Handbook of the marine fauna of north-west europe

808 Pages·2017·102.41 MB·English

by  Peter J. Hayward, John S. Ryland

HANDBOOK OF THE MARINE FAUNA
OF NORTH-WEST EUROPE

H A N D B O O K O F T H E
Marine Fauna of
North-West Europe
SECOND EDITION
EDITED BY
.
PETER J HAYWARD
.
AND JOHN S RYLAND
1
3
GreatClarendonStreet,Oxford,OX26DP,
UnitedKingdom
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PREFACE
The first edition of the Handbook of the Marine Fauna of North-West Europe was published in
1995,anditisperhapsameasureofitsusefulnessthatithasbeenreprintedinalmosteveryyear
since. It evolved from a two-volume predecessor, The Marine Fauna of the British Isles and North-
West Europe (Clarendon Press, 1990), which had originated as a manual intended to aid the
identification of the most familiar and widely distributed species of fish and benthic macrofauna of
the coastal waters of Britain and Ireland, and the adjacent coasts of Europe. Prior to then, marine
biologists,amateurandprofessional,hadperforcetorelyuponarangeoffieldguides,monographs,
andspecialistpublicationsfortheidentificationofcoastalmarineanimals.TheHandbookwasamore
compactwork,withfewerspeciesandratherlessdetailthanwasincludedinitsprogenitor,butitwas
notsimplyanabridgement.Whilethetotalnumberofspeciestreatedwasreduced,asmallnumberof
additionalspecieswasincluded,andrecenttaxonomicrevisionswereincorporated.
The wider application of, in particular, molecular genetic techniques for the characterization of
individual species has resulted in substantial changes in the understanding of the systematics and
phylogeny of practically all marine animal groups. Phylogenetic analyses have revealed previously
unsuspectedrelationships,demandingreappraisaloftraditionalclassificationsandinvolvingconsid-
erablemodificationoftaxonomicnomenclature.Whilerelativelyfewentirelynewspeciesofmacro-
fauna have been revealed, the marine communities of the north-west European shelf have changed.
The sources of change are various. In a period of rising concern for the health of the natural
environment,therehasbeenanincreaseintheextentofmonitoringandrecordingofmarinespecies
andcommunities;oneconsequenceofthisisthatsomespeciesformerlyconsideredtobeuncommon,
and infrequently encountered, are now seen to be quite widely distributed within the region. The
application of more selective sampling techniques also allows the occurrence of delicate or deep-
burrowing infaunal animals to be more accurately recorded. Anthropogenic disturbance is another
significantsourceofchange;benthicfishinggear,anddredgesemployedinaggregateextraction,may
result in the decline of some components of the benthic community, and the spread of others.
Environmentalchangeisalsoasignificantfactordrivingchangeinthemarinebenthiccommunities
of north-west Europe. The geographical distributions of, for example, south-western, warm-water
species are spreading northwards as sea temperatures rise, while the ranges of northern, cold-water
species may be expected to contract. Exotic, invasive species are an especially conspicuous new
element in the marine fauna of north-west Europe, and the number of newly occurring species is
likelytocontinuetoincrease,partlyasaconsequenceofseawaterwarming,butalsoasadirectresult
of anthropogenic introductions, as part of the ship-borne fouling community, and as associates of
translocatedoysterstock.Thespreadofsuchspeciesisfurtherfacilitatedbycoastalleisurecraftand
thedevelopmentofmarinas,allowingstepping-stonedispersalthroughtheregion.
The primary purpose of this book is to provide swift and simple identifications of the animal
specieslikelytobeencounteredduringshore-basedandcoastalmarinefieldstudies,andduringthe
vi PREFACE
courseofinshoresurveyandmonitoringwork.Theselectionofanimalsincludedvariesbetweenthe
differentgroups.Thecoverageofmolluscsanddecapodsisstillextensive,asthesetendtobeamong
the ecologically more important components of the benthos and attract the eye more readily than
hydroids, small worms, or copepods. Some groups requiring specialist techniques for collection,
examination, and identification have been excluded entirely, for example, Protozoa, Pogonophora,
and Tardigrada. The selection of fish is limited to intertidal and coastal species, as are those of
sponges, amphipods, and mysids. The taxonomy and classification of each animal group have been
updated. A few species described in Marine Fauna but excluded from the first edition of the
Handbookhavebeenrestoredinthissecondedition,asresultofnewinformationontheirdistribution
andoccurrence,andanumberofnew,exoticspeciesareaddedtotheoriginalselection.
Weareconfidentthatthisbookwillserveitspurpose,allowingstudent,amateur,andprofessional
biologists to identify with accuracy the preponderance of animal species encountered between the
tidemarks andintheshallowsubtidal aroundtheBritishIslesandneighbouring coastsofmainland
Europe. Inevitably, rare animals are occasionally encountered and will not be identifiable using this
text.Insuchcases,referenceshouldbemadetothespecialisttextsindicatedforeachanimalgroup;
wherever possible, the confirmatory notes for each family covered include estimates of its total
taxonomic diversity in the region. With the exception of the ctenophores and large pelagic scypho-
zoans,whichfrequentlybeachonsandyshores,alloftheinvertebrateanimalsincludedbelongtothe
benthos.Theplanktonandthemeiofauna,althoughinterestingandabundant,andcertainlydeserv-
ingofattention,aretoonumerousanddiverseforevenarepresentativeselectiontobeincludedhere.
Togetherwithseabirdsandmarinemammals,theyhaveanextensiveliteratureoftheirown.
Swansea P.J.H.
2016 J.S.R.
ACKNOWLEDGEMENTS
MostofthecontributorstothefirsteditionoftheHandbookarelongretired,andafewarenow
nolongerwithus.Preparationofarevisedsecondeditionhasthusbeenademandingtaskthat
has taken longer than anticipated to achieve. The editors are extremely grateful to those original
contributors who felt able to return to systematic zoology, and to update their accounts. In some
cases, this has been a considerable task, as nomenclature and phylogenetic classifications have
changed extensively, and new data on distribution patterns have accumulated. We are also deeply
indebtedtothoseyoungercolleagueswhohavemadetimeinbusycareerstoundertaketherevisionof
those chapters, including some of the largest, for which no original contributor could take
responsibility.

CONTENTS
ListofContributors xi
ListofIllustrators xiv
1. The Marine Environment of North-West Europe 1
PeterJ.Hayward
2. Using This Book 19
PeterJ.Hayward
3. Sponges (Phylum Porifera) 35
ClaireGoodwin,BernardE.Picton,ChristineC.Morrow,andPeterE.J.Dyrynda
4. Hydroids, Sea Anemones, Jellyfish, and Comb Jellies (Phyla Cnidaria
and Ctenophora) 65
PaulF.S.Cornelius,RichardL.Manuel,JohnS.Ryland,
PeterSchuchert,andChrisWood
5. Flatworms and Ribbon Worms (Phyla Xenacoelomorpha, Platyhelminthes
and Nemertea) 133
RayGibsonandE.WynKnight-Jones
6. Annelids (Phylum Annelida) 165
PhyllisKnight-Jones,E.WynKnight-Jones,KateMortimer-Jones,
AnthonyNelson-Smith,RüdigerM.Schmelz,andTarmoTimm
7. Priapulids, Sipunculans, Echiurans, and Entoprocts 271
E.WynKnight-JonesandJohnS.Ryland
8. Crustaceans (Phylum Arthropoda, Subphylum Crustacea) 283
GailV.Ashton,AngelikaBrandt,MichaelJ.Isaac,PhilipMakings,JohnMoyse,
ErnestNaylor,GeoffreySmaldon,andJohnI.Spicer
9. Mites and Sea Spiders (Phylum Arthropoda, Subclass Acari, Class Pycnogonida) 441
RogerN.Bamber,PhilipE.King,andPhilipJ.A.Pugh
10. Molluscs (Phylum Mollusca) 455
A.LouiseAllcock,PeterJ.Hayward,GeoffreyD.Wigham,andNathalieYonow
11. Bryozoa 603
JohnS.Ryland