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I<br />

BULLETIN DE L'INSTITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE<br />

BULLETIN VAN HET KONINKLIJK BELGISCH INSTITUUT VOOR NATUURWETENSCHAPPEN<br />

<strong>Species</strong> <strong>composition</strong> <strong>and</strong> <strong>diversity</strong> <strong>of</strong> <strong>free</strong>-<strong>living</strong> <strong>Plathelminthes</strong><br />

(Turbellaria) from s<strong>and</strong>y beaches at the Belgian coast<br />

P. JOUK & E. SCHOCKAERT<br />

Abstract<br />

Ten s<strong>and</strong>y beaches at the Belgian coast were sampled monthly on a<br />

qualitative basis. Six <strong>of</strong> them were also sampled quantitatively every<br />

two months. In total, 130 species were encountered, 38 % <strong>of</strong> them<br />

occur only in I or 2 localities, 32 % have a more general distribution (in<br />

3 to 7 localities) <strong>and</strong> 30 % are found in at least 8 localities. Schizorhynchia<br />

generally fonn the dominant taxon in number <strong>of</strong> species,<br />

followed by the Proseriata. In Knokke <strong>and</strong> Het Zwin, equal numbers<br />

<strong>of</strong> species were found for both taxa.<br />

Mariakerke, Bredene, Knokke <strong>and</strong> Het Zwin have the most diverse<br />

turbellarian fauna's, with the lowest dominance ratio's. In the semiexposed<br />

beach <strong>of</strong> Oostende the fauna is slightly less diverse, dominated<br />

by a few species, while the sheltered beach in Heist clearly has the least<br />

diverse fauna, strongly dominated by one species, Archilopsis arenaria.<br />

Keywords: <strong>free</strong>-<strong>living</strong> <strong>Plathelminthes</strong>, Belgian coast, species richness,<br />

<strong>diversity</strong>.<br />

Samenvatting<br />

Tien z<strong>and</strong>str<strong>and</strong>en aan de Belgische kust werden ma<strong>and</strong>elijks bemonsterd<br />

op kwalitatieve basis. Zes ervan werden ook tweema<strong>and</strong>elijks<br />

kwantitatief bemonsterd. In totaal werden 130 soorten aangetr<strong>of</strong>fen,<br />

waarvan 38 % maar in I <strong>of</strong> 2 localiteiten voorkomen, 32 % een meer<br />

algemene verspreiding hebben (in 3 tot 7 str<strong>and</strong>en) en 30 % in minstens<br />

8 localiteiten aangetr<strong>of</strong>fen worden. Schizorhynchia vonnen doorgaans<br />

het grootste taxon in aantal soorten, gevolgd door de Proseriata. In<br />

Knokke en Het Zwin worden gelijkaardige aantallen soorten van beide<br />

taxa gevonden.<br />

Mariakerke, Bredene, Knokke en Het Zwin hebben de meest diverse<br />

turbellariënfauna's, met de laagste dominantie verhoudingen. Het<br />

semi-geëxposeerde str<strong>and</strong> van Oostende heeft een iets minder diverse<br />

fauna, die gedomineerd wordt door enkele soorten. Het beschutte<br />

str<strong>and</strong> van Heist heeft duidelijk de minst diverse fauna, die sterk<br />

gedomineerd wordt door één soort, Archilopsis arenaria.<br />

Trefwoorden: vrij levende <strong>Plathelminthes</strong>, Belgische kust, soortenrijkdom,<br />

diversiteit.<br />

Introduction<br />

Free-<strong>living</strong> <strong>Plathelminthes</strong> <strong>and</strong> other s<strong>of</strong>t-bodied mei<strong>of</strong>aunal<br />

taxa are <strong>of</strong>ten disregarded in bio<strong>diversity</strong> studies or<br />

numbers are highly underestimated when they have been<br />

considered. In most bio<strong>diversity</strong> surveys, fixed samples<br />

- - -- ---<br />

BIOLOGIE, 72-SUPPL.: 35-41, 2002<br />

BIOLOGIE, 72-SUPPL.: 35-41, 2002<br />

are used, while the s<strong>of</strong>t-bodied fauna should be studied<br />

alive to allow proper identification (after which they can<br />

be recognised in fixed samples). Data on species <strong>diversity</strong><br />

<strong>of</strong> <strong>Plathelminthes</strong> are very rare: H' values <strong>of</strong> 1.20-3.35<br />

were recorded in beaches <strong>of</strong> the German Isl<strong>and</strong> <strong>of</strong> Sylt<br />

(REISE1983a, b, 1984; WEHRENBERG1988) <strong>and</strong> <strong>of</strong> 2.55<br />

maximum in mangrove s<strong>and</strong>flats <strong>of</strong> northem Queensl<strong>and</strong>,<br />

Australia (DlTIMANN1991). MARTENSel al. (1985) <strong>and</strong><br />

MARTENS& SCHOCKAERT (1986) stated that turbellarian<br />

<strong>diversity</strong> can be <strong>of</strong> the same magnitude as that <strong>of</strong> nematodes<br />

or harpacticoids, <strong>and</strong> that turbellarians may represent<br />

up to more than 25 % <strong>of</strong>the mei<strong>of</strong>aunal density. The<br />

data presented here are in the same order <strong>of</strong> magnitude.<br />

A general species list <strong>of</strong> the Belgian coast <strong>and</strong> adjacent<br />

areas, listing about 250 species, is given by SCHOCKAERT<br />

el al. (1990). In this contribution, detailed species lists are<br />

given for ten s<strong>and</strong>y beaches at the Belgian coast. <strong>Species</strong><br />

<strong>composition</strong>, efficiency <strong>of</strong> specific extraction methods<br />

for s<strong>of</strong>t-bodied mei<strong>of</strong>auna <strong>and</strong> <strong>diversity</strong> <strong>of</strong> the turbellarian<br />

populations are discussed.<br />

Material <strong>and</strong> methods<br />

In ten localities along the 64 km long Belgian coast, live<br />

samples from the s<strong>and</strong>y beaches have been investigated<br />

monthly from April 1985 till June 1986 (see table 2). In<br />

six <strong>of</strong> these localities, every two months, 2 cores <strong>of</strong> 10 cm2<br />

(40 cm deep) were taken <strong>and</strong> fixed for quantitative analysis,<br />

<strong>and</strong> one for sediment analysis. Samples were taken<br />

in the medio-litoral zone, where high numbers <strong>of</strong> individuais<br />

<strong>and</strong> <strong>of</strong> species are to be found (JOUKel al. 1988).<br />

Living Turbellaria are extracted from the s<strong>and</strong> by the<br />

MgClz-decantation method (HULINGS& GRAY 1971,<br />

MARTENS1984, see SCHOCKAERT 1996), identified under<br />

the compound microscope <strong>and</strong> mounted in lactophenol<br />

for later comparison. Fixed samples are extracted with the<br />

Bamett-method (BARNETI1968, MARTENS1984), aUflatworms<br />

mounted in lactophenol <strong>and</strong> identified.


36 P. lOUK & E. SCHOCKAERT<br />

Table I - Sedimentcharacteristics<strong>and</strong> expositionper locality.<br />

* S<strong>and</strong>:<br />

Results <strong>Species</strong> <strong>composition</strong><br />

The /ocalities <strong>and</strong> the sediment<br />

The s<strong>and</strong> become coarser from west to east (see table 1).<br />

Roughly spoken, there is a gradient in the sediment from<br />

west to east, with the coarse <strong>and</strong> less well sorted s<strong>and</strong> in<br />

the eastern localities. The most sheltered locality Heist<br />

however has the finest s<strong>and</strong> <strong>and</strong> the highest silt content <strong>of</strong><br />

the ten localities. The relatively sheltered beach <strong>of</strong> Oostende<br />

("Klein Str<strong>and</strong>") has finer s<strong>and</strong> than the neighbouring<br />

localities. Except for the two most western (very<br />

wide) beaches <strong>and</strong> the most eastern locality, jetties <strong>and</strong>/or<br />

piers have been built on the beaches. If sediment characteristics<br />

reflect the degree <strong>of</strong> exposure, it can be said<br />

that Het Zwin <strong>and</strong> Knokke are the most exposed beaches,<br />

which can indeed be confinned by (subjective) observation<br />

<strong>of</strong> the sites.<br />

De Panne St.-Idesbald Mariakerke Oostende Bredene<br />

- mean median 0.200 0.211 0.215 0.208 0.234<br />

partic1e size (mm)<br />

- c1assification fine fine fine fine fine<br />

- sorting very well very well to very well very well very well to<br />

less well well<br />

* mean silt % 0.04 0.04 0.01 0.03 0.01<br />

* mean s<strong>and</strong> % 098 96.54 96.51 91.56 099<br />

* mean gravel % 2.45 3.42 3.48 8.39 1.19<br />

* S<strong>and</strong>:<br />

Blankenberge Zeebrugge Heist Knokke Zwin<br />

- mean median 0.243 0.226 0.197 0.332 0.316<br />

partic1e size (mm)<br />

- c1assification fine fine fine medium medium<br />

- sorting very well to very well to very well to less well less well<br />

well well well<br />

* mean silt % 0 0.02 0.16 0.02 0.02<br />

* mean s<strong>and</strong> % 099 099 99.01 97.35 88.57<br />

* mean gravel % 0.75 0.77 0.84 2.36 11.41<br />

In total, about 130 species <strong>of</strong> <strong>free</strong>-<strong>living</strong> <strong>Plathelminthes</strong><br />

were encountered (tabie 2). Within the Acoela, only<br />

Paratomelia rubra was identified to species level. At<br />

least 15 other species were present, but not identified<br />

(sectioned material necessary for species identification).<br />

For the same reason, the three Catenulida-species were<br />

only identified to family level.<br />

Based on the live samples, 108 species (Acoela <strong>and</strong><br />

Catenulida not inc1uded) were found throughout the survey.<br />

Generally, 50 to 60 species were recorded per 10cality.<br />

In Heist, only 20 species were found. After study<br />

<strong>of</strong> the fixed material, only 3 additional species were<br />

added to the overall list. However, there is a significant<br />

increase from 10 to 50 % <strong>of</strong> the number <strong>of</strong> species per<br />

locality (table 3): 10-14 % for Mariakerke, Oostende <strong>and</strong><br />

Bredene, 19 <strong>and</strong> 29 % resp. for Knokke <strong>and</strong> Het Zwin <strong>and</strong><br />

--- -- -- - - - - - - - -


<strong>Species</strong> <strong>composition</strong> <strong>and</strong> <strong>diversity</strong> <strong>of</strong> <strong>free</strong>-<strong>living</strong> <strong>Plathelminthes</strong> (Turbellaria) 37<br />

Table 2 - <strong>Species</strong> list per locality. X: present in live samples (10 localities); 0: present in fixed samples (6 localities).<br />

Acoela<br />

IJePanne S-Idesbald Mariakm, lmIende BI


38 P. JOUK & E. SCHOCKAERT<br />

*Kalyptorhynchia<br />

**Eukalyplorhynchia<br />

Gnathorbynchidae<br />

Carcharodagnalhus slil<strong>of</strong>er (Schilke, 1970)<br />

X<br />

Gnalhorhynehus conocaudatus Meixner, 1929 X X XO X X X X XO<br />

Psillaeorhynehus verweyi den Hartog, 1968<br />

XO<br />

UneinorhynehusfIavidus Karling, 1947<br />

- Cicerinidae<br />

0 XO<br />

Cicerina brevieirrus Meixner, 1928<br />

Cicerina remanei Meixner, 1928 X X XO XO XO x x<br />

X<br />

XO<br />

XO<br />

XO<br />

X<br />

XO<br />

Cieerina lelradaetyla Giard, 1904 X X X X XO XO<br />

Paraeieerina delloides Martens & Sehockaert, 1981 X X X XO X XO<br />

Ptyalorhynehus eaeeus Meixner in Ax, 1951<br />

- Polyeystidae<br />

X X 0 XO XO X X XO XO<br />

Neopolycyslis lridenlala Karting, 1955 X X X X<br />

Polycyslidarum sp.<br />

- Cystiplanidae<br />

X X<br />

Nigerrhynehus opislhoporus Schilke, 1970<br />

- Psammorhynehidae<br />

XO<br />

Psammorhynehus tubulipenis Meixner, 1938 X X 0 X X X XO XO XO<br />

- <strong>Species</strong> incertae eedis<br />

Elver/ia krusei Noldt, 1989<br />

**Schizorhyochia<br />

X<br />

- Sehizorhynehidae<br />

Amphirhynehus caudolus Sehilke, 1970<br />

0<br />

Carcharodorhynehus isolalus Schilke, 1970 X X XO XO XO X 0<br />

Careharodorhynehus lislensis Sehilke, 1970 X 0 X<br />

Limirhynehus danicus Schilke, 1970 X X XO XO XO X X X<br />

Neosehizorhynehus brevipharynggus Sehilke, 1970 X XO X 0 X X<br />

Neosehizorhynehus longipharynggus Sehilke, 1970 X X XO XO XO X X X<br />

Neosehizorhynehus parvoroslro Ax & Heller, 1970 X X XO XO XO X X XO XO<br />

Neosehizorhynchus sp. X X XO XO XO X X 0 0<br />

Parasehizorhynehoides gl<strong>and</strong>ulis Schi1ke, 1970 X XO<br />

Prosehizorhynehus bivaginatus Schilke, 1970 X X XO XO X X 0 0<br />

Prosehizorhynehus helgol<strong>and</strong>ieus I'Hardy, 1965 X X XO 0 XO X X 0<br />

Prosehizorhynehus gullmarensis Karting, 1950<br />

XO<br />

Prosehizorhynchus /TiduelibusSehilke, 1970 X X XO XO XO X X<br />

Schizochilus ehoriurus Boaden, 1963 X X XO XO XO X X XO XO<br />

Schizochilus marcusi Boadeu, 1963 X X XO XO XO X X XO<br />

Schizorhynehoides aculeatus rHardy, 1963<br />

X<br />

Sehizorhynehoides karlingi Martens & Sehockaert, 1981 X X X XO<br />

Schizorhynehoides symmelricus Martens & Sehockaert, 1981 X XO XO XO X XO<br />

Thylaeorhynehus ambronensis Schilke, 1970 X X XO XO XO X X XO XO<br />

Thylaeorhynehus caudalus Meixner, 1928<br />

X<br />

Thylacorhynehus eonglobalus Meixner, 1928 X X XO XO XO X X 0 0<br />

Thylaeorhynehuspyriferus Karling, 1950<br />

XO<br />

Sehizorhynehoidarum sp.<br />

Nernatorbynehidae<br />

XO<br />

Nematorbynehus parvoacumine Schilke, 1969<br />

- Karkinorbynehidae<br />

X<br />

Cheliplana baadeni Schilke, 1970 X X X X X XO XO<br />

Cheliplana gemmifera Noldt, 1989 X XO X XO X X<br />

Cheliplana marcusi (Karting, 1956) X X 0 X 0 0<br />

Cheliplana microcirrus Noldt, 1989<br />

XO<br />

Cheliplana remanei (Meixner, 1928) X X X XO X X X<br />

Cheliplana stylifera Karling, 1949 X X XO XO XO X X XO XO 0<br />

Cheliplanilla eaudala Meixner, 1938 X X XO XO XO X X 0 XO 0<br />

Karkinorhynehus bruneli Schilke, 1970 X X XO XO X X X X<br />

Karkinorhynehides purpureus Sehilke, 1970 XO X 0 XO<br />

Diascorbynehidae<br />

Diascorbynehus rubros Baaden, 1963<br />

*Dalyellioida<br />

Provortieidae<br />

X X XO XO XO X X XO XO<br />

Hangelhellia ealeeifera Karting, 1940<br />

X<br />

Pagaina kinnei Ax, 1970 X X XO XO XO X X XO XO XO<br />

Pogaina nolons (Ax, 1951) X X 0 0 XO X<br />

Provortex tubiferus Lutber, 1948<br />

XO<br />

Provorlex cf psammophilus Meixner in Ax, 1951<br />

Graflillidae<br />

X X X 0 XO X X X XO XO<br />

Bresslauilla reliela Reisinger, 1929 X X<br />

Dolyellioida sp. 1 X X XO 0 XO X 0 XO 0<br />

Dolyellioida sp. 2<br />

X<br />

Dolyellioida sp. 3<br />

X<br />

Dalyellioida sp. 4 X X XO XO XO X 0<br />

Dolye/lioida sp. 5 XO 0<br />

Dolyellioida sp. 6 X 0 0<br />

Dalyellioida sp. 7 X 0 XO 0 0<br />

--


- -<br />

<strong>Species</strong> <strong>composition</strong> <strong>and</strong> <strong>diversity</strong> <strong>of</strong> <strong>free</strong>-<strong>living</strong> <strong>Plathelminthes</strong> (Turbellaria) 39<br />

Table 3 - Number <strong>of</strong> species per locality <strong>and</strong> taxon (X: present, not identified to species level).<br />

De Sinl. Marial:e!te Oostende Btedene Blanken- Ztt- Heist Knokke HdZwin<br />

Panne ldesbald bag< brugge<br />

Live Live Live Fixed Live Fixed Live Fixed Live Live Live Fixed Live Fixed Live Fixed<br />

Acoela X X X X X X X X X X X X X X X X<br />

Catenulida X X X X X X X X X X X X 0 0 X X<br />

Macrostomida 4 4 3 3 3 3 3 3 4 5 I 3 3 3 3 4<br />

Prolecithophora 1 I I 1 1 1 1 I I I 0 0 I 1 0 0<br />

Proseriata 11 14 11 IJ 9 10 10 11 11 10 3 4 17 19 18 18<br />

Rhabdocoela<br />

*Typhloplanoida 4 6 7 7 5 5 6 7 6 6 4 6 12 IJ 9 15<br />

*Kalyptorhynchia<br />

uEukalyptorhynchia 7 5 4 6 6 7 4 4 6 9 6 6 8 8 5 5<br />

**Schizorhynchia 18 26 24 24 18 19 22 25 20 22 1 3 15 21 11 17<br />

*Dalyellioida 6 7 4 5 4 8 4 6 5 4 5 8 3 5 3 4<br />

Total 51 63 54 59 46 53 50 57 53 57 20 30 59 70 49 63<br />

50 % for Heist. These results show that the MgChdecantation<br />

method is clearly less efficient than the Barnett<br />

method, <strong>and</strong> that it has a better efficiency in fine,<br />

clean, (very) weil sorted s<strong>and</strong> than in medium, less weil<br />

sorted s<strong>and</strong> <strong>and</strong>lor silty, fine s<strong>and</strong>.<br />

Thirty percent <strong>of</strong> the species were found in 8 to 10<br />

localities, with only 8 species found in all 10 beaches.<br />

32 % have a rather general distribution, <strong>and</strong> are encountered<br />

in 3 to 7 beaches. About 38 % <strong>of</strong> the species occur<br />

only in 1 or 2 localities: 16 species were only found in<br />

Knokke <strong>and</strong>lor Het Zwin, <strong>of</strong> which 5 were unique for<br />

Knokke <strong>and</strong> 2 unique for Het Zwin.<br />

Schizorhynchia generally are the dominant taxon in<br />

number <strong>of</strong> species. In Heist however, only a few individuals<br />

<strong>of</strong> 3 species are present. The second species richest<br />

taxon is Proseriata. In Knokke <strong>and</strong> Het Zwin, the Proseriata<br />

are represented by the same number <strong>of</strong> species as the<br />

Schizorhynchia. These localities contain the most Otoplanidae-species<br />

(weU known for their preference for<br />

dynamic habitats) <strong>and</strong> the most Typhloplanoida-species.<br />

Strikingly, 9 species with a wide distribution at the Belgian<br />

coast (found in minimum 8localities) do not occur in<br />

Het Zwin.<br />

Table 4 - Diversity<strong>and</strong> evennessindices for <strong>Plathelminthes</strong>in six localities.<br />

Diversity<br />

The <strong>diversity</strong> indices H', SI, No, Nb N2 <strong>and</strong> Ninf. (HlLL<br />

1973, SOETAERT& HElP 1990 <strong>and</strong> references therein) aU<br />

show a significant correlation (Spearman-rank correlation<br />

test, p


40 P. lOUK & E. SCHOCKAERT<br />

100<br />

90<br />

80<br />

Propor1ional<br />

cunUative<br />

abl.M1dance<br />

10<br />

60<br />

so<br />

40<br />

JO<br />

20<br />

o<br />

o<br />

10<br />

.<br />

1 ,.<br />

<strong>Species</strong> order<br />

Fig. I - K-dominancecurves (6 localities).<br />

For the evenness-indices N10,N' 10,N2I, N' 21>no significant<br />

differences were found between localities (Kruskall-Wallis<br />

test, p>0.05), nor was there a significant<br />

correlation with the sediment characteristics (Spearmanrank<br />

correlation test, p>0.05).<br />

Discussion<br />

With over 130 species in total <strong>and</strong> 50 to 60 species per<br />

locality, the Belgian beaches can be considered rich in<br />

turbellarian species, with a <strong>composition</strong> <strong>of</strong>the fauna very<br />

similar to that <strong>of</strong> similar beaches <strong>of</strong> the German Isl<strong>and</strong><br />

Sylt, where studies on the <strong>Plathelminthes</strong> has been going<br />

on since the 50-ties (see REISE1988 <strong>and</strong> references therein).<br />

As the studies in Sylt show, <strong>and</strong> as does this one on<br />

the Belgian coast, some species have a wide distribution,<br />

while others are restricted to limited habitats. As a general<br />

mIe, it can be said that the more the habitats diverge,<br />

the smaller the similarities <strong>of</strong> the turbellarian fauna are.<br />

But considering that on the Belgian coast, 38 % <strong>of</strong> the<br />

species occur only in one or two beaches, it is clear that<br />

minor differences in the habitat (even those difficult to<br />

detect) may have an impact on the turbellarian fauna.<br />

This also explains why several previously unknown species<br />

can be found at our coast.<br />

The <strong>diversity</strong> <strong>of</strong> the turbellarian fauna at the Belgian<br />

coast is also very similar to that noted for similar habitats<br />

on the German Isl<strong>and</strong> Sylt (ARMONIES& HELLWIG-AR-<br />

MONIES 1987, REISE 1983 a, b, WEHRENBERG 1988, WELL-<br />

NER& REISE1989), with the H' ranging from 1.20 to 3.35,<br />

however with higher values for sheltered areas. DrIT-<br />

MANN(1998) found a H' value <strong>of</strong> 2.55 or less in s<strong>and</strong><br />

flats adjacent to mangroves in northem Queensl<strong>and</strong>, Australia,<br />

<strong>and</strong> own unpublished data on the <strong>diversity</strong> <strong>of</strong><br />

turbellarians in Ecuadorian open beaches <strong>and</strong> mangrove<br />

s<strong>and</strong> flats <strong>of</strong> Zanzibar, Tanzania, again all result in H'<br />

values <strong>of</strong> 2.0 to 3.0.<br />

The <strong>diversity</strong> <strong>of</strong>the <strong>free</strong>-<strong>living</strong> <strong>Plathelminthes</strong> is <strong>of</strong>the<br />

same magnitude as that <strong>of</strong> the nematodes <strong>and</strong> harpacti-<br />

100<br />

coid copepods, generally considered the two most dominant<br />

taxa in the mei<strong>of</strong>auna, in s<strong>and</strong>y habitats (BODIN<br />

1988, HElPet al. 1985, HICKS& COULL1983, WARWICK<br />

& GEE 1984). It is clear that <strong>free</strong>-<strong>living</strong> <strong>Plathelminthes</strong><br />

should be considered when assessing the bio<strong>diversity</strong> <strong>of</strong><br />

any marine habitat.<br />

Acknowledgements<br />

The first author thanks the "Instituut tot Aanmoediging van het Wetenschappelijk<br />

Onderzoek in Nijverheid en L<strong>and</strong>bouw" (IWONL) for<br />

the assignment <strong>of</strong> a scholarship from 1985 till 1987.<br />

References<br />

ARMONIES,W. & HELLWIG-ARMONIES, M., 1987. Synoptic pattems<br />

<strong>of</strong> mei<strong>of</strong>aunal <strong>and</strong> macr<strong>of</strong>aunal abundances <strong>and</strong> specific<br />

<strong>composition</strong> in littoral sediments. Helgoländer Meeresunters.,<br />

41: 83-lIl.<br />

BARNETI,p.R.a., 1968. Distribution <strong>and</strong> ecology <strong>of</strong> harpacticoid<br />

copepods <strong>of</strong> an intertidal mudflat. Int. Revue ges. Hydrobiol.,<br />

53: 177-209.<br />

BODIN,P., 1988. Results <strong>of</strong> ecological monitoring <strong>of</strong> three<br />

beaches polluted by the "Amoco Cadiz" oil spill: development<br />

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Philippe lOUK<br />

Royal Zoological Society <strong>of</strong> Antwerp<br />

Koningin Astridplein 26<br />

B-2018 Antwerp<br />

Ernest SCHOCKAERT<br />

Dept. SBO<br />

Limburg University Center<br />

University Campus<br />

B-3590 Diepenbeek<br />

41

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