Chronic Bottom Trawling Alters the Functional Composition Of

The following appendices accompany the article

Chronic bottom trawling alters the functional composition of

benthic invertebrate communities on a sea-basin scale

H. M. Tillin1,3, J. G. Hiddink1,*, S. Jennings2, M. J. Kaiser1

1School of Ocean Sciences, University of Wales, Bangor, Menai Bridge, Anglesey LL59 5AB, UK

2Centre for Environment, Fisheries and Aquaculture Science (CEFAS), Lowestoft Laboratory, Pakefield Road, Lowestoft, Suffolk NR33 0HT, UK

3Present address: School of Biological Sciences, Biosciences Building, University of Liverpool, Crown Street, Liverpool L69 7ZB, UK

Corresponding author. Email:

Marine Ecology Progress Series 318:31–45 (2006)

1

Appendix 1. Fuzzy coding of ecological characteristics of benthic invertebrates in the North Sea

Ecological characters: / Adult mobility / Habitat / Feeding type / Food type
Species / Perm attach / Temp attach / Burrower / Crawler / Swimmer / Infauna / Epifauna / Epizoic / Deposit / Filter/susp / Browser / Scavenger / Predator / Algae / Inverte-brates / Carrion / Detritus / Plankton / Susp OM / Micro-
organisms
Acanthocardia echinata / 0 / 0 / 3 / 0 / 1 / 4 / 0 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 0
Actinauge richardi / 3 / 3 / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 2 / 0 / 2 / 2 / 0 / 2 / 0 / 0 / 2 / 2 / 0
Adamsia carciniopados / 3 / 1 / 0 / 0 / 0 / 0 / 1 / 3 / 1 / 1 / 0 / 3 / 1 / 0 / 0 / 2 / 2 / 0 / 0 / 0
Aequipecten opercularis / 0 / 2 / 0 / 0 / 2 / 0 / 4 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 3 / 1 / 1
Alcyonidium (Genus) / 4 / 0 / 0 / 0 / 0 / 0 / 3 / 2 / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 4
Alcyonium digitatum / 4 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 4
Amphiura (Genus) / 0 / 0 / 3 / 2 / 0 / 3 / 1 / 0 / 2 / 3 / 0 / 0 / 0 / 0 / 0 / 0 / 1 / 3 / 0 / 0
Anapagurus laevis / 0 / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 3 / 0 / 1 / 2 / 0 / 0 / 0 / 2 / 3 / 0 / 0 / 0
Aphrodita aculeata / 0 / 0 / 3 / 1 / 0 / 4 / 0 / 0 / 1 / 0 / 0 / 3 / 2 / 0 / 0 / 3 / 2 / 0 / 0 / 0
Aporrhais serresianus / 0 / 0 / 3 / 1 / 0 / 3 / 1 / 0 / 3 / 0 / 1 / 0 / 0 / 2 / 0 / 0 / 2 / 0 / 0 / 0
Arctica islandica / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 0 / 1 / 3 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 3 / 1 / 1
Ascidiella (Genus) / 4 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 2 / 2 / 2
Asterias rubens / 0 / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 1 / 3 / 0 / 3 / 2 / 0 / 0 / 0 / 0
Astropecten irregularis / 0 / 0 / 3 / 2 / 0 / 2 / 2 / 0 / 0 / 0 / 0 / 1 / 3 / 0 / 3 / 2 / 0 / 0 / 0 / 0
Bolocera tuediae / 3 / 1 / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 1 / 0 / 0 / 3 / 0 / 4 / 0 / 0 / 0 / 0 / 0
Brissopsis lyrifera / 0 / 0 / 3 / 1 / 0 / 3 / 1 / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 0
Buccinum undatum / 0 / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 3 / 2 / 0 / 2 / 3 / 0 / 0 / 0 / 0
Cancer pagurus / 0 / 0 / 2 / 3 / 0 / 1 / 3 / 0 / 0 / 0 / 0 / 2 / 3 / 0 / 3 / 2 / 0 / 0 / 0 / 0
Colus gracilis / 0 / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 3 / 2 / 0 / 2 / 3 / 0 / 0 / 0 / 0
Corystes cassivelaunus / 0 / 0 / 3 / 1 / 0 / 3 / 1 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 4 / 0 / 0 / 0 / 0 / 0
Crangon allmanni / 0 / 0 / 1 / 3 / 1 / 1 / 3 / 0 / 0 / 0 / 0 / 1 / 3 / 0 / 3 / 1 / 0 / 0 / 0 / 0
Crossaster pappossus / 0 / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 1 / 3 / 0 / 3 / 1 / 0 / 0 / 0 / 0
Dichelopandalus bonnieri / 0 / 0 / 0 / 3 / 3 / 0 / 4 / 0 / 0 / 0 / 0 / 2 / 3 / 0 / 2 / 2 / 0 / 2 / 0 / 0
Ditrupa arietina / 0 / 1 / 3 / 2 / 0 / 3 / 1 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 2 / 2 / 0
Echinocardium (Genus) / 0 / 0 / 3 / 1 / 0 / 3 / 1 / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 0
Echinus esculentus / 0 / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 2 / 0 / 1 / 2 / 3 / 2 / 2 / 2 / 2 / 0 / 0 / 0
Ensis ensis / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 4 / 0
Flustra foliacea / 4 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 4
Liocarcinus holsatus / 0 / 0 / 2 / 2 / 2 / 2 / 2 / 0 / 0 / 0 / 1 / 2 / 2 / 1 / 2 / 3 / 0 / 0 / 0 / 4
Luidia ciliaris / 0 / 0 / 2 / 3 / 0 / 3 / 2 / 0 / 0 / 0 / 0 / 1 / 3 / 0 / 3 / 1 / 0 / 0 / 0 / 0
Luidia sarsi / 0 / 0 / 2 / 3 / 0 / 3 / 2 / 0 / 0 / 0 / 0 / 1 / 3 / 0 / 3 / 1 / 0 / 0 / 0 / 0
Macropodia (Genus) / 0 / 0 / 0 / 3 / 1 / 0 / 4 / 0 / 0 / 0 / 2 / 1 / 3 / 1 / 3 / 0 / 0 / 0 / 0 / 0
Neanthes fucata / 2 / 2 / 0 / 0 / 0 / 2 / 3 / 0 / 1 / 0 / 3 / 1 / 0 / 1 / 3 / 1 / 0 / 1 / 0
Necora puber / 0 / 0 / 0 / 1 / 3 / 0 / 4 / 0 / 0 / 0 / 1 / 2 / 3 / 2 / 3 / 2 / 0 / 0 / 0 / 0
Nephrops norvegicus / 0 / 0 / 2 / 2 / 1 / 3 / 2 / 0 / 0 / 1 / 0 / 1 / 3 / 0 / 3 / 1 / 0 / 0 / 1 / 0
Neptunea antiqua / 0 / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 2 / 3 / 0 / 1 / 3 / 0 / 0 / 0 / 0
Nucula turgida / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 0
Ophiura ophiura / 0 / 0 / 0 / 3 / 2 / 0 / 4 / 0 / 0 / 3 / 0 / 1 / 1 / 0 / 2 / 2 / 0 / 2 / 2 / 2
Opthiothrix fragilis / 0 / 0 / 1 / 3 / 1 / 1 / 3 / 0 / 0 / 3 / 0 / 1 / 1 / 0 / 1 / 1 / 0 / 2 / 2 / 2
Pagurus bernhardus / 0 / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 1 / 1 / 1 / 2 / 1 / 0 / 0 / 2 / 3 / 1 / 1 / 1
Pagurus prideaux / 0 / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 1 / 1 / 1 / 2 / 1 / 0 / 0 / 2 / 3 / 1 / 1 / 1
Pandalus borealis / 0 / 0 / 0 / 3 / 1 / 0 / 4 / 0 / 2 / 0 / 0 / 1 / 3 / 0 / 2 / 0 / 2 / 2 / 0 / 0
Pecten maximus / 0 / 1 / 0 / 0 / 2 / 0 / 4 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 0
Pennatula phosphorea / 4 / 0 / 2 / 0 / 0 / 2 / 2 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 2 / 2 / 0
Polinices (Genus) / 0 / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 4 / 0 / 0 / 0 / 0 / 0
Porania pulvillus / 0 / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 1 / 1 / 0 / 1 / 3 / 0 / 3 / 2 / 0 / 2 / 2 / 2
Porifera (Phylum) / 4 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 4
Processa canaliculata / 0 / 0 / 2 / 2 / 2 / 2 / 2 / 0 / 1 / 0 / 0 / 1 / 3 / 0 / 3 / 1 / 1 / 0 / 0 / 0
Psammechinus miliaris / 0 / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 1 / 0 / 2 / 2 / 2 / 2 / 2 / 2 / 1 / 0 / 0 / 0
Pseudamussium septemradiatum / 0 / 2 / 0 / 0 / 3 / 0 / 4 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 0
Spatangus purpureus / 0 / 0 / 3 / 1 / 0 / 4 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 0
Turritella communis / 0 / 0 / 2 / 2 / 0 / 4 / 0 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 4 / 0

Appendix 2. Fuzzy coding of life history characteristics of benthic invertebrates in the North Sea

Size / Adult longevity yrs / Age at sexual maturity % lifespan / Reproductive technique / Reproductive frequency / Mobility/dissemination
Species / V small / Small / Med / Med-large / Large / V large / <2 / 2-5 / 5-10 / 10+ / <2 / 2-5 / 5-10 / 10+ / Asexual / Sexual spawn / Sex planktonic / Sex mini -adults / Annual once / Annual (2 or more) / Biennial / Semelparous (Ani-mals live to be >2) / No pelagic life stage / Pelagic life stage / Low mobility adult / Highly mobile adult / Migratory
Acanthocardia echinata / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 0 / 1 / 3 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 2 / 3 / 0 / 0
Actinauge richardi / 0 / 2 / 0 / 0 / 2 / 0 / 0 / 0 / 0 / 4 / 2 / 2 / 0 / 0 / 1 / 3 / 0 / 0 / 2 / 2 / 0 / 0 / 0 / 4 / 4 / 0 / 0
Adamsia carciniopados / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 1 / 3 / 0 / 3 / 1 / 0 / 0 / 0 / 4 / 0 / 0 / 2 / 2 / 0 / 0 / 0 / 4 / 2 / 2 / 0
Aequipecten opercularis / 0 / 0 / 1 / 3 / 1 / 1 / 0 / 0 / 4 / 0 / 4 / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 2 / 2 / 0 / 0 / 0 / 4 / 2 / 2 / 0
Alcyonidium (Genus) / 0 / 1 / 3 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 2 / 2 / 0 / 0 / 0 / 2 / 2 / 0 / 4 / 0 / 0 / 0 / 0 / 4 / 4 / 0 / 0
Alcyonium digitatum / 1 / 0 / 1 / 1 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 4 / 0 / 0 / 1 / 3 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 4 / 4 / 0 / 0
Amphiura (Genus) / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 1 / 3 / 2 / 0 / 4 / 0 / 0 / 0 / 4 / 0 / 0 / 3 / 2 / 0 / 0 / 0 / 4 / 3 / 1 / 3
Anapagurus laevis / 4 / 0 / 0 / 0 / 0 / 0 / 1 / 3 / 0 / 0 / 3 / 1 / 0 / 0 / 0 / 0 / 4 / 0 / 1 / 3 / 0 / 0 / 0 / 4 / 3 / 1 / 0
Aphrodita aculeata / 0 / 0 / 0 / 2 / 2 / 0 / 0 / 1 / 3 / 0 / 2 / 2 / 0 / 0 / 0 / 0 / 4 / 0 / 4 / 0 / 0 / 0 / 2 / 2 / 3 / 1 / 0
Aporrhais serresianus / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 1 / 3 / 1 / 3 / 1 / 0 / 0 / 0 / 0 / 4 / 0 / 4 / 0 / 0 / 0 / 0 / 4 / 3 / 1 / 0
Arctica islandica / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 0 / 4 / 0 / 4 / 0 / 0 / 3 / 1 / 0 / 0 / 0 / 4 / 3 / 1 / 0
Ascidiella (Genus) / 0 / 0 / 4 / 0 / 0 / 0 / 3 / 1 / 0 / 0 / 4 / 0 / 0 / 0 / 3 / 3 / 0 / 0 / 3 / 1 / 0 / 0 / 0 / 4 / 4 / 0 / 0
Asterias rubens / 0 / 3 / 1 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 3 / 2 / 0 / 0 / 0 / 4 / 0 / 0 / 3 / 1 / 0 / 0 / 0 / 4 / 2 / 2 / 1
Astropecten irregularis / 1 / 1 / 1 / 0 / 0 / 0 / 0 / 3 / 1 / 0 / 1 / 3 / 0 / 0 / 0 / 4 / 0 / 0 / 3 / 1 / 0 / 0 / 0 / 4 / 2 / 2 / 0
Bolocera tuediae / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 0 / 4 / 2 / 2 / 0 / 0 / 0 / 4 / 0 / 0 / 3 / 1 / 0 / 0 / 0 / 3 / 4 / 0 / 0
Brissopsis lyrifera / 0 / 0 / 2 / 2 / 0 / 0 / 0 / 4 / 0 / 0 / 0 / 4 / 0 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 4 / 3 / 1 / 0
Buccinum undatum / 0 / 0 / 0 / 2 / 2 / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 3 / 1 / 0 / 0 / 0 / 4 / 3 / 0 / 0 / 1 / 4 / 0 / 1 / 3 / 0
Cancer pagurus / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 0 / 4 / 0 / 3 / 2 / 0 / 0 / 0 / 4 / 0 / 3 / 0 / 1 / 0 / 0 / 4 / 2 / 3 / 2
Colus gracilis / 0 / 2 / 0 / 2 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 3 / 1 / 0 / 0 / 0 / 4 / 3 / 0 / 0 / 1 / 4 / 0 / 1 / 3 / 0
Corystes cassivelaunus / 0 / 0 / 2 / 2 / 0 / 0 / 0 / 4 / 0 / 0 / 3 / 1 / 0 / 0 / 0 / 0 / 4 / 0 / 4 / 0 / 0 / 0 / 0 / 4 / 3 / 1 / 0
Crangon allmanni / 4 / 0 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 3 / 1 / 0 / 0 / 0 / 0 / 4 / 0 / 1 / 3 / 0 / 0 / 0 / 4 / 1 / 3 / 1
Crossaster pappossus / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 4 / 2 / 2 / 0 / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 4 / 3 / 1 / 0
Dichelopandalus bonnieri / 2 / 2 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 3 / 1 / 0 / 0 / 0 / 0 / 4 / 0 / 3 / 1 / 0 / 0 / 0 / 4 / 1 / 3 / 1
Ditrupa arietina / 4 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 4 / 4 / 0 / 0
Echinocardium (Genus) / 0 / 0 / 2 / 2 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 4 / 0 / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 4 / 3 / 1 / 0
Echinus esculentus / 0 / 0 / 2 / 2 / 0 / 0 / 0 / 0 / 4 / 0 / 2 / 2 / 0 / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 4 / 3 / 1 / 0
Ensis ensis / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 4 / 0 / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 4 / 2 / 2 / 2
Flustra foliacea / 0 / 2 / 0 / 0 / 2 / 0 / 0 / 0 / 3 / 1 / 2 / 2 / 0 / 0 / 0 / 1 / 3 / 0 / 4 / 0 / 0 / 0 / 0 / 4 / 4 / 0 / 0
Liocarcinus holsatus / 1 / 1 / 3 / 1 / 1 / 1 / 0 / 2 / 2 / 1 / 4 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 2 / 2 / 0 / 0 / 0 / 4 / 1 / 3 / 3
Luidia ciliaris / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 3 / 1 / 2 / 2 / 0 / 0 / 0 / 4 / 0 / 0 / 3 / 1 / 0 / 0 / 0 / 4 / 3 / 1 / 1
Luidia sarsi / 0 / 1 / 1 / 1 / 0 / 0 / 0 / 3 / 2 / 0 / 4 / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 3 / 1 / 0 / 0 / 0 / 4 / 3 / 1 / 1
Macropodia (Genus) / 4 / 0 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 3 / 1 / 0 / 0 / 0 / 0 / 4 / 0 / 1 / 3 / 0 / 0 / 0 / 4 / 3 / 1 / 0
Neanthes fucata / 3 / 1 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 2 / 2 / 0 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 4 / 2 / 2 / 0
Necora puber / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 0 / 1 / 3 / 4 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 1 / 3 / 0 / 0 / 0 / 4 / 0 / 4 / 1
Nephrops norvegicus / 0 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 4 / 0 / 3 / 0 / 2 / 0 / 0 / 4 / 2 / 3 / 0
Neptunea antiqua / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 4 / 0 / 2 / 3 / 2 / 0 / 0 / 0 / 4 / 2 / 0 / 0 / 2 / 4 / 0 / 1 / 3 / 0
Nucula turgida / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 3 / 1 / 0 / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 4 / 4 / 0 / 0
Ophiura ophiura / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 1 / 3 / 0 / 0 / 0 / 4 / 0 / 0 / 3 / 1 / 0 / 0 / 0 / 4 / 3 / 1 / 0
Opthiothrix fragilis / 1 / 3 / 0 / 0 / 0 / 0 / 0 / 1 / 3 / 0 / 3 / 1 / 0 / 0 / 0 / 4 / 0 / 0 / 3 / 1 / 0 / 0 / 0 / 4 / 3 / 1 / 0
Pagurus bernhardus / 0 / 3 / 1 / 0 / 0 / 0 / 0 / 1 / 3 / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 2 / 3 / 0 / 0 / 0 / 4 / 3 / 1 / 0
Pagurus prideaux / 2 / 2 / 0 / 0 / 0 / 0 / 0 / 2 / 2 / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 2 / 2 / 0 / 0 / 0 / 4 / 3 / 1 / 0
Pandalus borealis / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 2 / 3 / 0 / 3 / 1 / 0 / 0 / 0 / 0 / 4 / 0 / 4 / 0 / 0 / 0 / 0 / 4 / 1 / 3 / 1
Pecten maximus / 0 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 4 / 0 / 0 / 0 / 4 / 0 / 0 / 2 / 2 / 0 / 0 / 0 / 4 / 3 / 1 / 0
Pennatula phosphorea / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 1 / 3 / 0 / 0 / 0 / 4 / 0 / 4 / 0 / 0 / 0 / 0 / 4 / 4 / 0 / 0
Polinices (Genus) / 1 / 1 / 1 / 0 / 0 / 0 / 0 / 2 / 3 / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 2 / 2 / 0 / 4 / 0 / 0 / 0 / 4 / 2 / 2 / 0
Porania pulvillus / 0 / 0 / 0 / 0 / 4 / 0 / 0 / 0 / 3 / 1 / 3 / 1 / 0 / 0 / 0 / 4 / 0 / 0 / 3 / 1 / 0 / 0 / 0 / 4 / 3 / 1 / 0
Porifera (Phylum) / 0 / 0 / 0 / 0 / 4 / 0 / 2 / 2 / 2 / 2 / 3 / 1 / 0 / 0 / 2 / 2 / 1 / 0 / 3 / 1 / 0 / 0 / 2 / 3 / 4 / 0 / 0
Processa canaliculata / 4 / 0 / 0 / 0 / 0 / 0 / 1 / 3 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 3 / 1 / 0 / 0 / 0 / 4 / 2 / 2 / 0
Psammechinus miliaris / 1 / 3 / 0 / 0 / 0 / 0 / 0 / 0 / 3 / 2 / 2 / 3 / 1 / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 4 / 3 / 1 / 0
Pseudamussium septemradiatum / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 0 / 4 / 0 / 3 / 1 / 0 / 0 / 0 / 4 / 0 / 0 / 3 / 1 / 0 / 0 / 0 / 4 / 3 / 1 / 0
Spatangus purpureus / 0 / 0 / 0 / 2 / 2 / 0 / 0 / 0 / 1 / 3 / 2 / 2 / 0 / 0 / 0 / 4 / 0 / 0 / 4 / 0 / 0 / 0 / 0 / 4 / 3 / 1 / 0
Turritella communis / 3 / 1 / 0 / 0 / 0 / 0 / 0 / 0 / 3 / 1 / 4 / 0 / 0 / 0 / 0 / 2 / 2 / 0 / 2 / 2 / 0 / 0 / 0 / 4 / 4 / 0 / 0

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Appendix 3. Ecological characteristics of benthic invertebrates in the North Sea

Movement type / Environmental position /

Feeding mode

/ Food
Acanthocardia echinata / Burrower (leaps to avoid predators) (Fish & Fish 1996) / Infauna, top few cm (Fish & Fish 1996) / Filter feeder (Kiorboe & Mohlenberg 1981) / Phytoplankton (Kiorboe & Mohlenberg 1981)
Actinauge richardi / Temporary attachment or free living but sessile (Manuel 1988) / Epifauna (Manuel 1988) / Predator and suspension feeder (Shick 1991) / Invertebrates, suspended organic matter, phytoplankton and zooplankton (inferred from Shick 1991)
Adamsia carciniopados / Permanently attached (Manuel 1988) / Young on rocks and shells, adults almost exclusively on Pagurus prideaux (Peckett 2002) / Scavenger on food scraps from host, sweeps bottom (Warner 1971) / Carrion, organic detritus (inferred)
Aequipecten opercularis / Attached when young, free living as adult, swims swiftly & freely to escape predators (Fish & Fish 1996) / Epifauna (Fish & Fish 1996) / Filter feeder (Fish & Fish 1996) / Suspended organic matter (inferred)
Alcyonidium (Genus) / Permanently attached (Fish & Fish 1996) / Epifauna (Fish & Fish 1996) / Suspension feeder (Fish & Fish 1996) / Micro-organisms (Fish & Fish 1996)
Alcyonium digitatum / Permanently attached (Fish & Fish 1996) / Epifauna (Fish & Fish 1996) / Active suspension feeder (Budd 2001a) / Phytoplankton & zooplankton (Roushdy 1962, Roushdy & Hansen 1961)
Amphiura (Genus) / Crawler/burrower (Buchanan 1964) / Infaunal 4 cm deep (Buchanan 1964, Rosenberg et al.1997). A. filiformis active burrower excavating down to 1173 cm (Solan & Kennedy 2002) / A. chiajei primarily surface deposit feeder (Buchanan 1964). A. filiformis primarily a suspension feeder (Buchanan 1967, Ockelman & Muus 1978) but switches to deposit feeding where current velocities are low (Solan & Kennedy 2002) / Plankton and detritus (Budd 2002a, Hill & Wilson 2001)
Anapagurus laevis / Crawler (inferred) / Epifauna (inferred) / Omnivorous detritivore with some scavenging (most hermit crabs, Hazlett 1981) / Organic detritus, carrion (Hazlett 1981)
Aphrodita aculeata / Crawler/burrower (Mettam 1971) / Infauna (Mettam 1971) / Predator (Mettam 1980) / Active and sedentary polychaetes (Mettam 1980)
Aporrhais serresianus1 / Crawler (Graham 1953) / Shallow infauna (Graham 1953) / Detritus feeder & browser (Graham 1955) / Plant debris (Fish & Fish 1996)
Arctica islandica / Burrower (Sabbatini & Pizzola 2004) / Shallow infauna (Witbaard & Bergman 2003) / Active suspension feeder / surface deposit feeder (Sabatini & Pizzola 2004) / Phytoplankton (Sabatini & Pizzola 2004)
Ascidiella (Genus) / Permanently attached (Fish & Fish 1996) / Epifauna (Fish & Fish 1996) / Suspension feeder (Fish & Fish 1996) / Suspended detritus and phytoplankton (Fish & Fish 1996)
Asterias rubens / Crawler (Budd 2001b) / Epifauna (Budd 2001) / Predator/scavenger (Budd 2001, Howarth & Moore 1996) Mainly predatory (Anger 1977) scavenger on trawl discards (Ramsay et al.1998) / Molluscs, polychaetes, echinoderms (Fish & Fish 1996)
Astropecten irregularis / Burrower & walker (Freeman 2001, Heddle 1967) / Shallow infauna while resting (Christensen 1970); forages over sediment (Freeman & Seed 2001) / Predator (Fish & Fish 1996) / Bivalves, brittle stars, polychaete worms, crustaceans (Fish & Fish 1996)
Bolocera tuediae / Temporarily attached to soft bottoms (Gemill 1921) / Epifauna (Manuel 1988) / Predator / suspension feeder (Schick 1991) / Crustaceans (Vader 1970)
Brissopsis lyrifera / Burrower (Hollertz 1999) / Infauna (Budd 2002b) / Sub-surface deposit feeder (Nichols 1969), occassional surface deposit feeder (Hollertz 1999) / Non-selective (Nichols 1959); feeds on organic detritus, foraminiferans, small organisms (Budd 2002b)
Buccinum undatum / Crawler, relatively sedentary (Hancock 1963, Himmelman & Hamel 1993) / Epifauna (from other gastropods) / Active predator / scavenger; predominantly scavenger (Darkin 1912, Hancock 1960). Scavenger on trawl discards (Ramsay et al.1998) / Carrion, bivalves, polychaetes (Neilsen 1975, Taylor 1978)
Cancer pagurus / Crawler (Neal & Wilson 2004) burrower and slow mover (Watt et al. 1991) / Epifauna (Neal & Wilson 2004) Burrows at night (Houlihan pers. Comm. in Watt et al. 1991) / Predator and scavenger (Neal & Wilson 2004) / Molluscs (Fish & Fish 1996, Neal & Wilson 2004), crustaceans and carrion (Neal & Wilson 2004)
Colus gracilis / Crawler (from other gastropods) / Epifauna (from other gastropods) / Predator/scavenger (from other gastropods) / Carrion, bivalves, polychaetes (Neilsen 1975, Taylor 1978), after Buccinum undatum
Corystes cassivelaunus / Burrower (Warner 1977) / Infauna (Fish & Fish 1996) / Predator (Fish & Fish 1996) / Burrowing invertebrates (Fish & Fish 1996)
Crangon allmanni / Crawler, moves through furrows, tail flip escape action (Burrows et al.2003) / Mobile epifauna, shallow infauna; rests with just antennae exposed (Burrows et al.2003) / Predator (Allen 1960) / Crustacea and annelida preferentially, then mollusca, foraminifera and ophiuroidea (Allen 1960)
Crossaster papposus / Crawler (Sloan 1980) / Epifauna (Sloan 1980) / Active predator (Hancock 1974, Legault & Himmelman 1993). Very infrequent scavenger (Carlson & Pfister 1999) / Wide range, including bivalves, particularly other echinoderms (Hancock 1974)
Dichelopandalus bonnieri / Swimmer / walker (Al-Adhub & Naylor 1977) / Epifauna, demersal (Al-Adhub & Bowers 1977) / Active predator and scavenger (Legardere 1973) / Spring: benthic invertebrates, summer: bathypelagic crustaceans, winter: dead fishes (Legardere 1973)
Ditrupa arietina / Juveniles attached by mucus to substratum, otherwise free living (Gremare et al.1998) / Epifauna/shallow infauna (Jordana et al. 2000) / Suspension feeder (Jordana et al. 2000) / Organic matter, phytoplankton (Jordana et al.2000)
Echinocardium (Genus) / Ecordatum burrower (Lawrence 1987) / Ecordatum infaunal (Buchanan 1967) / Ecordatum Surface deposit feeder (Buchanan 1966) / Ecordatum organic detritus
Echinocardium flavescens / Burrower (inferred after E cordatum) / Infaunal (inferred after E cordatum)) / Deposit Feeder (inferred after E cordatum) / Organic detritus (inferred after E cordatum)
Echinus esculentus / Crawler (Tyler-Walters 2003) / Epifaunal (Tyler-Walters 2003) / Active and passive omnivore (Lawrence 1975) / Worms, barnacles, tunicates, bryozoans, macroalgae, bottom material, detritus (Lawrence 1975)
Ensis ensis / Burrower (Hill 2000) / Lives in deep vertical burrows (Hill 2000) / Active suspension feeder (Hill 2000) / Suspended organic detritus (inferred)
Flustra foliacea / Permanently attached (Tyler-Walters & Ballerstedt 2002) / Epibenthic/epifaunal (Tyler-Walters & Ballerstedt 2002) / Active suspension feeder (Tyler-Walters & Ballerstedt, 2002) / Phytoplankton, detritus, dissolved organic matter (Tyler-Walters & Ballerstedt 2002)
Hairy hydroids, and bryozoans (mixed) / Both permanently attached / Both epifauna / Bryozoans: filter feeders. Hydroids: filter feeders and passive carnivores (Ruppert & Barnes 1996) / Hydroids: suspended organic matter, but mainly zooplankton and small invertebrates (Gili & Hughes 1995, Ruppert & Barnes 1996)
Liocarcinus holsatus / Swimmer / crawler / burrower (Venema et al. 1973) / Epifauna (inferred) / Scavenger/ opportunist (Groenewold & Frond 2000) / Polychaetes, small bivalves, echinoderms, small crustaceans, juvenile fish (Choy 1986). Carrion (Groenewold & Frond 2000)
Luidia ciliaris / Burrower / walker (Heddle 1967) / Mainly epifaunal (Heddle 1976) / Predator (inferred from L sarsi) / Echinoderms, polychaetes, molluscs, crustaceans (inferred from Lsarsi)
Luidia sarsi / Burrower (Fish & Fish 1996), crawler (Brun 1972, Fenchel 1965) / Shallow infaunal by day, epifaunal at night (Picton & Morrow, Fenchel 1965) / Predator (Hayward & Ryland 1996, Brun 1972, Fenchel 1965) / Echinoderms, polychaetes, molluscs, crustaceans, prefers echinoderms –92% of diet (Brun 1972)
Macropodia (Genus) / Crawler, occasionnal swimmer (Hartnoll 1960) / Epifauna (Fish & Fish 1996) / Predator/browser (Hartnoll 1963) / Primarily crustaceans and algae then polychaetes (Hartnoll 1963)
Nereis fucata / Juveniles: benthic, living in mucus tubes; adults: commensal with hermit crabs (Hayward & Ryland 1996) / Commensal with hermit crab and free living (Hayward & Ryland 1996) / Scavenger (Fish & Fish 1996) also filter feeds and predates on hermit crab eggs (inferred from other Nereidae) / Carrion (Fish & Fish 1996) and suspended organic matter (inferred from other Nereidae)
Necora puber / Swimmer and crawler (Watt et al. 1999) / Epifaunal (Phillipart 1998) / Omnivore, herbivore and predator (Norman & Jones 1992) Fast moving predator (Watt et al. 1999) and scavenger (Moore & Howarth 1996) / Brown algae, crustaceans and molluscs, proportion depth; related with deeper crabs more likely to be predators (Norman & Jones 1992) Also carrion (Moore & Howarth 1996)
Nephrops norvegicus / Swimmer and burrower (Sabatini & Hill 2004), walker (Farmer 1974a) / Infauna by day, occupying burrows up to 30 cm deep (Rice and Chapman 1971, large males found in depressions Farmer 1974b). Forage on surface at night, females remain in burrows while carrying eggs (Hughes 1998a, Chapman & Rice 1971) / Predator (Figueiredo & Thomas 1967, Sabatini & Hill 2004) with ability to filter feed (Loo et al.1993) / Crustaceans, molluscs, polychaetes, echinoderms (Sabatini & Hill 2004) and suspended organic particles (Loo et al. 1993)
Neptunea antiqua / Active crawler (Pearce & Thorson 1967) / Epifauna (Pearce & Thorson 1975) / Predator and scavenger (Younge & Thompson 1976, Graham 1953, 1955) Preferentially a scavenger (Pearce &Thorson 1967) / Invertebrate fauna (Fish & Fish 1996) lamellibranchs, macoma, abra, gastropod eggs, polychaetes, ascidians (Blegvad 1914)
Nucula turgida / Burrower (inferred) / Infauna (inferred) / Obligate deposit feeder (Davis & Wilson 1985) / Organic detritus (Davis & Wilson 1985)
Ophiura ophiura / Primarily crawler (inferred) / Epifauna (Hughes 1998b) / Suspension feeder (Fish & Fish 1996), scavenger, predator and suspension feeder (Feder 1981) / Small bivalves, polychaetes, crustaceans, (Tyler 1977, Feder 1981) carrion (Ramsay et al.1998)
Opthiothrix fragilis / Crawler (Jackson 1999,Broom 1975) / Epifauna (Warner 1971) / Suspension feeder, scavenger and predator (Vevers 1956, Nagabushanam & Colman 1959) Mainly suspension feeder (Brun 1969, Warner 1971, Davoult & Gounin 1995) / Detritus/diatoms, (Fish & Fish 1996); silt and detritus (Warner & Woodley 1975); small invertebrates, polychaetes, crustaceans, phytoplankton and diatoms (Brun 1969)
Pagurus bernhardus / Crawler (from other hermit crabs) / Epifauna (from other hermit crabs) / Primarily deposit feeder, then predator, then filter feeder (Gerlach et al.1976, Hazlett 1981) / Organic detritus, carrion (Hazlett 1981)
Pagurus prideauxi / Crawler (from other hermit crabs) / Epifauna (from other hermit crabs) / Primarily deposit feeder, then predator, then filter feeder (Gerlach et al.1976, Hazlett 1981) from Pbernhardus / Organic detritus, carrion (Hazlett 1981)
Pandalus borealis / Mobile, crawler (Shumway et al.1985) / Epifauna (Shumway et al.1985) / Predator, deposit feeder (Allen 1959) / Small invertebrates and detritus (Allen 1959)
Pecten maximus / Swimmer, juveniles attached, adults free living (Fish & Fish 1996) / Epifauna (Fish & Fish 1996) / Suspension feeder (Fish & Fish 1996) / Suspended organic particles (from other pectinids)
Pennatula phosphorea / Permanent attachment (Hill & Wilson 2000) / Epifaunal/infaunal (Hill & Wilson 2000) / Passive suspension feeder (Hill & Wilson 2000) / Plankton & organic particles (Hill & Wilson 2000)
Polinices (Genus) / Crawler (Copeland 1922) / Infaunal/epifaunal, forage over sediment (Ansell 1982) / Predator (Ansell 1982) / Bivalves (Ansell 1982)
Porania pulvillus / Crawler (inferred from body type) / Epifauna (inferred from other Asteroidea) / Ciliary feeder and predator (Ericsson & Hansson 1974) / A digitatum, Brachiopod–Crania anomala, ascidians (Ericsson & Hansson 1974)
Porifera (Phylum) / Permanently attached (Fish & Fish 1996, Hayward & Ryland 1996) / Epibenthic (Fish & Fish 1996) / Filter feeder (Fish & Fish 1996, Hayward & Ryland 1996) / Micro-organisms (Fish & Fish 1996)
Processa canaliculata / Crawler (Smaldon 1979) / Infaunal/epifaunal (Smaldon 1979) / Active predator (Smaldon 1979) / Small polychaetes, sipunculan worms (Smaldon 1979)
Psammechinus miliaris / Crawler (inferred from environmental position) / Epifauna (Jackson 2003, Kelly & Cook 2001) / Predator (Jackson 2003, Hancock 1957) and grazer (Lawrence 1975) / Macroalgae, hydroids, bryozoa, boring sponges, barnacles, molluscs, cockles (Lawrence 1975)
Pseudamussium septemradiatum / Swimmer, juveniles attached, adults free living (from P. maximus) / Epifauna (from other Pectinids) / Filter feeder (from other Pectinids) / Suspended organic particles and phytoplankton
Spatangus purpureus / Burrower (Lawrence 1987) / Shallow infauna (Fish and Fish 1996) / Deposit Feeder ( Rees & Dare 1993) / Organic detritus (inferred)
Turritella communis / Slow-moving, creeping / burrowing (Thorson 1965) / Infauna (Thorson 1965) / Suspension feeder (Graham 1938, Yonge 1946) / Organic detritus (inferred)

Appendix 4. Life history characteristics of benthic invertebrates in the North Sea

Species / Maximum size
(g wet weight) / Adult longevity / Age at sexual maturity / Reproductive technique / Reproductive frequency / Dispersal
Acanthocardia echinata / 77.6 / 10 yr (inferred from size and other bivalves). / Not known, coded as zero. / Sexual, spawners, pelagic larvae (bivalves generally, Fish & Fish 1996). / Annual (C. edule Fish & Fish 1996). / Pelagic larvae (Fish & Fish 1996) but generally low mobility.
Actinauge richardi / 171 / 10 + (inferred from other similarly sized Cnidaria). / 1–5 (inferred from other similarly sized Cnidaria). / Most Cnidarians reproduce asexually and by spawning to produce pelagic larvae (Fautin 2002). / Annual (inferred from other Cnidaria). / Pelagic larvae (Fautun 2002).
Adamsia carcinopados / 42 / 5+ (inferred from other similarly sized Cnidaria). / 1–2+ (inferred from other similarly sized Cnidaria). / Sexual, planktonic, planktotrophic larvae (Chia 1976, Gemill 1921, Shick 1991) / Annual, usually more than 1 cycle, (Shick 1991). / Pelagic larvae (Shick 1991).
Aequipecten opercularis / 49.1 / 6 yr (Heilmayer et al.2004). / 1 yr (Fish & Fish 1996). / Hermaphroditic, sexual spawner. / Annual (Fish & Fish 1996). / Larvae pelagic for
3–4 wk (Fish & Fish 1996).
Alcyonidium (Genus) / 23.4 / 5–10 (inferred from other bryozoa). / 5–10 (inferred from other bryozoa). / Hermaphroditic / sexual pelagic larvae. / Annual (inferred from other bryozoa). / Pelagic larvae (Fish & Fish 1996).
Alcyonium digitatum / 233.6 / 20+ (Hartnoll 1988). / 2–3 yr (Hartnoll 1975). / Colonies gonochoristic and hermaphroditic. Sexual-spawners (Fish & Fish 1996, Hartnoll 1975). / Annual, episodic (Budd 2001a, Hartnoll 1975). / Greater than 10 km (Hartnoll 1975).
Amphiura (Genus) / 1 / A filiformis = 2–4 yr (Ursin 1960, Glemarec & Menesguen 1978)
2–6 yr (Buchanan 1964, O'Connor & McGrath 1980) 10–20 yr (Muus 1981, O'Connor et al.1983). A. Chiajei = 10 yr (Munday & Keegan 1992, Buchanan 1964). / 3–4 yr (Buchanan 1964, Munday & Keegan 1992). / Sexual spawner (Bowmer 1982). / Annual (Buchanan 1964, Fenaux 1970) Annual x 2 for 10–43% of individual A filiformis Bowmer 1982) A. filiformis semelparous but A. chiajei not (Lawrence 1975). / Larvae of A chiajei long pelagic life (Skold et al.1994). Adult A. filiformis can migrate over 2 km (Rosenberg et al.1997).
Anapagurus laevis / 8.2 / Possibly less than 1, maybe 2–5 (inferred from size and other Paguridae). / Most likely below 2 possibly older (inferred from size and other Paguridae). / Gonochoristic, sexual, pelagic larvae (Hazlett 1981). / Annual x 2-broods (Pike & Williamson 1959). / Long range transport as larvae (Hazlett 1981).
Aphrodita aculeata / 68.5 / 6 (Wildish & Peer 1983). / 6 (Wildish & Peer 1983). / Sexual, fertilisation internal (Tyler-Walters & Hughes 2004, Fordham 1925). / Annual protracted (Tyler-Walters & Hughes 2004). / Short pelagic life or non-pelagic (Thorson 1946).
Aporrhais serresianus / 7 / 5–10 (inferred from other similarly sized gastropods). / Most likely below 2 , possibly 2–5 and above (inferred from other similarly sized gastropods). / Sexual (Fish & Fish 1996). / Annual (Fish & Fish 1996). / Long larval life (Lebour 1933).
Arctica islandica / 189.6 / 105 (Fish & Fish 1996) 100+ (Ropes1985, Witbaard & Duineveld 1990). / 11 (Thoriarinsdottir 1999). / Sexual, spawner (Sabatini & Pizzola 2004). / Annual (Sabatini & Pizzola 2004). / Planktotrophic larvae (Sabatini & Pizzola 2004).
Ascidiella (Genus) / 32 / 1–2 yr (Fish & Fish 1996). / 1 yr (Fish & Fish 1996). / Hermaphroditic, spawner. / 1–2 spawning episodes yr–1 (Fish & Fish 1996). / Pelagic larvae (inferred).,
Asterias rubens / 163.1 / 5–10 yr (Budd 2001) 7-8 years (Schafer 1972). / 2nd yr (Fish & Fish 1996) 1 yr (Budd 2001b, Vevers 1949). / Sexual (Budd 2001b) Spawner (Nichols & Barker 1984, Vevers 1949). / Annual episodic (Jangoux & van Impe 1977). / >10 km (Barker & Nichols 1983) Adults do not migrate over large distances (Sparck 1932).
Astropecten irregularis / 9.2 / 10 yr (Christensen 1970),2 4 yr Nojima 1982, Ventura 1999). / 2 yr (Nojima 1982). / Sexual (Fish & Fish 1996). / Annual (inferred from life history). / Pelagic larvae (inferred).
Bolocera tuediae / 452 / 10+ (Vader 1970). / Between 1–5 yr (inferred from longevity). / Sexual pelagic larvae (Gemill 1921) also asexual (Fautin 2002). / Annual Feb–March (Chia 1976, quoted in Shick 1991). / Larvae pelagic for 1 mo (Vader 1970).
Brissopsis lyrifera / 44 / 4 yr (Buchanan 1967). / 4 yr (Buchann 1967). / Sexual spawner (Budd 2002b). / Semelparous (Buchanan 1967). / >10 km (Nichols 1969).
Buccinum undatum / 169 / 50 (Philippart 1998). / 6–9 yr (Valentinsson et al.1999). / Gonochoristic, direct development (Martel et al.1986). / Annual (from N. antiqua). / Adults mobile, but no plantonic larval stage.
Cancer pagurus / 1157.5 / 20 yr (Fish & Fish 1996, Philippart 1998). / 3–5 yr (Fish & Fish 1996) 10 yr (Neal & Wilson 2004). / Sexual, pelagic larvae (Neal & Wilson 2004) / Annual (Neal & Wilson 2004, Fish & Fish 1996) Possibly biennial (Bennett 1995). / >10 km (Neal & Wilson 2004) Adults mobile (Sparck 1932). Females migrate long distances (Bennett 1995).