269

Senckenbergiana maritima

31

(2)

269-281

Frankfurt am Main, 28.12.2001

Senckenberg am Meer 500

Epibenthic Diversity in the North Sea

RUTH Z

& JOHN ALVSV~.G & INGEBORG DE BOOIS & JOHN COTTER &
SIEGFRIED EHRICH & ALEX FORD & HILMAR HINZ &
ASTRID JARRE-TEICHMANN & SIMON JENNINGS & INGRID KR6NCKE &
JOHN LANCASTER & Gr.RJAN PIET & P H I H P PRINCE
With 9 Text-Figures and 1 Table

Keywords: epifauna, diversity, fishing effort, depth, N o r t h Sea.

A bstract

[Z
R. & ALVSV~G,J. & BooTs, I. DE ~( COTTER,J. & EHRICH,S. & FORD,A. & Hl.xz, H.
& JARRE-TEICHMANN,A. & JENNINGS, S. & KRONCKE, I.. & LANCASTER,J. & PIET G. & PRINCE,
P. (200l): Epibenthic diversity in the North Sea. - Senckenbergiana marit., 31 (2): 26%281, 9 figs.,
1 tab.; Frankfurt a. M.]
In 1999 the epibenthic fauna of the North Sea was investigated using the 3'J quarter 'International Bottom Trawl Survey' of five European countries. Altogether 241 stations were sampled
covering 143 ICES rectangles.
The objectives of the project were (i) to analyse epibenthic diversity patterns in the North Sea,
(ii) to identify the spatial distribution of faunal communities and (iii) to relate environmental
factors as well as fishing effort to species diversity.
Epibenthic fauna was clearly divided between the southern North Sea and the centralnorthern North Sea, roughly along the 50 m depth Iine. The separation was based on an overall
higher number of species in the central and northern North Sea a n d a change in the species
composition from north to south.
Sessile fauna including erect, branching species like bryozoans and hydrozoans were particularly diverse along a corridor in the central-northern North Sea between 56~ and 58~ coinciding
with the area between the 50 m and 100 m depth line. Cluster analysis, based on the structure of
the community, confirmed the north-south gradient found for species diversity. Separation of
clusters was driven to a great extent by species occurring predominantly or exclusively north of
the 50 m contour line. Few species were exclusive to the south, but a number of scavenging
species were found here more frequently and in higher numbers.
Depth was positively correlated with the diversity of free-living fauna, whereas the type of
sediment showed no significant relationship with variations in numbers of species. Beam-trawling
effort was negatively correlated with the diversity of sessile fauna.

Authors' addresses:
RUTH Z
ALEXFORD,JOHN LANCASTER,University of Wales Swansea, Biological Sciences, Singleton Park, Swansea, SA2 8PP, U.K.,
e-mail: bdzuhlke@swansea.ac.uk - JOHN COTTER, SIMONJENNINGS, CEFAS, Lowestoft, England, U.K. - SlEGrIED EHRICH, Bundesforschungsanstalt f Fischerei, Institut f Seefischerei, Palmaille 9, D-22767 Hamburg, Germany, e-mail: ehrich.ish@bfa-fisch.de HILMaa HtNZ, INGRiD Ka6NC~E, Forschungsinstitut Senckenberg, Schleusenstrasse 39a, D-26382 Wilhelmshaven, Germany. -JoHt~
ALVSV.~G, Institute for Marine Research, Bergen, Norway. - INGrBORG DE Bools, GERJAN PtET, Netherlands Institute for Fisheries
Research (RIVO), P.O. Box 58, 1970 AB Ijmuiden, The Netherlands. - ASTRtDJARaE-TEICHMANY,PHILIP PRINCE, Danish Institute for
Fisheries Research, Hirtshals, Denmark.

270

Introduction
Epibenthos constitutes a major part of the benthic
community and in the North Sea studies on epibenthic
species were among the first benthic investigations
(PETERSEN 1914, 1918). Since then several epibenthic
studies throughout the North Sea have been carried out
(DxER et al. 1983; F~UZNHHM et al. 1989; JENmNGS et al.
1999a). Re~s et al. (1999) studied the English part of the
North Sea, BaSFORD et al. (1989) the northern North Sea,
DUtNEWLD et al. (1991) the southern North Sea and
KR6NCK~ (1992) described the benthos of the Dogger
Bank. These studies generally support the zonation proposed by GL~MAREC (1973), showing a division between
faunal communities in the southern and northern North
Sea. Environmentat factors like depth, sediment composition, temperature and currents are regarded as being
mainly responsible for distribution patterns of epibenthic

communities (FRAUENHE[M et al. 1989; BASFORD et al.

1990; JENmN6S et al. 1999a). Furthermore, experimental
studies showed considerable effects of fishing on the
composition of benthos (BsatZEmO & DEL~EO 1999;
PRENA et al. 1999; TUCK et al. 2000; KAtSER et al. 2000).
Detailed reviews of potential effects of fishing are given in
LmDEBOOM & DEGROoT (1998) and JeNNINGS & KAISER
(1998). However, sampling grids were usually too coarse
to relate benthos and fishing effort on a large spatial scale.
In this study we attempt a detailed description of the
epibenthic community of the North Sea with a higher
resolution than former surveys. We present spatial distribution patterns of individual species, epibenthic communities and species diversity. Furthermore, we investigate
the relationship of fishing effort and environmental factors with epibenthic diversity in the North Sea.

Material and Methods

In 1999 epibenthos samples were taken at 241 stations
in the North Sea during the 3~d quarter 'International
Bottom Trawl Survey' (IBTS) of five European countries:
Germany, The Netherlands, England, Norway and
Denmark (Fig. 1). The survey covered 143 ICES rectangles
(boxes of 0.5 ~ latitude by 1o longitude); one sample was
taken per station.

E6

E8

I C E S rectangles
FO
F2
F4

F6

F8

52
50
48

46
(t)
O~
t-- 4 4

~Z
e--

42

O9
LU

Sampling Gear and Sample Treatment

All samples were taken with a 2 m beam-trawl fitted
with a 20 mm stretched mesh (10 mm knot to knot) a n d a
liner of 4 mm knotless mesh (2 mm 'knot' to 'knot') fitted
inside the codend. Detail of the equipment is given in
JENNINGS et al. (1999a). Towing warp was 14 mm, 6/19
construction. Due to technical considerations, the Netherlands used 16 mm warp and Norway 10 mm warp. The
tow was shot with a 3:1 ratio of warp length to depth and
towed at approximately 1 knot for 5 minutes. However,
due to local current or wind conditions towing speed
varied. The average length of tows was 197_+56 m.
Samples were washed through a 5 mm sieve and
epibenthic fauna was separated from the remains. The
majority of species were identified on board. Unidentified
fauna was preserved in a 4% seawater formalin solution
and processed later in the respective laboratories. Freeliving species were counted and weighed with an accuracy
of 1 g, sessile animals were recorded as present or absent.
Species names were standardised to the nomenclature of
HowsoN & PICTON (1997). We are aware that some
species names have changed since the publicafion of this
account or ate in the process of being renamed. We have
indicated these changes to the best of our knowledge.

L~ 4O
38

Environmental Factors and Beam-Trawling Effort

36

Depth was recorded at every station sampled and

information about sediment characteristics was derived
from BAS~ORDet al. (1993). Beam-trawling effort data for

34

32

4"

2"

0 ~

2 ~

1ongitude

4"

6 a

8 ~

Fig. 1. Epibenthos sampling stations in the North Sea, 1999. GFR: Germany, NOR: Norway, NED: The Netherlands, ENG:
England, DEN: Denmark.

271

1998 were supplied by Germany, England and The

Netherlands. All data were summed as annual hours
fishing by ICES statistical rectangle. Size of vessel was
not taken into account. Data from Denmark and Belgium,
countries with major fishing interests in the North Sea,
were not available.

Data Analysis
Species were separated into free-living and sessile
benthos. Abundances were standardised to a tow length
of 200 m (400 m0, numbers of species were estimated as
species per haul. For free-living fauna Hill's diversity
indices were calculated (HILL 1973) which incorporate
abundances to a different degree. However, in this paper
species diversity is mainly expressed as numbers of species
per station.
Hierarchical cluster analysis was used to separate
groups of stations with similar communities. We use the
term 'community' for assemblages of similar sessile or
free-living species characteristic for certain areas. Ecological interactions are not implied. Analysis was carried
out with the PRIMER statistical package (CLARKE &

WARXVlCK 1994). For free-living fauna, abundances were

double-square-root transformed, sessile fauna was analysed as present or absent. The Bray-Curtis index was
calculated between each possible pair of samples. Species
which were predominantly responsible for the separation
of clusters were determined with the PRIMER program
SIMPER. This examines the percentage contribution each
species makes to the difference between two groups.
Cluster analysis was based on 73 stations of the English
survey, since dendrograms covering more samples could
not be processed and plotted in a meaningful way and the
English survey covered the whole North Sea.
Beam-trawling effort and environmental factors were
examined for their degree of correlation with numbers of
sessile and free-living species to evaluate possible effects
of fishing on the benthic fauna. The correlation was
carried out for stations <50 m depth. Restricting the
analysis to the relatively shallow stations of the southern
and central North Sea avoided severe interaction with
depth as a factor influencing the benthic community and
hence increased the likelihood of establishing the extent
to which beam-trawling, in comparison to other environmental factors, might influence species diversity.

Results
A total of 423 benthic species were recorded. Of those
270 were classified as free-living and 153 as sessile. Tab. 1
lists species that occurred at 4% (10 of 241 stations) or
more of all stations sampled.

N2, based on the Shannon-Weaver index and Simpson's

index, resulted in the same pattern of higher diversity in
both the whole of the north and parts of the central/
southern English coast.

Spatial Diversity Pattern in the North Sea

Distribution of Communities

Generally the total number of species was lower in the

southern North Sea than in the central and northern
North Sea (Fig. 2). A maximum of 51 species per haul was
found in the middle of the northern North Sea (48F0).
The lowest number of species (6) occurred at a station
close to the coast of southern Denmark (39F7).
This general spatial trend was found for sessile as well
as free-living fauna. High numbers of sessile species were
found in particular along a corridor in the central North
Sea between 56 ~ and 58~ west of 5~ (Fig. 2A). Few
species were found in the German Bight and southern
North Sea a n d a t several stations no sessile fauna was
recorded. However, low numbers were also found in
several parts of the northern North Sea (e.g. 47F3, 51F1
and 52E9). These species-poor stations were not located
in any one particular region, but were scattered around
the northern North Sea.
The number of free-living species in the northern
North Sea north of 56~ (20.7_+7.6) was about double
that in the southern North Sea (10.9+4.5) (Fig. 2B). Along
the central/southern English coast, however, particularly
high numbers of free-living species, up to 31 per haul,
were found. Other diversity indices, such as Hill's N1 and

Cluster analysis indicated that, similar to species diversity, communities were geographically separated along
a south-north gradient.
For sessile fauna, six clusters were identified (Fig. 3),
with two clusters being restricted to the northern North
Sea, north of 56~ The other four groups were located
in the southern North Sea, with only some stations of
one cluster reaching beyond 56~ Sessile species mainly
responsible for the separation of communities between
north and south are shown in Fig. 5. The most diverse
cluster (cluster 5, Fig. 3) dominated the region between
56 ~ and 58~
Erect or branching species like Hustra
foliacea, Securiflustra securifrons, Alcyonium digitaturn or
Ascidiella scabra were mainly found in this zone and
several sessile species were exclusive to this cluster (e.g.
the branching bryozoans Eucratea loricata and Buskea

dichotorna).
Free-living fauna was separated into three main
clusters (Fig. 4). Clusters one and three were located in
the southern and northern North Sea respectively. A
third group occurred in the centre and formed a transition
zone between north and south. Free-living species mainly
responsible for the clustering ate shown in Fig. 6.

272

Table
1. Epibenthic species in the N o r t h Sea, IBTS 1999. - Species p r e s e n t a t 10 stations (4.1% presence) or
m o r e are listed. - Several species strictly belong to infauna but were recorded because they occurred regularly or
were f o u n d a m o n g erect sessile fauna. Species classified as 'erect species' are marked (*).

Free-living fauna
Ditrupa arietina
Aphrodita aculeata
Nephtys caeca
Hyalinoecia tubicola
Nothria conchylega
Eunice pennata
Pycnogonum littorale
Cirolana borealis
Spirontocaris lilljeborgi
Spirontocaris spinus
Processa canaliculata
Pandalina sp.
Pandalus borealis
Pandalus montagui
Crangon allmanni
Crangon crangon
Pontophilus spinosus
Sabinea sarsi
Nephrops norvegicus
Anapagurus laevis
Pagurus bernhardus
Pagurus prideaux
Pagurus pubescens
Galathea dispersa
Galathea intermedia
Galathea strigosa
Munida rugosa
Ebalia cranchii
Ebalia tuberosa
Hyas araneus
Hyas coarctatus
Macropodia rostrata
Macropodia tenuirostris
Corystes cassivelaunus
Atelecyclus rotundatus
Liocareinus depurator
Liocarcinus holsatus
Tmetonyx cicada
Ampelisca maerocephala
Antalis vulgaris
Nucula nitidosa
Modiolus modiolus
Pseudamussium
septemradiatum
Aequipecten opercularis
Astarte sulcata
Tridonta elliptica

presence
[%]
10.0
34.4

4.1
29.0

11.6
5.8
14.9
7.5

21.6
6.2
5.0
5.4
6.2
49.0
53.9

19.1
26.6
6.6
13.3
54.8
88.8
26.6

41.1
12.0

11.6
13.7
7.9
5.8
7.5

9.1
42.3

19.1
5.4
18.3
8.7
14.1
50.6
4.1

5.0
5.4
8.3
5.8

5.8
18.3

13.7
5.0

Free-living fauna
Acanthocardia echinata
Cerastoderma edule
Spisula elliptica
Ensis ensis
Phaxas pellucidus
Abra prismatica
Arctica islandica
Chamelea gallina
Timoelea ovata
Corbula gibba
Sepiola atlantica
Turritella communis
Aporrhais pespelecani
Velutina velutina
Euspira catena
Polinices pulchellus
Buccinum undatum
Neptunea antiqua
Colus gracilis
Colus islandicus
Colusjeffreysianus
Scaphander lignarius
Henricia oculata
Asterias rubens
Leptasterias muelleri
Ophiothrix fragilis
Ophiopholis aculeata
Amphiura chiajei
Ophiura albida
Ophiura ophiura
Ophiura sarsi
Psammechinus miliaris
Echinus acutus
Echinus elegans
Strongylocentrotus
droebachiensis
Spatangus purpureus
Echinocardium cordatum
Echinocardium flavescens
Aslropecten irregularis
Brissopsis lyrifera
Luidia sarsi
Hippasteria phrygiana

presence
[%]
8.7
6.6
8.7
4.6
10.8
7.9
4.6
8.3
5.4
6.2
17.4
12.9
16.6
6.6

7.1
6.6
44.4
43.2
45.6
7.9
5.0
12.4

5.4
74.3

9.1
17.0
8.3

4.1
27.8
42.3
6.2
18.3
25.3
5.8
4.6
20.7
23.7
22.8
75.1

9.1
29.0

4.1

Sessile fauna

presence
[%]

Suberites ficus*
12.9
Suberites pagurorum
14.9
Phakellia ventilabrum*
4.6
Alcyonium digitatum*
22.8
Pennatula phosphorea*
16.2
Epizoanthus incrustatus* 31.1
Bolocera tuediae*
14.9
Urticina felina*
7.1
Urticina eques*
4.6
Hormathia digitata
25.3
Actinauge richardi*
5.0
Adamsia carciniopados
I 1.6
Caryophyllia smithii*
6.2
Tubularia indivisa*
12.9
Hydractinia echinata
41.1
L afoea dumosa*
19.1
Halecium beanii*
4.6
Nemertesia antennina*
5.8
Nemertesia ramosa*
5.4
Abietinaria abietina*
6.6
Diphasia attenuata*
5.8
Hydrallmania falcata*
24.5
Thuiaria thuja*
7.1
Sertularia cupressina*
4.1
Thelepus cincinnatus*
24. I
Hydroides norvegica
14.1
Pomatoceros triqueter
6.6
Serpula vermicularis
10.8
Balanus balanus
8.7
Balanus crenatus
10.4
Verruca stroemia
10.8
Anomia ephippium
18.3
Hiatella arctica
10.0
Alcyonidium diaphanum* 14.1
Alcyonidium parasiticum I 1.6
Tubulipora liliacea
7.5
Turbicellepora avicularis* 6.2
Eucratea lorieata*
7.5
Electra pilosa
17.0
Flustra foliacea*
32.4
SecurijTustra securifrons* 10.4
Amphiblestrum flemingii
4.1
Dendrobeania murrayana* 6.2
Corella parallelogramma* 17.0
Ascidiella aspersa*
4.6
Ascidiella scabra *
23.2

273

A. Sessile
4 a

2 ~

0 ~

fauna

2 ~

4 ~

B. Free-living
6 ~

8 ~

E6

E8

FO

fauna

F2

F4

F6

F8

62 ~

52

~-..~o-,.

61 ~

~-~ 9

60 ~

Oo

~~,.~.

~e

50

48

59 ~

46
c)
m
4 4 oo

58 ~

57 ~

42 ~"

"O

56 ~

40 ~

55 ~

38

54 ~
36
53 ~
34
52 ~
32
51E6

E8

FO
ICES

F2

F4

F6

F8

4~

2 ~

rectangles

Several species did not occur throughout the North

Sea but were restricted to defined geographical regions
(Figs. 5, 6). The spatial distribution patterns could be
divided into three categories.
a. Species limited to an area north of the 50 m contour:
Suberites pagurorum, Epizoanthus incrustatus (= E.

papillosus), Hormathia digitata, Thelepus cincinnatus,

Verruca stroemia, Pandalus rnontagui, Pontophilus spinosus, Hyas coarctatus, Anapagurus laevis.
b. Species limited to an atea north of the 80 m depth
contour: Hyalinoecia tubicola, Pagurus prideaux.
c. Species predominantly occurring south of the 100 m
contour: Electra pilosa, Ophiura ophiura.

Alcyonium digitatum was regularly recorded in the

central and southern North Sea, but not in the German
Bight (Fig. 5). No species were restricted to the southern
North Sea, but brittle stars as well as Asterias rubens and
Liocarcinus holsatus were found here in higher numbers.
The distribution of several sessile species was determined
by their free-living host, e.g. the distribution of Hydractinia echinata depended on Pagurus bernhardus and
Adamsia carciniopados on Pagurus prideaux.

2"

4"

6 ~

8 ~

Iongitude

Fig. 2. Spatial distribution of species diversity [numbers of species haul 1l'

species.

D i s t r i b u t i o n of I n d i v i d u a l Species

0 ~

--

A. Numbers of sessile species; B. Numbers of free-living

Beam-Trawling Effort and Environmental

Factors

Beam-trawling was recorded in the central and southern North Sea, but intensive beam-trawling efforts were
confined to areas close to the German, Dutch and
Belgium coast (Fig. 7). Up to 60,000 beam-trawling hours
per ICES rectangle per year were recorded in 1998. In
half of the fished ICES rectangles, beam-trawling effort
was less than 5,000 hours per year.
Numbers of sessile species were negatively correlated
with beam-trawling effort (Fig. 8A). Restricting the list of
sessile fauna to erect species, which potentially improve
the structural complexhy of the habitat, also showed a
negative correlation with beam-trawling effort (r=-0.262,
n=69, p<0.05; species included in this analysis are marked
in Tab. 1). No correlation was found between numbers of
free-living species and beam-trawling effort (Fig. 8B), but
they were positively correlated with depth (Fig. 8D) and
latitude. Sessile fauna, in contrast, was not correlated with
depth (Fig. 8C). The total species richness did not vary
significantly between different types of sediment (Fig. 9).
Ahhough the mean number of sessile species was higher
in areas with coarser sediment, this trend was also not
statistically significant.

274

Sessile fauna

ctuster I
mm

cluster 2
~ 0

Q 9

mo

9
o

cluster 3

9
m

~7 O

9 9

~7

9
o

Oo

cluster 4
& 9

,~

[3

~7

r-1 0

cluster 5

cluster 6

0 cluster 1

[ ] cluster 2

~ cluster 3

9 cluster 4

9 cluster 5

9 cluster 6

~, outlying sites

10 20 30 40 50 60 70 80 90 100
Bray-Curtis similarity
Sampling latitude:

O51 ~
~ []53%55*
9 57 o- 59 ~ 9 59*- 62 ~

X55 ~

Fig. 3. Dendrogram and spatial distribution of c]usters of sessile epibemhos. - The grouping is based on hierachical clustering of 73 sites (English survey), using group-average linking of Bray-Curtis similarities calculated on
presence/absence data.

Discussion
Epibenthic surveys in the North Sea, on the spatial
scale and with coverage presented by this study, ate
scarce. Former studies h a d a similar spatial range, but
less than half the number of stations were sampled
(FRAUENHEIM et al. 1989; DxrR et al. 1983; J~NNINCS et al.
1999a). A clear separation between communities in the
south-eastern North Sea and the central-northern North
Sea, roughly along the 50 m depth line, was found for
free-living as well as sessile fauna. This coincides with
former descriptions of epibenthic communities in the
North Sea (GLi~MA~Er 1973; DXE~ et al. 1983; Fv.At3~I~}trI~ et al. 1989).
The separation of the faunal communities was based:
1. on the overall higher number of species inhabiting
central and northern parts of the North Sea and

2. on a change in species composition from north to

south.
Little information is published on epibenthic diversity
patterns in the North Sea, bnt a trend towards higher
numbers of species from south to north was mentioned
by FRAUENHEIM et al. (1989) and DYER et al. (1983).
Infauna diversity increased along the same gradient (HEIv
et al. 1992; HEIV & CV,ArYM~ERSr 1995). However,
diversity of sessile species did not increase steadily, but
was highest between 56 ~ and 58~ west of 5~ This
coincides with the area between the 50 m and 100 m
contour lines (LEE & RAMSTER 1981). These depth lines
are known to be boundaries between epibenthic communities (GL~MaREC 1973; DYER et al. 1983; BASrORD et al.
1989), but our results suggest that sessile species in par-

275

Free-living fauna

9 9

.#

--_
clusterl

---k

~o

:ii-

!~

f!

,-

nmm 9

o. ~ T i /

L~.o......~ #
cluster 2

"~.

",oooook~

~; ~o oooo~
"

!i

:|

cluster 3

~~

0 cluster 1 9 cluster 2
outlying sites
10

20

30

40

-~

50

60

9 cluster 3

370

80

90 100

Bray-Cu~is similarity

Sampling latitude:
C) 51~
9

57 ~ 59 ~

Vq53 ~ 55 ~
9
~ 62 ~

X 5 5 ~ 57 ~

Fig. 4. Dendrogram and spatial distribution of clusters of free-living epibenthos. - The grouping is based on
hierachical clustering of 73 sites (English survey), using group-average ]inking of Bray-Curtis similarities calculated
on 44-transformed abundance data.

ticular are responsible for changes in the epibenthic community. The sessile community dominating this central
zone was more diverse than others, and several bryozoans, anthozoans and ascidians were either found exclusively or more frequently in this area. Spatial distribution
patterns of Hustra foliacea, Securiflustra securifrons and
to some extent Alcyonium digitatum indicate that these
erect species predominantly inhabit this central corridor
between 56~ and 58~ which coincides with patterns
shown by BAS;ORD et al. (1989). Similar to other habitat
structuring species they possibly have a promoting effect
on species diversity (RIEsEN & REISE 1982; DAUER et al.
1982; BARRV& DAWrON 1991) or at least actas indicator
for areas of higher diversity. Infauna diversity, for example, was also found to peak between 56~ and 58~

(overall infauna: HElv et al. 1992; Mollusca: HEW &

CtL*EVMEERSCH 1995). However, free-living epibenthos
was not more diverse in this zone.
Early descriptions of the distribution of E foliacea, A.
digitaturn and Suberites ficus indicate that these species
were not always scarce in the SE North Sea (ARNDT 1928;
PAX 1928; BORG 1930). PAx (1928) calculated that about
10,000 kg of A. digitatum were caught during 20 trips of a
fishing vessel in the German Bight and even in the early
eighties DYER et al. (1983) found A. digitatum at several
stations in the southern North Sea. E foliacea was an
abundant species in the German Bight at the beginning
of the last century according to BORC (1930), but about
20 years ago DVER already recorded little E foliacea in
the SE North Sea.

276

~,~~

Suberites pagurorum

j;~

II

Alcyonium digitatum

"~"

%./

Pennatula phosphorea

.;.

.-~

Hormathia digitata

I~~~'~:.::.~.~l

~
~-"

i jjll:,l,,,Sl

Adamsia carciniopados

.~-.. ~ .
o0 ~ ~ t ~ ' - J

~~

7~.S

..~'i~
~ . * . " "i~~.
Epizoanthus incrustatus

~ I

Hydractinia echinata

~"

"" % /

Hydrallmania falcata

#. o.o. .

~o

1~0" 0 0

~.

"..'.'c

Thelepus cincinnatus

Hydroides norvegica

Balanus crenatus

~JL: ~
~:.:~~
..,-..~,~

~~:~

~>'-..:...~A l ' ~ - i " - ~ '

Electra pilosa

Flustra foliacea

Verruca stroemia

~ " " " ~ ~J

~~'"~-

Securiflustra securifrons

Ascidiella scabra

Fig. 5. Distribution of sessile species mainly responsible for the separation between communities in the southern
and central-northern North Sea. - Species were determined by the program SIMPER of the P R I M E R package.

277

+" + +--++".~++i-,~ ~+89

~'-~,'-.

_,

? .. ,. _ , ~

++,,_,..-..

'+-

Hyalinoecia t u b i c o l a

~,.~we~,

k?

~-.i

, -

" ~

J+'~

.++....+~

Pandalusmontagui

+'-;'Z', ~+ /

~--+-T_.~II" I "

'

9 +~

...

,,"

Pontophilus
spinosus

,+~

.e
I-

Crangon a l l m a n n i

R'~q" .,~

,,+-+,I

I ~ln:':7:_.,~r'+

l~':;"::"

~.t

~~"'-" ~

k ~:~ ;~=i'..: ~I

~ ~ "

Anapagurus laevis

Pagurus bemhardus

Pagurus pubescens

9 9 ~

9 o~

~.. '.,

ii",.~~
9 o,

~'_ :~
i~~.

I ~,::'"

~
Hyas coarctatus

~ /

~...

~.:~--r
~.Z""
" :"~~.'~
-'."

~-

,:,,.~

-~.~

~ h ~-

Liocarcinus holsatus

Asteriasrubens

Astropecten irregularis

Ophiura albida

Echinus a c u t u s

Echinocardium
cordatum

~'.!~".ii~'il J
Ophiura ophiura

Fig. 6. Distribution of free-living species mainly responsible for the separation between communities in the
southern and central-northern North Sea. - Species were determined by the program SIMPER of the PRIMER
package. The sizes of the circles indicate abundances (low, medium, high).

278

61 ~

,i

i
~o.;,

L__'.~___ L~ _

~0"' _~~.

'~,
_

p.

.
58"

57"

.=a~"

i
t

9 I

o;o

,(
,i

56"

~~"

54"

....

Dl

'

~~

9 I )lo
i~~

i~~ 7~i''

aD=

oio

51 9 ,-

II

5,000 h

25,O0O h

60,000 h

50 ~

4"

2"

0"

2"

4"

6"

8"

Fig. 7. Beam trawling effort 1998 [h ICES-rectangle -1 year-l].Accumulated data from The Netherlands, Germany and
England.

Information on the distribution and ecology of epifaunal species is scarce, but it appears that similar to our
study species like Hyas coarctatus, Epizoanthus incrustatus, Hyalinoecia tubicola and Thelepus cincinnatus were
found mainly in the northern North Sea during past
surveys (WALTON 1907; HARTMANN-ScHRODER 1996;
INGLE 1996; B(DGGEMANN 1998). Their distribution may
be determined by temperature of prevailing currents.
The epibenthic community in the southern North Sea,
excluding the English coast, was determined by freeliving, scavenging species like Ophiura spp., Asterias
rubens, Pagurus bernhardus and Liocarcinus holsatus. The
range of species coincides with the 'Southern North Sea
sandy'- group described by REES et al. (1999) as well as
FV,A, ENHEIM et al. (1989) and JENNINGS et al. (1999a).
Regarding potentiaI factors ruling diversity on a
large scale, correlations were found with depth and beamtrawling effort. Depth and sediment characteristics have
been described as the main environmental factors influencing epibenthic communities (BAsFOR, et al. 1990; REES
et al. 1999). In this survey we did not find significant
variations in species diversity related to sediment type and
former epibenthos studies report contrasting results regarding the importance of sediment. REES et al. (1999)
regard sediment type as the main structuring force for

the epifauna, while DOINEVELI3 et al. (1991) assume an

'obscure or lacking relation between the organisms and
the sediment type'. For this study, sediment categories
were derived from maps published by BASFORD et al.
(1993), which might not always be detailed enough and
hence might have affected the analysis.
Depth, reflecting differences in temperature and food
availability, and diversity of free-living fauna was positively correlated. This agrees with most other epibenthic
studies in the North Sea (e.g. FRAUENHEIM et al. 1989;
BASVORD et al. 1990; JZNNINGS et al. 1999a). However,
sessile fauna was not correlated with depth. For the SE
North Sea H m z (2000) found a significant correlation
between fauna and both temperature and depth.
Beam-trawling was concentrated a]ong the continental
coast. Highest values of over 60,000 hrs trawling were
estimated for single ICES rectang]es in Dutch and
German waters. Fishing effort data was available from
main beam-trawling countries, namely The Netherlands,
Germany and England, but data from Belgium and Denmark are missing. Hence, beam-travAing pressure might
be higher than indicated in some areas. Our estimates of
beam-trawling in the southern North Sea also ignore the
horse power of trawlers, which certainly has a considerable effect on the impact of trawling on the benthic fauna.
However, the distribution of beam-trawling effort we
estimated for 1998 is similar to accounts of JENNINGS et al.
(1999b).
The total number of sessile species, as well as the
number of erect, branching sessile species, was negatively
correlated with hours of beam-traw]ing carried out per
year within the respective ICES rectangle. Free-living
fauna, on the other hand, did not show any correlation
with beam-traw]ing, but the dominance of scavengers in
the southern North Sea indicates high levels of physical
disturbance (KAIsEI~ et al. 2000). The results suggest that
beam-trawling may affect the distribution of sessile fauna
on a large spatial scale. The absence of A. digitatum and
the bryozoan E foliacea in most parts of the SE North
Sea but their presence in unfished areas like the
'Steingrund' (KOHNE & RACHOR 1996) supported this
hypotheses. Several studies found changes in benthic communities related to trawling (eg. FHI) & HALL 1999;
RAMSEY et al. 1998; KAISER & SVENCER 1996; BERGMAN &
HUP 1992; several reports in LINnEBOOM & DEGROOT
1998) and sessile species have been shown to be particularly vulnerable to fishing gear towed over the seafloor
(FREESE et al. 1999; KAISER et al. 1998, 2000). They
become detached from their settlement substratum and,
unlike free-living fauna, most sessile species do not have
the ability to disperse and colonise elsewhere after detachment. It can not be ruled out that beam-trawling removes
some of the species creating habitar structure, which
would thereby negativeiy affect associated organisms
(RIEsrN & REIS~ 1982; SAINSBUXX 1987; LANGTON &
ROBINSOiX 1990; COLLIE et al. 1997). However, such
interpretation has to be regarded with some caution,
because (i) other environmental factors might be responsible for the variation in sessile species, (ii) fishing vessels
may be actively avoiding areas with ah abundant sessile
fauna and (iii) fishing effort can vary greatly within any
one ICES rectangle (RIJNSI)ORV et al. 1998), hence the
spatial scale of this analysis may be too coarse.

279

Beam-trawling

2O

3O

la 9

Sessile fauna
* negaUve
correlation,

14
12

10

"6
o

l i

8oo~e

25

16 9

R
ce

Beam-trawling

9~
p<0.05

9
O

Free-living fauna

~ "]..~ "

..:

= 0'F':'~1"

"

9
O9

2 |
O

m_

wo 9

9 9
9

:~~

20000

60000

40000

Depth

Sessile fauna

Depth

Free-living faur~ 9

9 9 9

25

16

no correlation

14

'~

12

`6
o

60000

30

20
18

,==

40000

20000

fishJng effort [h ICES rectanglos "~]

fi$hing effort [h ICES rectangles -t]

no correlation

20 9

* ~siUve
correlation,

p<O.05

9 9 9

==
15;

O0

10
8

OO
9 N
9
9

~
O0
9 OO N

15

9 o,:t,:,i, :.

,.:o~o%,

o% 8 ~

5
N O

40

50

0
0

10

20

:30

60

depth [m]

10

20

30

40

50

60

depth [mi

~oo',

o 9 ~

~J

~ 1 4 9o

Fig. 8. Relationship between species diversity and fishing effort (sum of hours beam-trawling per ICES rectangle and year) and
depth. - (A)-(D). Product Momem Correlation Coefficient r, rc05~21.67=0.237: (A) r=-0.253, p<0.05; (B) r=0.021; (C) r=0.025; (D)
r=0.391, p<0.05; (E) Location of sampling stations considered for the analysis (<50 m depth).

In the near future more detailed fishing effort maps,

based on satellite tracking of fishing vessels, will become
available. This will allow a more reliable analysis of the
relationship between fishing effort and benthos and will
clarify to what extent fishing might contribute to the
spatial variation in diversity of epibenthos in the N o r t h
Sea,

Acknowledgements
We are grateful to the various governmental agencies and
fishery institutes which have hosted sampling personnel on their
vessels during their groundfish surveys, or otherwise co-operated
with this project. E[KE RACHOR and HUBE8T REES gave valuable
comments on the first version of the manuscript. This study was
funded by the European Community (DG IVX, 98/021).

280

3o

sessile species

all species

25

12,

"~
~

2o

10,

"15

8'

4,
c
2

(5)

(62)
(130)
(211
Sediment type
(number of stations)

mll
2

(7)

(5)

(62)
(130)
(21)
Sediment type
(number of stations)

(7)

Fig. 9. Species richness (mean

in relation to sediment types (according to BASFORD et al.,
1993). - 2: coarse sand; 3: medium sand; 4: fine sand; 5: coarse sih; 6: medium silt. Differences
between sediment types were not significant (ANOVA, TUKEYtest).

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Submitted: 10 November 2000

Reviewed: 5 March 2001
Accepted: 22 May 2001