leptocylindrus sp. v B vB 2
Licmophora flabellata vB 1
Licmophora gracilis vB 1
Licmophora lyngbyei xB 1
Licmophora sp. xB 1
Lithodesmium undulatum vB x B 2
Lithodesmium sp. vBvB 2
Melosira arctica xB 1
Melosira sp. v B, S v B v B x B v B v B 6
LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP Total
findings
Eng&WaBe Dk Ger 1 Ger 2 Lit Nl Nor 1 Nor 2 Sco Swe EUCA*
Shipping study
Taxon Engl
Melosira nummuloides vB x B v B 3
Melosira varians vBvB 2
Meridion sp. vB 1
Meuniera membranacea vBvB x B vB 4
Meuniera sp. v B 1
Navicula elegans vB 1
Navicula finimarchia vB 1
Navicula pelagica xB 1
Navicula transitrans (indet. var.) vB 1
Navicula sp. v B vBvBvB x BvB vB 7
Neostreptotheca subindica vB 1
Nitzschia acicularis vB 1
Nitzschia accuminata vB 1
Nitzschia bicapitata vB 1
Nitzschia bilobata vB 1
Nitzschia (cf.) droebachensis vB 1
Nitzschia frigida vB 1
Nitzschia frustulum vB 1
Nitzschia linearis vB 1
Nitzschia longissima vB v B vB x B vB 5
Nitzschia (cf.) nana vB 1
Nitzschia sigma vB 1
Nitzschia sp. v B, S v B v B x B v B v B 6
Nitzschia ventricosa vB 1
Odontella alternans vB 1
Odontella aurita vBvB x B vB 4
Odontella granulata vB x B vB 3
Odontella mobiliensis vB x B vBvB 4
Odontella regia vB x B v B 3
Odontella rhombus (forma rhombus) v S v B x B v B 4
Odontella sinensis vB x B v B vB 4
Odontella sp. v S x B v* B v B 4
Oestrupia musea vB 1
Paralia sp. v B 1
Paralia sulcata v B, S v B x B v B v B 5
Pinnularia sp. v B v B 2
Pinnularia viridis vB 1
Plagiogramma staurophorum xB 1
Plagiogramma sp. xB 1
Plagiogrammopsis vanheurckii vBv B 2
Planktoniella sol xB 1
Planktoniella sp. v B 1
Pleurosigma itium vB 1
Pleurosigma sp. v B v B v B x B v B 5
Podocystis sp. v B 1
Podosira stelliger vS vB vB 3
Porosira glacialis xB 1
Proboscia alata v B vB x B vBvB 5
Pseudoeunotia doliolus vB 1
Pseudo-nitzschia delicatissima vB x B v B 3
Pseudo-nitzschia cf. seriata xB vB 2
Pseudo-nitzschia seriata vB vB vB 3
Pseudo-nitzschia sp. vBvB vBvB 4
Pseudosolenia calcar avis xB 1
Rhabdonema sp. v B 1
Rhaphoneis alata vB 1
Rhaphoneis amphiceros v B v B, S v B v B 4
Rhaphoneis suriella vB 1
Rhaphoneis sp. v B x B v B v B 4
Rhizosolenia delicatula vB 1
Rhizosolenia fallax vB 1
Rhizosolenia hebetata (forma hebetata) vB xB vB 3
Rhizosolenia hebetata (forma semispina) vB 1
Rhizosolenia imbricata vB x B v B vB 4
Rhizosolenia indica vB 1
Rhizosolenia longiseta vB 1
Rhizosolenia pungens vB 1
Rhizosolenia robusta vB xB vB 3
Rhizosolenia setigera v B vBvB x B vBvB 6
Rhizosolenia sp. v B vB vB x B vB 5
Rhizosolenia styliformis vB xB vB 3
Rhopalodia musculus vB 1
Roperia tessellata xB 1
Skeletonema costatum v B v B, S v B v B v B x B v, v* B, S v B 8
Skeletonema sp. v B vBvB 3
Skeletonema subsalsum vB 1
Stephanodiscus astraea vB 1
Stephanodiscus sp. v B, S x B 2
Stephanopyxis turris v B vB x B vB 4
Striatella unipunctata xB vB 2
Subsilicea fragilarioides vB 1
Surirella brightwell vB 1
Surirella sp. v B x B v B 3
Synedra sp. v B, S v B v B v B 4
Tabellaria fenestrata vB vB 2
Tabellaria sp. vB vB vB 3
Thalassionema sp. v B 1
LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP Total
findings
Eng&WaBe Dk Ger 1 Ger 2 Lit Nl Nor 1 Nor 2 Sco Swe EUCA*
Shipping study
Taxon Engl
Thalassionema frauenfeldii vB 1
Thalassionema nitzschioides vB v B vBvB x B vBvB 7
Thalassiosira allenii xB 1
Thalassiosira anguste-lineata vB x B 2
Thalassiosira antarctica xB 1
Thalassiosira eccentrica vB x B 2
Thalassiosira gravida vB 1
Thalassiosira hendeyi vB 1
Thalassiosira hyalina xB 1
Thalassiosira leptosus vB 1
Thalassiosira nordenskioeldii vB x B 2
Thalassiosira punctigera vB 1
Thalassiosira rotula vB x B v*B 3
Thalassiosira subtilis vB vB 2
Thalassiosira sp. v B v B v B v B x B v, v* B, S v B 7
Thalassiothrix nitzschioides 0
Triceratium alternans vB 1
Triceratium favus xB vB 2
Triceratium sp. vBvB 2
Trigonium sp. xB vB 2
Undatella sp. v B 1
Xanthophyceae vB 1
Goniochloris muticum vB x B 2
Tribonema sp. v, r B 1
Dinophyceae (Dinoflagellates), indet. r B, S v B r B x, r B, S v B 5
Alexandrium cf. affine rS rB 2
Alexandrium cf. lusitanicum rS rS 2
Alexandrium cf. minutum rS rS 2
Alexandrium cf. tamarense/catenella x, r B, S v, r B , S 2
Gonyaulax excavata rS 1
Alexandrium sp. v, r B, S r B, S v, r B, S 3
Amphidinium sphenoides vB vB 2
Amphidinium sp. vB x B vB 3
Bitectatodinium tepikiense *** rS 1
Ceratium furca vBv B x B vB 4
Ceratium fusus v B x B v B v B 4
Ceratium cf. horridum vB x B vBvB 4
Ceratium lineatum vB x B vB 3
Ceratium longipes vB x B vB 3
Ceratium macroceros xB vBvB 3
Ceratium massiliense vB 1
Ceratium minutum xB 1
Ceratium tripos (var. tripos) vB x B vBvB 4
Ceratium sp. v B vBvB 3
Diplopelta parva rS rS 2
Diplopelta symmetrica rS 1
Diplopsalis lenticula vB x B rS 3
Diplopsalis sp. xB vB 2
cf. Dissodinium asymmetricum vB 1
Dissodinium pseudolunula xB rS 2
Ebria tripartita vB x B 2
Fragilidium subglobosum xB 1
Glenodinium foliaceum xB 1
Glenodinium sp. xB 1
Gonyaulax digitale xB 1
Gonyaulax grindleyi xB rS 2
Gonyaulax polygramma xB 1
Gonyaulax scrippsae xB rS 2
Gonyaulax spinifera xB rS 2
Gonyaulax verior vB rS 2
Gonyaulax sp. v B v, r B, S x B v B 4
Gymnodinium cf. catenatum rS 1
Gymnodinium filum xB 1
Gymnodinium cf. mikimotoi xB 1
Gymnodinium sanguineum xB 1
Gymnodinium vulffi vB 1
Gymnodinium sp. v B x, r B, S v* B, S v B 4
Gyrodinium fusiforme x B 1
Gyrodinium pellucidum x B 1
Gyrodinium spirale x B 1
Gyrodinium sp. v B x B 2
Heterocapsa rotundata vB r S 2
Heterocapsa triquetra vB v B x B r S 4
Heterocapsa sp. xB 1
Lingulodinium machaerophorum *** rS 1
Lingulodinium polyedrum rS 1
Mesoporos sp. v B 1
Nematoshaeropsis labyrinthea rS 1
Noctiluca scintillans vB v B x B 3
Noctiluca sp. vB 1
Oblea rotunda rS vB rS 3
Operculodinium centrocarpum *** r B, S 1
Orthopithonella granifera *** rS 1
Oxhyrris marina vB 1
Oxytoxum oxytoxoides vB 1
Oxytoxum scolopax xB 1
Oxytoxum sp. xB vB 2
LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP Total
findings
Eng&WaBe Dk Ger 1 Ger 2 Lit Nl Nor 1 Nor 2 Sco Swe EUCA*
Shipping study
Taxon Engl
Pentapharsodinium dalei rS rS 2
Pentapharsodinium thyrrhenicum rS 1
Peridinella catenata vB 1
Peridinium palatinum v S 1
Peridinium sp. v B 1
Peridinium willei v S 1
Phalacroma (Dinophysis) acuminata vB v B x B 3
Phalacroma (Dinophysis) acuta vB x B 2
Phalacroma (Dinophysis) caudata xB 1
Phalacroma (Dinophysis) norvegica v S v B x B v B 4
Phalacroma (Dinophysis) ovum vB 1
Phalacroma (Dinophysis) rotundata vB x B 2
Phalacroma (Dinophysis) sp. v B xB vBvB 4
Pheopolykrikos hartmannii r B, S r S 2
Polykrikos kofoidii rS 1
Polykrikos schwartzii r S x, r B, S v, r B, S 3
Prorocentrum balticum vB 1
Prorocentrum compressum xB 1
Prorocentrum dentatum xB 1
Prorocentrum gracile xB 1
Prorocentrum micans v B v B v B x B v B 5
Prorocentrum minimum vB vBvB xB 4
Prorocentrum sp. vB x B vBvB 4
Protoperidinium americanum r B, S r S 2
Protoperidinium bipes v B x B, S 2
Protoperidinium brevipes v B x B, S v B 3
Protoperidinium claudicans rS rS vB 3
Protoperidinium compressum r S r B, S r S 3
Protoperidinium conicoides rS 1
Protoperidinium conicum v B x, r B, S v, r B, S 3
Protoperidinium curtipes xB 1
Protoperidinium depressum vB x B vBvB 4
Protoperidinium diabolum xB 1
Protoperidinium divaricatum rS rS 2
Protoperidinium divergens vBvB 2
Protoperidinium excentricum rS vB rS rS 4
Protoperidinium cf. grande vB 1
Protoperidinium leonis r S x, r B, S v, r B, S 3
Protoperidinium cf. minutum rS rS 2
Protoperidinium oblongum r S v B x, r B, S r S 4
Protoperidinium ovatum xB 1
Protoperidinium pallidum vB 1
Protoperidinium cf. pellucidum vB x B 2
Protoperidinium pentagonum rS xB rS 3
Protoperidinium pyriforme xB vB 2
Protoperidinium subinerme rS rS 2
Protoperidinium sp. v B r S v B v, r B, S 4
Protoperidinium steinii vB x B 2
Protoperidinium subinerme rS vB 2
Pyrophacus horologium rS 1
Pyrophacus steinii var. vancampoae *** rS 1
Scrippsieila crystallina rS rS 2
Scrippsiella hangoei vB r S 2
Scrippsiella lachrymosa rS rS rS 3
Scrippsiella precaria rS 1
Scrippsiella rotunda rS rS 2
Scrippsiella trifida rS rS 2
Scrippsiella trochoidea r S v B x'''', r'' B, S v, r B, S 4
Scrippsiella sp. v B x'', r' B, S 2
Spiniferites bentori rS 1
Spiniferites sp. r B, S r S 2
Zygabikodinium lenticulatum rS vB rS rS 4
Zygabikodinium sp. vB 1
Cryptophyceae, indet. vBvBvB x B vB 5
Cryptomonas ovata vB 1
Cryptomonas sp. vB 1
Leucocryptus sp. cf. vB v B 2
Teleaulax acuta vB 1
Prasinophyceae, indet. vB 1
cf. Nephroselmis sp. vB 1
Pachysphaera cf. pelagica vB 1
Pterosperma cf. nationalis vB 1
Pterosperma sp. vB 1
Pyramimonas sp. vBv B 2
Chlorophyceae, indet vB v*B v B 3
Actidesmium sp. v B 1
Actinastrum sp. v B vB 2
Ankistrodesmus sp. vB vB 2
Chlorella sp. vB 1
Chlorhormidium sp. v B 1
Coelastrum astroideum vB 1
Coelastrum microporum xB 1
Coelastrum sphaericum vB 1
Coelastrum sp. vBvB 2
Crucigenia fenestrata xB 1
Crucigenia sp. v B vB 2
Desmidales, indet. vB 1
LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP Total
findings
Eng&WaBe Dk Ger 1 Ger 2 Lit Nl Nor 1 Nor 2 Sco Swe EUCA*
Shipping study
Taxon Engl
Dictyosphaerium ehrenbergianum vB 1
Dictyosphaerium pulchellum vB v B 2
Enallax (Scenedesmus) acutiformis vB 1
Geminella sp. v B 1
Gloeocystis (Chlamydocapsa) sp. v B 1
Haematococcus sp. vB 1
Kirchneriella sp. vB 1
Koliella sp. (cf.) v* B 1
Lagerheimia sp. v B 1
Microactinium sp. v B 1
Monoraphidium arcuatum vB 1
Monoraphidium contortum vBvB 2
Monoraphidium mirabile vB 1
Oocystis borgei vB 1
Oocystis lacustris vB 1
Oocystis marssonii vB 1
Oocystis sp. v B 1
Pandorina sp. v B 1
Pediastrum boryanum xB vBvB 3
Pediastrum duplex var. duplex vB xB vB 3
Pediastrum duplex var. gracillimum vB 1
Pediastrum sp. v B, S v B v B x B v B 5
Pediastrum tetras vB 1
Planktonema lauterbornii vB 1
Planktonema sp. v B 1
Prasiola sp. vB 1
Pyramidomonas sp. vB 1
Scenedesmus aculeolatus vB 1
Scenedesmus acuminatus vB vB xB vBvB 5
Scenedesmus acutus vB vB vB 3
Scenedesmus armatus vB 1
Scenedesmus obliquus vB 1
Scenedesmus opoliensis vB x B 2
Scenedesmus quadricauda var. quadricauda vB xB vBvB 4
Scenedesmus quadricauda var. longispina vB 1
Scenedesmus sempervirens vB 1
Scenedesmus sp. v B, S v B v B v B 4
Schroederia sp. v B 1
Selenastrum capricornutum vB 1
Selenastrum sp. v B 1
Sorastrum sp. v B 1
Staurastrum paradoxum vB 1
Staurastrum sp. v B vB vB 3
Staurastrum tetracerum vB 1
Staurodesmus sp. vB 1
Tetraëdron caudatum vB xB 2
Tetraëdron sp. v B vB 2
Tetrastrum sp. v B vB vB 3
Tetrastrum staurogeniaeforme vB vB xB 3
Ulotrichales, indet. vB 1
Rhodophyta, parts xB 1
Filamentous green alga, indet. vB 1
Gloeotila sp. xB 1
Fauna
Cnidaria
Anthozoa, indet. aB 1
Octocorallia a B 1
Hydrozoa, indet. aB aBaB 3
Bougainvillae sp. aB 1
Manet sp. aB 1
Obelia sp. aB 1
Sarsia sp. aB 1
Rathkea octopunctata aB a B 2
Scyphozoa, indet. lB 1
Siphonophora, indet. aB a B 2
Abylidae, indet. aB 1
Ctenophora, indet. aB 1
Pleurobrachia pileus aB a B 2
Tentaculata, indet lB 1
Bryozoa, indet. lB lB 2
Conopeum reticulum aS 1
Crassimarginetella sp. a S 1
Hincksina sp. a S 1
Membranipora tuberculata aS 1
Watersipora arcuata aS 1
Turbellaria, indet. a B a B 2
Trematoda, indet. lB 1
Nemertea, indet. aB 1
Rotatoria, indet. l B, S a B l B a B a B a B a B a B 8
Anuraeopsis fissa aB 1
Brachionus budapestensis aB 1
Brachionus calyciflorus anuraeiformis a B,S 1
Brachionus diversicornis aB 1
Brachionus quadridentatus rhenanus aB 1
Brachionus urceolaris a B,S 1
Keratella cochlearis cochlearis a B aBaB aB 4
LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP Total
findings
Eng&WaBe Dk Ger 1 Ger 2 Lit Nl Nor 1 Nor 2 Sco Swe EUCA*
Shipping study
Taxon Engl
Keratella cochlearis tecta aB 1
Keratella irregularis aB 1
Keratella quadrata a B,S a B 2
Keratella sp. aB 1
Kellicottia sp. aB 1
Notholca sp. aB 1
Synchaeta baltica a, l B 1
Synchaeta sp. aB 1
Nematoda, indet. a S a B a B a B 4
Anaplostoma sp. a S 1
Axonolaimus sp. j S 1
Chromadorida, indet. a S 1
Chromaspirina sp. a S 1
Desmodora sp. a S 1
Diplolaimella sp. a B 1
Microlaimus sp. j, a S 1
Monhystera sp. j, a B 1
Oncholaimus oxyuris j, a B,S 1
Oncholaimus sp. a S 1
Prochromadorella sp. j B 1
Rhabditidae, indet j, a B 1
Teratocephalidae, indet a B 1
Theristus sp. a S 1
Tripoloides marinus j, a S 1
Mollusca
Gastropoda indet. l B l B l B l B l B l B l B 7
Aporrhais sp. l S 1
Atlanta oligogyra aS 1
Atlanta sp. a S 1
Barleeia sp. a S 1
Buccinidae, indet l S 1
Caecidae, indet l S 1
Cavolinia inflexa imitans aS 1
Cavolinia longirostris limbata aS 1
Cerithiimorpha, indet l B 1
Chaenogastropopda, indet l B,S 1
Clio pyramidata lanceolata aS 1
Clione limicina aB 1
Crepidula fornicata aS 1
Crepidula sp. l S 1
Fasciolariidae, indet j S 1
Hydrobiidae, indet a S 1
Limacina helicina helicina lS 1
Limacina inflata lS 1
Limacina lesueuri lS 1
Limacina trochiformis l B,S 1
Limacina sp. l S 1
Littorinidae indet l B,S eB 2
Muricidae, indet l S 1
Neogastropoda, indet l S 1
Neritidae, indet l S 1
Odostomia sp. a S 1
Opisthobranchia l S 1
Rissoa sp. l B,S 1
Turridae, indet. l S 1
Bivalvia, indet. l B l B l B l B l B l B l B 7
Anomia ephippium aS 1
Anomia simplex aS 1
Argopecten irradians aS 1
Astarte borealis aS 1
Cerastoderma edule aS 1
Cerastoderma lamarcki aS 1
Corbula barrattiana aS 1
Corbula gibba aS 1
Crassostrea gigas l, a S 1
Crassostrea virginica aS 1
Crassostrea sp. l B 1
Cryptomya californica aS 1
Dreissena polymorpha aS 1
Gryphaeidae, indet l S 1
Lyrodus pedicellatus lB 1
Macoma balthica aS 1
Mactra corallina a, l B, S 1
Modiolus barbatus lS 1
Modiolus demissus lS 1
Modiolus modiolus aS 1
Modiolus sp. l B,S 1
Montacutidae, indet l B,S 1
Mya arenaria aS lB 2
Mytilaster lineatus lB 1
Mytilus edulis a, l B, S a S 2
Mytilus galloprovincialis lS 1
Mytilus sp. a S 1
Ostrea edulis a, l S 1
Ostrea sp. a S 1
Parvicardium ovale aS 1
Pinnidae, indet l S 1
LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP Total
findings
Eng&WaBe Dk Ger 1 Ger 2 Lit Nl Nor 1 Nor 2 Sco Swe EUCA*
Shipping study
Taxon Engl
Placamen tiara aS 1
Pteriidae, indet l B 1
Semele sp. a S 1
Septifer sp. l S 1
Solenidae, indet a S 1
Tellina fabula aS 1
Tellina sp. l S 1
Teredinidae, indet l B 1
Teredo navalis lB 1
Venus fasciata aS 1
Scaphopoda
Pseudantalis cf. rubescens aS 1
Polychaeta, indet. l B, S a, l S l B l B l B l B l, a B 7
Capitella capitata aS 1
Ficopomatus enigmaticus aS 1
Hydroides elegans aS 1
Nephthyidae, indet a S 1
Nereidae, indet. a, e B 1
Nereis pelagica aS lB 2
Nereis succinea aS lB 2
Polydora ciliata lB 1
Polydora sp. l B l B j S l B 4
Scyllidae, indet. aB 1
Sedentaria l B 1
Spionidae, indet l B j S l B a B 4
Spio filiformis lB 1
Terebellidae, indet lB 1
Tomopteris septentrionalis aB 1
Oligochaeta, indet. l B 1
Nais sp. a S 1
Tubificidae jS 1
Pantopoda
Nymphon sp. j S 1
Crustacea, indet l B lB 2
Cladocera, indet r S a B a B 3
Alona rectangula aB 1
Bosmina coregoni jB aB aB aB 4
Bosmina coregoni maritima a B,S 1
Bosmina longirostris j,a B,S 1
Bosmina sp. l B a B a B a B 4
Ceriodaphnia sp. j B a B 2
Chydorus sphaericus aB 1
Chydorus sp. a S 1
Ctenodaphnia sp. r S 1
Daphnia ambigua aB 1
Daphnia cucullata incerta aB 1
Daphnia galeata j, a, r B,S a B 2
Daphnia pulex aBaB 2
Daphnia sp. a, r S 1
Evadne nordmani aB a B aB 3
Evadne spinifera aB 1
Evadne sp. aB 1
Hyalodaphnia sp. r B,S 1
Ilyocryptus sordidus jS 1
Leydigia acanthoceroides a, r B,S 1
Moina micrura rS 1
Penilia avirostris aB 1
Podon intermedius aB 1
Podon polyphemoides aB a B 2
Podon cf. leuckarti aB 1
Podon sp. aB a B 2
Simocephalus expinosus aB 1
Ostracoda, indet. aB lBaB 3
Aurila sp. l S 1
Conchoecia elegans aB 1
Cypria sp. a S 1
Cyprideis salebrosa aS 1
Cyprideis sp. a S 1
Cypridina sp. a B 1
Myodocopidae, indet l, a B 1
Paracypris sp. a B 1
Paradoxostoma sp. a B 1
Sarsiellidae, indet a S 1
Urocythereis sp. a S 1
Copepoda, indet. a B l, e B l, e B l, a B 4
Calanoida, indet. l B a B l, a B l B l B a B a B 7
Acartia bifilosa inermis aB a B 2
Acartia clausi j, a B a B l, a B a B a, l B 5
Acartia discaudata aB a B 2
Acartia grani aB a B 2
Acartia hudsonica aB 1
Acartia latisetosa aB 1
Acartia lilljeborgi aS 1
Acartia longiremis aB aB a B a B aB 5
Acartia tonsa j, a B a B 2
Acartia spinicauda aB 1
Acartia sp. j, a B l, a B a B a B 4
LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP Total
findings
Eng&WaBe Dk Ger 1 Ger 2 Lit Nl Nor 1 Nor 2 Sco Swe EUCA*
Shipping study
Taxon Engl
Acrocalanus longicornis aB 1
Anomalocera pattersoni aB 1
Anomalocera sp. lB 1
Calanoides cariantus aB 1
Calanus finmarchicus aB aB aB aBaB 5
Calanus helgolandicus aB a B aB 3
Calanus sp. lB aB 2
Candacia sp. aB 1
Centropages furcatus aB aB 2
Centropages hamatus a B a B a B l, a B a B 5
Centropages typicus a B aB a B aBaB 5
Centropages velificatus aB 1
Centropages sp. a B aB 2
Clausocalanus arcuricornis aB 1
Clausocalanus furcatus aB 1
Ctenocalanus vanus aB 1
Diaptomus sp. l, a B 1
Euchaeta sp. aB 1
Eurytemora affinis a B,S a B l, a B 3
Eurytemora hirundoides a B,S a B a B a B a B 5
Eurytemora lacustris aB 1
Eurytemora sp. j, a B a B a B 3
Isias clavipes aB 1
Labidocera woolastoni aB 1
Limnocalanus macrurus aB 1
Limnocalanus sp. aB 1
Metridia brevicauda aB 1
Metridia lucens aB aB 2
Microcalanus pusillus aB 1
Microcalanus sp. aB 1
Paracalanus denudatus aB 1
Paracalanus parvus a B a B l, a B a B a B 5
Paracalanus sp. lBaB 2
Pseudocalanus elongatus a B a B l, a B a B a B a B 6
Pseudocalanus sp. a B aB 2
Pseudocyclops crassiremis aB 1
Temora discuadata aB 1
Temora longicornis a B a B a B l, a B a B a B a B 7
Temora stylifera aB 1
Temora turbinata aB 1
Temora sp. j B l, a B a B a B 4
Tortanus discaudatus aB 1
Cyclopoida, indet. l B a B a B 3
Acanthocyclops robustus aB 1
Acanthocyclops sp. j B 1
Cyclops abyssorum aB 1
Diacyclops sp. j S 1
Megacyclops sp. a B 1
Mesocyclops leuckarti aB aB 2
Mesocyclops sp. a B 1
Oithona amazonica aB 1
Oithona attenuata aB 1
Oithona brevicornis aB aB 2
Oithona hebes aB 1
Oithona helgolandica aB aB 2
Oithona nana aB a B aB 3
Oithona oculata aB 1
Oithona plumifera aB 1
Oithona similis l, a B a B a, l B 3
Oithona simplex aB 1
Oithona sp. j, a B,S l, a B l B a B 4
Thermocyclops crassus aB 1
Thermocyclops sp. j S 1
Harpacticoida, indet. a B a S a B a B a B a B a, l B a B 8
Diosaccus sp. a B 1
Ectinosoma minor aS 1
Ectinosoma sp. j, a B,S 1
Enhydrosoma propinquum aS 1
Euterpina acutifrons aB aB 2
Euterpina sp. a B,S aB 2
Halectinosoma sp. a B 1
Laophonte sp. a S 1
Longipedia rosea aB 1
Macrosetella gracilis aB 1
Microarthridion littorale aB 1
Microsetella norvegica aB aB 2
Microsetella rosea a B,S 1
Microsetella sp. aB 1
Monstrilloida, indet. aB 1
Oculosetella sp. aB 1
Onychocamptus sp. a B 1
Pseudobradya sp. a S 1
Pseudonychocamptus koreni aB 1
Pseudozosime sp. a B 1
Robertsonia sp. a B 1
Stenocaris sp. a B 1
Strilloma grandis aB 1
LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP Total
findings
Eng&WaBe Dk Ger 1 Ger 2 Lit Nl Nor 1 Nor 2 Sco Swe EUCA*
Shipping study
Taxon Engl
Tachidius discipes aB 1
Tisbe elegantula j, a B 1
Tisbe furcata aB 1
Tisbe graciloides a B,S 1
Tisbe gurneyi aB 1
Tisbe inflata aB 1
Tisbe longicornis aB 1
Tisbe wilsoni aB 1
Tisbe sp. j, a B,S 1
Poecilostomatoida, indet. l B 1
Corycaeus anglicus aB aB 2
Corycaeus brehmi aB 1
Corycaeus latus aB 1
Corycaeus sp. aB a B aB 3
Lubbockia aculeata aB 1
Oncaea clevi aB 1
Oncaea conifera aB 1
Oncaea dentipes aB 1
Oncaea media aB 1
Oncaea mediterranea aB 1
Oncaea subtilis aB 1
Oncaea venusta a B,S aB 2
Oncaea sp. l, a B l, a B a B 3
Pachos sp. a B 1
Saphirella tropica aB 1
Siphonostomatoida, indet. aB 1
Cirripedia, indet. lB l B lB 3
Balanomorpha, indet. l B a, l S lB 3
Balanus amaryllis aS 1
Balanus amphitrite amphitrite aSaS 2
Balanus amphitrite cirratus aS 1
Balanus amphitrite communis aS 1
Balanus balanoides aS 1
Balanus balanus a, l B,S 1
Balanus campbelli aS 1
Balanus crenatus aS 1
Balanus eburneus aS 1
Balanus improvisus a, l B, S a S 2
Balanus perforatus aS 1
Balanus reticulatus aS 1
Balanus trigonus aS 1
Balanus uliginosis aS 1
Balanus variegatus aS 1
Balanus sp. l B l B l B 3
Elminius modestus aS 1
Elminius simplex aS 1
Megabalanus tintinnabulum tintinnabulum aS 1
Tetraclita aoranga aS 1
Lepadomorpha
Lepas sp. l B 1
Mysidacea, indet. aB a B lBaB 4
Gastrosaccus normanni aB 1
Gastrosaccus spinifer aB 1
Gastrosaccus sp. aB 1
Hemimysis sp. aB 1
Mesopodopsis slabberi j, a B a B 2
Neomysis integer aB aB 2
Paramysis kervillei aB 1
Paramysis helleri aB 1
Paramysis spiritus aB 1
Praunus flexuosus aB 1
Isopoda, indet. aB lB 2
Cirolanidae, indet. aB 1
Cymodoce truncata aB 1
Eurydice pulchra aB 1
Eurydice truncata aB 1
Sphaeroma rugicauda aS 1
Amphipoda , indet. aB 1
Bodotriidae, indet. a B 1
Caprella sp. jB 1
Corophium acherusicum a B,S 1
Corophium volutator aB 1
Ericthonius brasiliensis aB 1
Gammarus salinus aS 1
Gammarus zaddachi a B,S 1
Gammarus sp. a B,S 1
Jassa marmorata aS 1
Jassa sp. a S 1
Microdeutopus gryllotalpa aS 1
Paracaprella tenuis aB 1
Parathemisto sp. aB 1
Cumacea, indet. aB a B aB 3
Euphausiacea, indet. aB 1
Euphausiid calyptopsis jB 1
Euphausiid furcilia jB 1
Decapoda, indet. l B l B l B l, a B l B l B 6
Ashtoret maculata aS 1
LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP LS SP Total
findings
Eng&WaBe Dk Ger 1 Ger 2 Lit Nl Nor 1 Nor 2 Sco Swe EUCA*
Shipping study
Taxon Engl
Callinectes sapidus aS 1
Callinectes sp. lB 1
Carcinus maenas j, a S 1
Charybdis feriatus j S 1
Charybdis helleri aS 1
Charybdis sp. a B 1
Crangon crangon j, a B,S a B 2
Eriocheir sinensis aS 1
Panopeus sp. a S 1
Pirimela denticulata lB 1
Portunus pelagicus jS 1
Portunus petreus aS 1
Portunus sp. lB 1
Acari, indet. aB 1
Insecta, indet. l B l B 2
Clunio sp. a S 1
Trichoptera, indet. l S 1
Arachnida
Hydrachnellae indet. lB lB 2
Chaetognatha, indet. aB 1
Sagitta sp. aB a B aBaB 4
Sagitta elegans aB 1
Sagitta setosa aB 1
Sagitta macrocephala aB 1
Echinodermata, indet. lB lB 2
Ophiuroidea l B l B 2
Tunicata, indet. aB 1
Ascidiacae, indet. lB 1
Thaliacea, indet. aB 1
Dolioletta gegenbauri aB 1
Appendicularia, indet. aB lB 2
Oikopleura sp. aB a B aB 3
Pisces lB 1
Cyclostomata
Petromyzon marinus aS 1
Osteichthyes, indet. e B e B l B 3
Allosmerus elongatus lB 1
Ammodytes tobianus aS 1
Ammodytidae, indet. l, a B 1
Anguilla anguilla aS jB 2
Clupea harengus aB l B 2
Gaidropsarus sp. l, j B 1
Gasterosteus aculeatus a B,S 1
Gobiidae, indet. lB 1
Lipophrys pholis aB 1
Microstomus kitt lB 1
Osmerus eperlanus a B,S 1
Platychthes flesus lB 1
Pomatoschistus minutus a B,S 1
Sprattus sprattus a B,S 1
Syngnathidae aB 1
Syngnathus rostellatus aB 1
Total 28 4 502 19 90 88 181 3 327 41 67 252 68 990
