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
2Centre
of Ocean Sciences, University of Wales, Bangor, Menai Bridge, Anglesey LL59 5AB, UK
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: j.hiddink@bangor.ac.uk
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:
Species
Acanthocardia
Actinauge richardi
echinata
Adamsia carciniopados
Aequipecten
Alcyonidium (Genus)
opercularis
Alcyonium digitatum
Amphiura (Genus)
Anapagurus laevis
Aphrodita aculeata
Aporrhais serresianus
Arctica islandica
Ascidiella (Genus)
Asterias rubens
Astropecten irregularis
Bolocera tuediae
Brissopsis lyrifera
Buccinum undatum
Cancer pagurus
Colus gracilis
Corystes
Crangon allmanni
cassivelaunus
Crossaster pappossus
Dichelopandalus
Ditrupa arietina
bonnieri
Echinocardium
Echinus esculentus
(Genus)
Ensis ensis
Flustra foliacea
Liocarcinus holsatus
Luidia ciliaris
Luidia sarsi
Macropodia (Genus)
Neanthes fucata
Necora puber
Nephrops norvegicus
Neptunea antiqua
Nucula turgida
Ophiura ophiura
Opthiothrix fragilis
Pagurus bernhardus
Pagurus prideaux
Pandalus borealis
Pecten maximus
Pennatula phosphorea
Polinices (Genus)
Porania pulvillus
Porifera (Phylum)
Processa canaliculata
Psammechinus miliaris
Pseudamussium
septemradiatum
Spatangus purpureus
Turritella communis
Adult mobility
Habitat
Feeding type
Food type
Perm
attach
0
3
3
0
4
4
0
0
0
0
0
4
0
0
3
0
0
0
0
0
0
0
0
0
0
0
0
4
0
0
0
0
2
0
0
0
0
0
0
0
0
0
0
4
0
0
4
0
0
0
Temp
attach
0
3
1
2
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
2
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
2
Burrower
Crawler
Swimmer
Infauna
Epifauna
Epizoic
Deposit
Filter/susp
Browser
Scavenger
Predator
Algae
Carrion
Detritus
Plankton
0
0
0
0
0
0
0
0
0
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
0
0
1
0
0
1
0
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
0
Invertebrates
0
2
0
0
0
0
0
0
0
0
0
0
3
3
4
0
2
3
2
4
3
3
2
0
0
2
0
0
2
3
3
3
1
3
3
1
0
2
1
0
0
2
0
0
4
3
0
3
2
0
3
0
0
0
0
0
3
0
3
3
4
0
0
3
0
3
0
2
0
3
1
0
0
3
3
0
4
0
2
2
2
0
0
0
2
0
4
0
1
0
0
0
0
2
0
0
0
2
0
0
0
0
0
0
0
0
2
4
1
1
0
0
4
2
0
1
4
3
4
1
3
4
3
2
1
4
0
0
2
3
3
3
0
1
2
4
0
3
3
4
4
3
0
0
4
4
0
2
4
0
1
0
0
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
3
0
0
0
0
0
2
0
0
1
0
3
1
0
0
2
1
0
0
1
2
0
0
0
0
2
0
3
4
0
0
0
0
0
3
0
4
3
4
0
0
2
0
3
0
1
0
3
1
0
0
3
3
0
4
0
2
3
3
0
2
0
3
0
4
0
1
0
0
0
0
2
0
0
0
2
0
0
0
4
1
4
3
4
1
4
0
1
0
4
4
2
4
1
4
3
4
1
3
4
4
1
1
4
0
4
2
2
2
4
4
2
4
0
4
3
4
4
4
4
2
4
4
4
2
4
4
0
0
3
0
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
3
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
2
3
1
3
1
0
0
0
0
4
0
0
0
0
0
0
0
0
4
2
0
0
0
0
0
0
0
0
0
0
4
0
0
1
1
2
0
0
0
1
0
1
1
0
4
2
1
4
4
4
3
0
0
0
3
4
0
0
1
0
0
0
0
0
0
0
0
4
0
0
4
4
0
0
0
0
1
0
1
0
0
3
3
1
1
0
4
4
0
1
4
0
0
4
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
2
0
1
0
0
0
0
0
1
1
0
0
0
0
0
0
0
2
0
0
2
3
0
0
0
0
2
3
0
0
0
1
1
0
0
3
2
3
0
1
1
2
0
0
2
0
0
2
1
1
1
3
2
1
2
0
1
1
2
2
1
0
0
0
1
0
1
2
0
0
2
1
0
0
0
0
0
2
0
0
0
3
3
3
0
2
3
2
4
3
3
3
0
0
3
0
0
2
3
3
3
1
3
3
3
0
1
1
1
1
3
0
0
4
3
0
3
2
0
0
0
0
0
3
2
1
2
0
0
4
4
0
0
0
0
4
0
0
4
0
0
0
0
0
0
4
2
0
3
0
0
3
0
0
0
3
2
0
0
0
0
0
0
0
0
0
0
2
2
0
0
0
0
0
0
0
0
0
0
0
0
0
2
2
1
1
2
4
2
0
2
0
0
0
4
Susp
OM
0
2
0
1
0
0
0
0
0
0
1
2
0
0
0
0
0
0
0
0
0
0
0
2
0
0
4
0
0
0
0
0
1
0
1
0
0
2
2
1
1
0
0
2
0
2
0
0
0
0
Microorganisms
0
0
0
1
4
4
0
0
0
0
1
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
4
4
0
0
0
0
0
0
0
0
2
2
1
1
0
0
0
0
2
4
0
0
0
0
0
2
0
0
0
0
2
3
0
0
0
2
2
0
0
3
2
3
0
1
1
2
0
0
2
0
0
3
1
1
0
3
2
1
3
0
2
1
2
2
0
0
0
0
2
0
1
2
0
0
0
2
0
0
0
1
3
2
2
0
0
0
0
0
4
0
0
0
0
0
0
0
0
4
2
0
0
0
0
0
0
1
0
0
0
4
0
0
3
3
2
0
0
0
0
0
1
1
0
0
0
0
0
0
0
4
0
0
0
0
4
0
0
2
Appendix 2. Fuzzy coding of life history characteristics of benthic invertebrates in the North Sea
Size
Species
Acanthocardia echinata
Actinauge richardi
Adamsia carciniopados
Aequipecten opercularis
Alcyonidium (Genus)
Alcyonium digitatum
Amphiura (Genus)
Anapagurus laevis
Aphrodita aculeata
Aporrhais serresianus
Arctica islandica
Ascidiella (Genus)
Asterias rubens
Astropecten irregularis
Bolocera tuediae
Brissopsis lyrifera
Buccinum undatum
Cancer pagurus
Colus gracilis
Corystes cassivelaunus
Crangon allmanni
Crossaster pappossus
Dichelopandalus bonnieri
Ditrupa arietina
Echinocardium (Genus)
Echinus esculentus
Ensis ensis
Flustra foliacea
Liocarcinus holsatus
Luidia ciliaris
Luidia sarsi
Macropodia (Genus)
Neanthes fucata
Necora puber
Nephrops norvegicus
Neptunea antiqua
Nucula turgida
Ophiura ophiura
Opthiothrix fragilis
Pagurus bernhardus
Pagurus prideaux
Pandalus borealis
Pecten maximus
Pennatula phosphorea
Polinices (Genus)
Porania pulvillus
Porifera (Phylum)
Processa canaliculata
Psammechinus miliaris
Pseudamussium
septemradiatum
Spatangus purpureus
Turritella communis
Adult longevity yrs Age at sexual maturity %
V small Small Med Med-large Large V large <2 2-5 5-10 10+
0
0
0
0
0
1
0
4
0
0
0
0
0
1
0
0
0
0
0
0
4
0
2
4
0
0
0
0
1
0
0
4
3
0
0
0
0
0
1
0
2
0
0
0
1
0
0
4
1
0
2
4
0
1
0
4
0
0
4
0
0
3
1
0
0
0
0
2
0
0
0
2
0
0
0
4
2
1
0
1
0
1
0
0
0
4
0
3
3
2
4
0
4
1
0
0
0
3
0
0
0
1
3
1
0
0
0
0
0
4
1
1
0
2
0
0
0
2
0
0
0
0
2
2
0
0
3
0
1
0
0
0
0
0
0
4
0
1
0
0
0
0
1
0
0
0
0
0
0
0
3
0
1
0
0
2
0
0
0
0
0
0
2
2
0
2
2
0
0
0
0
2
2
0
0
1
0
1
0
0
0
4
0
0
0
0
0
0
0
4
0
0
0
0
0
0
4
2
0
1
0
0
0
0
2
0
4
0
0
0
0
0
4
0
3
0
1
0
0
lifespan
<2
2-5
5-10 10+
Reproductive technique
Sexual
Asexual
spawn
0
4
1
3
0
4
0
4
0
2
1
3
0
4
0
0
0
0
0
0
0
4
3
3
0
4
0
4
0
4
0
4
0
0
0
0
0
0
0
0
0
0
0
4
0
0
0
4
0
4
0
4
0
4
0
1
0
0
0
4
0
4
0
0
0
4
0
0
0
0
0
0
0
4
0
4
0
4
0
0
0
0
0
0
0
4
0
0
0
0
0
4
2
2
0
0
0
4
Sex
planktonic
0
0
0
0
2
0
0
4
4
4
0
0
0
0
0
0
0
4
0
4
4
0
4
0
0
0
0
3
4
0
0
4
0
4
4
0
0
0
0
4
4
4
0
4
2
0
1
4
0
Reproductive frequency
Sex mini Annual
-adults once
0
2
0
2
0
2
0
2
0
4
0
4
0
3
0
1
0
4
0
4
0
3
0
3
0
3
0
3
0
3
0
0
4
3
0
3
4
3
0
4
0
1
0
4
0
3
0
0
0
4
0
4
0
4
0
4
0
2
0
3
0
3
0
1
0
0
0
1
0
3
4
2
0
4
0
3
0
3
0
2
0
2
0
4
0
2
0
4
2
0
0
3
0
3
0
3
0
4
Mobility/dissemination
Annual (2 or
Semelparous (Ani- No pelagic
Biennial
more)
mals live to be >2) life stage
3
0
0
2
0
0
0
2
0
0
0
2
0
0
0
0
0
0
0
0
0
0
0
2
0
0
0
3
0
0
0
0
0
0
2
0
0
0
0
1
0
0
0
1
0
0
0
1
0
0
0
1
0
0
0
1
0
0
0
0
0
4
0
0
0
1
4
0
1
0
0
0
0
1
4
0
0
0
0
3
0
0
0
0
0
0
0
1
0
0
0
0
4
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
0
0
0
1
0
0
0
1
0
0
0
3
0
0
0
0
0
4
0
3
0
0
0
0
2
0
0
0
0
2
4
0
0
0
0
1
0
0
0
1
0
0
0
3
0
0
0
2
0
0
0
0
0
0
0
2
0
0
0
0
0
0
0
4
0
0
0
1
0
0
0
1
0
0
2
1
0
0
0
0
0
0
0
0
2
0
0
0
0
4
0
0
0
0
0
2
1
0
0
0
0
4
0
4
0
0
0
0
0
0
0
0
0
4
4
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
4
0
0
4
0
0
0
0
0
0
0
0
0
0
1
4
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
3
0
0
0
0
0
0
0
0
0
0
0
4
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
1
0
0
0
1
0
0
0
1
3
1
1
0
1
0
3
0
4
0
0
0
4
4
0
4
0
0
0
0
0
2
0
3
4
4
0
0
0
0
0
1
1
2
2
0
0
2
0
2
3
0
1
0
3
4
4
0
3
0
3
3
0
0
4
1
0
0
0
0
0
0
0
0
0
0
0
4
0
3
2
3
2
0
0
1
4
0
4
4
3
3
2
3
0
0
3
3
2
0
3
3
4
0
0
0
4
2
0
0
1
4
0
0
0
4
0
4
4
4
0
0
4
0
0
4
0
4
1
1
1
0
0
0
3
0
4
0
0
0
0
0
0
4
4
0
1
2
0
2
0
2
3
4
2
0
0
3
2
3
0
4
3
1
2
0
0
0
0
3
3
2
3
4
0
2
0
2
4
2
4
3
2
4
0
0
3
1
3
4
4
3
0
0
4
3
3
4
2
0
2
1
0
2
4
4
1
2
1
0
0
2
3
2
4
0
3
0
1
1
2
1
0
4
2
4
2
0
2
0
1
2
0
4
2
1
3
1
0
0
1
4
1
0
1
1
0
3
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
3
2
3
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
3
0
0
0
0
0
0
0
3
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
4
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
4
0
3
1
0
0
0
4
0
0
3
1
0
0
2
0
2
0
0
0
0
0
0
0
1
3
3
1
2
4
2
0
0
0
0
0
0
0
4
2
0
2
0
0
4
2
0
2
0
0
0
0
Pelagic life
stage
Low mobility Highly
Migratory
adult
mobile adult
4
4
4
4
4
4
4
2
4
4
4
4
4
3
4
0
4
0
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
4
0
4
4
4
4
4
4
4
4
4
4
3
4
4
4
2
2
4
4
3
3
3
3
3
4
2
2
4
3
1
2
1
3
1
3
1
4
3
3
2
4
1
3
3
3
2
0
2
1
4
3
3
3
3
1
3
4
2
3
4
2
3
0
2
2
0
0
1
1
1
1
1
0
2
2
0
1
3
3
3
1
3
1
3
0
1
1
2
0
3
1
1
1
2
4
3
3
0
1
1
1
1
3
1
0
2
1
0
2
1
0
0
0
0
0
3
0
0
0
0
0
1
0
0
0
0
2
0
0
1
0
1
0
0
0
2
0
3
1
1
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
4
3
1
0
0
0
4
4
3
4
1
0
0
0
3
Appendix 3. Ecological characteristics of benthic invertebrates in the North Sea
Movement type
Acanthocardia
echinata
Actinauge richardi
Burrower (leaps to avoid predators)
(Fish & Fish 1996)
Temporary attachment or free living
but sessile (Manuel 1988)
Environmental position
Infauna, top few cm (Fish & Fish
1996)
Epifauna (Manuel 1988)
Adamsia carciniopados Permanently attached (Manuel 1988) Young on rocks and shells, adults
almost exclusively on Pagurus
prideaux (Peckett 2002)
Aequipecten
Attached when young, free living as Epifauna (Fish & Fish 1996)
opercularis
adult, swims swiftly & freely to
escape predators (Fish & Fish 1996)
Alcyonidium (Genus) Permanently attached (Fish & Fish
Epifauna (Fish & Fish 1996)
1996)
Alcyonium digitatum Permanently attached (Fish & Fish
Epifauna (Fish & Fish 1996)
1996)
Feeding mode
Filter feeder (Kiorboe &
Mohlenberg 1981)
Predator and suspension feeder
(Shick 1991)
Scavenger on food scraps from
host, sweeps bottom (Warner 1971)
Filter feeder (Fish & Fish 1996)
Food
Phytoplankton (Kiorboe &
Mohlenberg 1981)
Invertebrates, suspended organic
matter, phytoplankton and
zooplankton (inferred from Shick
1991)
Carrion, organic detritus (inferred)
Suspended organic matter
(inferred)
Suspension feeder (Fish & Fish
1996)
Active suspension feeder (Budd
2001a)
Micro-organisms (Fish & Fish
1996)
Phytoplankton & zooplankton
(Roushdy 1962, Roushdy &
Hansen 1961)
Amphiura (Genus)
Crawler/burrower (Buchanan 1964) Infaunal 4 cm deep (Buchanan 1964, A. chiajei primarily surface deposit Plankton and detritus (Budd 2002a,
Rosenberg et al.1997). A. filiformis
feeder (Buchanan 1964). A.
Hill & Wilson 2001)
active burrower excavating down to filiformis primarily a suspension
1173 cm (Solan & Kennedy 2002)
feeder (Buchanan 1967, Ockelman
& Muus 1978) but switches to
deposit feeding where current
velocities are low (Solan &
Kennedy 2002)
Anapagurus laevis
Crawler (inferred)
Epifauna (inferred)
Omnivorous detritivore with some Organic detritus, carrion (Hazlett
scavenging (most hermit crabs,
1981)
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 Plant debris (Fish & Fish 1996)
1955)
Arctica islandica
Burrower (Sabbatini & Pizzola 2004) Shallow infauna (Witbaard &
Active suspension feeder / surface Phytoplankton (Sabatini & Pizzola
Bergman 2003)
deposit feeder (Sabatini & Pizzola 2004)
2004)
Ascidiella (Genus)
Permanently attached (Fish & Fish
Epifauna (Fish & Fish 1996)
Suspension feeder (Fish & Fish
Suspended detritus and
4
Asterias rubens
1996)
Crawler (Budd 2001b)
Epifauna (Budd 2001)
Astropecten irregularis Burrower & walker (Freeman 2001,
Heddle 1967)
Bolocera tuediae
Brissopsis lyrifera
Shallow infauna while resting
(Christensen 1970); forages over
sediment (Freeman & Seed 2001)
Temporarily attached to soft bottoms Epifauna (Manuel 1988)
(Gemill 1921)
Burrower (Hollertz 1999)
Infauna (Budd 2002b)
Buccinum undatum
Crawler, relatively sedentary
(Hancock 1963, Himmelman &
Hamel 1993)
Epifauna (from other gastropods)
Cancer pagurus
Crawler (Neal & Wilson 2004)
burrower and slow mover (Watt et al.
1991)
Crawler (from other gastropods)
Epifauna (Neal & Wilson 2004)
Burrows at night (Houlihan pers.
Comm. in Watt et al. 1991)
Epifauna (from other gastropods)
Colus gracilis
Corystes cassivelaunus Burrower (Warner 1977)
Infauna (Fish & Fish 1996)
Crangon allmanni
Crawler, moves through furrows, tail Mobile epifauna, shallow infauna;
flip escape action (Burrows et
rests with just antennae exposed
al.2003)
(Burrows et al.2003)
Crossaster papposus
Crawler (Sloan 1980)
Epifauna (Sloan 1980)
Dichelopandalus
bonnieri
Swimmer / walker (Al-Adhub &
Naylor 1977)
Epifauna, demersal (Al-Adhub &
Bowers 1977)
1996)
Predator/scavenger (Budd 2001,
Howarth & Moore 1996) Mainly
predatory (Anger 1977) scavenger
on trawl discards (Ramsay et
al.1998)
Predator (Fish & Fish 1996)
phytoplankton (Fish & Fish 1996)
Molluscs, polychaetes,
echinoderms (Fish & Fish 1996)
Bivalves, brittle stars, polychaete
worms, crustaceans (Fish & Fish
1996)
Crustaceans (Vader 1970)
Predator / suspension feeder
(Schick 1991)
Sub-surface deposit feeder (Nichols Non-selective (Nichols 1959);
1969), occassional surface deposit feeds on organic detritus,
feeder (Hollertz 1999)
foraminiferans, small organisms
(Budd 2002b)
Active predator / scavenger;
Carrion, bivalves, polychaetes
predominantly scavenger (Darkin (Neilsen 1975, Taylor 1978)
1912, Hancock 1960). Scavenger
on trawl discards (Ramsay et
al.1998)
Predator and scavenger (Neal &
Molluscs (Fish & Fish 1996, Neal
Wilson 2004)
& Wilson 2004), crustaceans and
carrion (Neal & Wilson 2004)
Predator/scavenger (from other
Carrion, bivalves, polychaetes
gastropods)
(Neilsen 1975, Taylor 1978), after
Buccinum undatum
Predator (Fish & Fish 1996)
Burrowing invertebrates (Fish &
Fish 1996)
Predator (Allen 1960)
Crustacea and annelida
preferentially, then mollusca,
foraminifera and ophiuroidea
(Allen 1960)
Active predator (Hancock 1974,
Wide range, including bivalves,
Legault & Himmelman 1993).
particularly other echinoderms
Very infrequent scavenger (Carlson (Hancock 1974)
& Pfister 1999)
Active predator and scavenger
Spring: benthic invertebrates,
(Legardere 1973)
summer: bathypelagic crustaceans,
winter: dead fishes (Legardere
1973)
5
Ditrupa arietina
Echinocardium
(Genus)
Echinocardium
flavescens
Echinus esculentus
Ensis ensis
Flustra foliacea
Hairy hydroids, and
bryozoans (mixed)
Liocarcinus holsatus
Luidia ciliaris
Luidia sarsi
Macropodia (Genus)
Nereis fucata
Necora puber
Juveniles attached by mucus to
Epifauna/shallow infauna (Jordana et Suspension feeder (Jordana et al.
substratum, otherwise free living
al. 2000)
2000)
(Gremare et al.1998)
Ecordatum burrower (Lawrence 1987) Ecordatum infaunal (Buchanan 1967) Ecordatum Surface deposit feeder
(Buchanan 1966)
Burrower (inferred after E cordatum) Infaunal (inferred after E cordatum)) Deposit Feeder (inferred after E
cordatum)
Crawler (Tyler-Walters 2003)
Epifaunal (Tyler-Walters 2003)
Active and passive omnivore
(Lawrence 1975)
Organic matter, phytoplankton
(Jordana et al.2000)
Ecordatum organic detritus
Organic detritus (inferred after E
cordatum)
Worms, barnacles, tunicates,
bryozoans, macroalgae, bottom
material, detritus (Lawrence 1975)
Burrower (Hill 2000)
Lives in deep vertical burrows (Hill Active suspension feeder (Hill
Suspended organic detritus
2000)
2000)
(inferred)
Permanently attached (Tyler-Walters Epibenthic/epifaunal (Tyler-Walters & Active suspension feeder (Tyler- Phytoplankton, detritus, dissolved
& Ballerstedt 2002)
Ballerstedt 2002)
Walters & Ballerstedt, 2002)
organic matter (Tyler-Walters &
Ballerstedt 2002)
Both permanently attached
Both epifauna
Bryozoans: filter feeders.
Hydroids: suspended organic
Hydroids: filter feeders and passive matter, but mainly zooplankton and
carnivores (Ruppert & Barnes
small invertebrates (Gili & Hughes
1996)
1995, Ruppert & Barnes 1996)
Swimmer / crawler / burrower
Epifauna (inferred)
Scavenger/ opportunist
Polychaetes, small bivalves,
(Venema et al. 1973)
(Groenewold & Frond 2000)
echinoderms, small crustaceans,
juvenile fish (Choy 1986). Carrion
(Groenewold & Frond 2000)
Burrower / walker (Heddle 1967)
Mainly epifaunal (Heddle 1976)
Predator (inferred from L sarsi)
Echinoderms, polychaetes,
molluscs, crustaceans (inferred
from Lsarsi)
Burrower (Fish & Fish 1996), crawler Shallow infaunal by day, epifaunal at Predator (Hayward & Ryland 1996, Echinoderms, polychaetes,
(Brun 1972, Fenchel 1965)
night (Picton & Morrow, Fenchel
Brun 1972, Fenchel 1965)
molluscs, crustaceans, prefers
1965)
echinoderms –92% of diet (Brun
1972)
Crawler, occasionnal swimmer
Epifauna (Fish & Fish 1996)
Predator/browser (Hartnoll 1963) Primarily crustaceans and algae
(Hartnoll 1960)
then polychaetes (Hartnoll 1963)
Juveniles: benthic, living in mucus
Commensal with hermit crab and free Scavenger (Fish & Fish 1996) also Carrion (Fish & Fish 1996) and
tubes; adults: commensal with hermit living (Hayward & Ryland 1996)
filter feeds and predates on hermit suspended organic matter (inferred
crabs (Hayward & Ryland 1996)
crab eggs (inferred from other
from other Nereidae)
Nereidae)
Swimmer and crawler (Watt et al.
Epifaunal (Phillipart 1998)
Omnivore, herbivore and predator Brown algae, crustaceans and
1999)
(Norman & Jones 1992) Fast
molluscs, proportion depth; related
moving predator (Watt et al. 1999) with deeper crabs more likely to be
and scavenger (Moore & Howarth predators (Norman & Jones 1992)
1996)
Also carrion (Moore & Howarth
6
1996)
Infauna by day, occupying burrows up Predator (Figueiredo & Thomas
Crustaceans, molluscs, polychaetes,
to 30 cm deep (Rice and Chapman
1967, Sabatini & Hill 2004) with echinoderms (Sabatini & Hill
1971, large males found in
ability to filter feed (Loo et
2004) and suspended organic
depressions Farmer 1974b). Forage on al.1993)
particles (Loo et al. 1993)
surface at night, females remain in
burrows while carrying eggs (Hughes
1998a, Chapman & Rice 1971)
Neptunea antiqua
Active crawler (Pearce & Thorson
Epifauna (Pearce & Thorson 1975) Predator and scavenger (Younge & Invertebrate fauna (Fish & Fish
1967)
Thompson 1976, Graham 1953,
1996) lamellibranchs, macoma,
1955) Preferentially a scavenger abra, gastropod eggs, polychaetes,
(Pearce &Thorson 1967)
ascidians (Blegvad 1914)
Nucula turgida
Burrower (inferred)
Infauna (inferred)
Obligate deposit feeder (Davis & Organic detritus (Davis & Wilson
Wilson 1985)
1985)
Ophiura ophiura
Primarily crawler (inferred)
Epifauna (Hughes 1998b)
Suspension feeder (Fish & Fish
Small bivalves, polychaetes,
1996), scavenger, predator and
crustaceans, (Tyler 1977, Feder
suspension feeder (Feder 1981)
1981) carrion (Ramsay et al.1998)
Opthiothrix fragilis
Crawler (Jackson 1999,Broom 1975) Epifauna (Warner 1971)
Suspension feeder, scavenger and Detritus/diatoms, (Fish & Fish
predator (Vevers 1956,
1996); silt and detritus (Warner &
Nagabushanam & Colman 1959) Woodley 1975); small
Mainly suspension feeder (Brun
invertebrates, polychaetes,
1969, Warner 1971, Davoult &
crustaceans, phytoplankton and
Gounin 1995)
diatoms (Brun 1969)
Pagurus bernhardus Crawler (from other hermit crabs)
Epifauna (from other hermit crabs)
Primarily deposit feeder, then
Organic detritus, carrion (Hazlett
predator, then filter feeder (Gerlach 1981)
et al.1976, Hazlett 1981)
Pagurus prideauxi
Crawler (from other hermit crabs)
Epifauna (from other hermit crabs)
Primarily deposit feeder, then
Organic detritus, carrion (Hazlett
predator, then filter feeder (Gerlach 1981)
et al.1976, Hazlett 1981) from
Pbernhardus
Pandalus borealis
Mobile, crawler (Shumway et al.1985) Epifauna (Shumway et al.1985)
Predator, deposit feeder (Allen
Small invertebrates and detritus
1959)
(Allen 1959)
Pecten maximus
Swimmer, juveniles attached, adults Epifauna (Fish & Fish 1996)
Suspension feeder (Fish & Fish
Suspended organic particles (from
free living (Fish & Fish 1996)
1996)
other pectinids)
Pennatula phosphorea Permanent attachment (Hill & Wilson Epifaunal/infaunal (Hill & Wilson
Passive suspension feeder (Hill & Plankton & organic particles (Hill
2000)
2000)
Wilson 2000)
& Wilson 2000)
Polinices (Genus)
Crawler (Copeland 1922)
Infaunal/epifaunal, forage over
Predator (Ansell 1982)
Bivalves (Ansell 1982)
sediment (Ansell 1982)
Porania pulvillus
Crawler (inferred from body type)
Epifauna (inferred from other
Ciliary feeder and predator
A digitatum, Brachiopod–Crania
Asteroidea)
(Ericsson & Hansson 1974)
anomala, ascidians (Ericsson &
Nephrops norvegicus
Swimmer and burrower (Sabatini &
Hill 2004), walker (Farmer 1974a)
7
Hansson 1974)
Micro-organisms (Fish & Fish
1996)
Infaunal/epifaunal (Smaldon 1979)
Small polychaetes, sipunculan
worms (Smaldon 1979)
Psammechinus miliaris Crawler (inferred from environmental Epifauna (Jackson 2003, Kelly &
Predator (Jackson 2003, Hancock Macroalgae, hydroids, bryozoa,
position)
Cook 2001)
1957) and grazer (Lawrence 1975) boring sponges, barnacles,
molluscs, cockles (Lawrence 1975)
Pseudamussium
Swimmer, juveniles attached, adults Epifauna (from other Pectinids)
Filter feeder (from other Pectinids) Suspended organic particles and
septemradiatum
free living (from P. maximus)
phytoplankton
Spatangus purpureus Burrower (Lawrence 1987)
Shallow infauna (Fish and Fish 1996) Deposit Feeder ( Rees & Dare
Organic detritus (inferred)
1993)
Turritella communis Slow-moving, creeping / burrowing Infauna (Thorson 1965)
Suspension feeder (Graham 1938, Organic detritus (inferred)
(Thorson 1965)
Yonge 1946)
Porifera (Phylum)
Permanently attached (Fish & Fish
1996, Hayward & Ryland 1996)
Processa canaliculata Crawler (Smaldon 1979)
Epibenthic (Fish & Fish 1996)
Filter feeder (Fish & Fish 1996,
Hayward & Ryland 1996)
Active predator (Smaldon 1979)
Appendix 4. Life history characteristics of benthic invertebrates in the North Sea
Species
Maximum size
(g wet weight)
Acanthocardia echinata 77.6
Actinauge richardi
171
Adamsia carcinopados
42
Aequipecten opercularis 49.1
Alcyonidium (Genus)
23.4
Alcyonium digitatum
233.6
Adult longevity
Reproductive
technique
10 yr (inferred from size Not known, coded as
Sexual, spawners,
and other bivalves).
zero.
pelagic larvae (bivalves
generally, Fish & Fish
1996).
10 + (inferred from other 1–5 (inferred from other Most Cnidarians
similarly sized Cnidaria). similarly sized Cnidaria). reproduce asexually and
by spawning to produce
pelagic larvae (Fautin
2002).
5+ (inferred from other 1–2+ (inferred from
Sexual, planktonic,
similarly sized Cnidaria). other similarly sized
planktotrophic larvae
Cnidaria).
(Chia 1976, Gemill
1921, Shick 1991)
6 yr (Heilmayer et
1 yr (Fish & Fish 1996). Hermaphroditic, sexual
al.2004).
spawner.
5–10 (inferred from
other bryozoa).
20+ (Hartnoll 1988).
Age at sexual maturity
Reproductive frequency
Dispersal
Annual (C. edule Fish & Pelagic larvae (Fish &
Fish 1996).
Fish 1996) but generally
low mobility.
Annual (inferred from
other Cnidaria).
Pelagic larvae (Fautun
2002).
Annual, usually more
than 1 cycle, (Shick
1991).
Pelagic larvae (Shick
1991).
Annual (Fish & Fish
1996).
Larvae pelagic for
3–4 wk (Fish & Fish
1996).
5–10 (inferred from
Hermaphroditic / sexual Annual (inferred from
Pelagic larvae (Fish &
other bryozoa).
pelagic larvae.
other bryozoa).
Fish 1996).
2–3 yr (Hartnoll 1975). Colonies gonochoristic Annual, episodic (Budd Greater than 10 km
and hermaphroditic.
2001a, Hartnoll 1975). (Hartnoll 1975).
Sexual-spawners (Fish &
8
Amphiura (Genus)
1
Anapagurus laevis
8.2
Aphrodita aculeata
68.5
Aporrhais serresianus
7
Arctica islandica
189.6
Ascidiella (Genus)
32
Asterias rubens
163.1
Astropecten irregularis 9.2
Bolocera tuediae
452
Fish 1996, Hartnoll
1975).
3–4 yr (Buchanan 1964, Sexual spawner
Munday & Keegan
(Bowmer 1982).
1992).
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).
Possibly less than 1,
Most likely below 2
Gonochoristic, sexual,
maybe 2–5 (inferred
possibly older (inferred pelagic larvae (Hazlett
from size and other
from size and other
1981).
Paguridae).
Paguridae).
6 (Wildish & Peer 1983). 6 (Wildish & Peer 1983). Sexual, fertilisation
internal (Tyler-Walters
& Hughes 2004,
Fordham 1925).
5–10 (inferred from
Most likely below 2 ,
Sexual (Fish & Fish
other similarly sized
possibly 2–5 and above 1996).
gastropods).
(inferred from other
similarly sized
gastropods).
105 (Fish & Fish 1996) 11 (Thoriarinsdottir
Sexual, spawner
100+ (Ropes1985,
1999).
(Sabatini & Pizzola
Witbaard & Duineveld
2004).
1990).
1–2 yr (Fish & Fish
1 yr (Fish & Fish 1996). Hermaphroditic,
1996).
spawner.
5–10 yr (Budd 2001) 7-8 2nd yr (Fish & Fish
Sexual (Budd 2001b)
years (Schafer 1972).
1996) 1 yr (Budd 2001b, Spawner (Nichols &
Vevers 1949).
Barker 1984, Vevers
1949).
10 yr (Christensen
2 yr (Nojima 1982).
Sexual (Fish & Fish
1970),2 4 yr Nojima
1996).
1982, Ventura 1999).
10+ (Vader 1970).
Between 1–5 yr (inferred Sexual pelagic larvae
from longevity).
(Gemill 1921) also
asexual (Fautin 2002).
Annual (Buchanan 1964, Larvae of A chiajei long
Fenaux 1970) Annual x 2 pelagic life (Skold et
for 10–43% of individual al.1994). Adult A.
filiformis can migrate
A filiformis Bowmer
1982) A. filiformis
over 2 km (Rosenberg et
semelparous but A.
al.1997).
chiajei not (Lawrence
1975).
Annual x 2-broods (Pike Long range transport as
larvae (Hazlett 1981).
& Williamson 1959).
Annual protracted (Tyler- Short pelagic life or nonWalters & Hughes 2004). pelagic (Thorson 1946).
Annual (Fish & Fish
1996).
Long larval life (Lebour
1933).
Annual (Sabatini &
Pizzola 2004).
Planktotrophic larvae
(Sabatini & Pizzola
2004).
1–2 spawning episodes
yr–1 (Fish & Fish 1996).
Annual episodic
(Jangoux & van Impe
1977).
Pelagic larvae (inferred).,
Annual (inferred from
life history).
>10 km (Barker &
Nichols 1983) Adults do
not migrate over large
distances (Sparck 1932).
Pelagic larvae (inferred).
Annual Feb–March (Chia Larvae pelagic for 1 mo
1976, quoted in Shick
(Vader 1970).
1991).
9
Brissopsis lyrifera
44
4 yr (Buchanan 1967).
Buccinum undatum
169
50 (Philippart 1998).
Cancer pagurus
1157.5
Colus gracilis
106
Corystes cassivelaunus
13.5
Crangon allmanni
6.6
Crossaster papposus
162.1
Dichelopandalus
bonnieri
4
Ditrupa arietina
0.3
Echinocardium (Genus) 40.6
4 yr (Buchann 1967).
Sexual spawner (Budd
2002b).
6–9 yr (Valentinsson et Gonochoristic, direct
al.1999).
development (Martel et
al.1986).
20 yr (Fish & Fish 1996, 3–5 yr (Fish & Fish
Sexual, pelagic larvae
Philippart 1998).
1996) 10 yr (Neal &
(Neal & Wilson 2004)
Wilson 2004).
Semelparous (Buchanan >10 km (Nichols 1969).
1967).
Annual (from N.
antiqua).
Adults mobile, but no
plantonic larval stage.
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).
50 (Phillipart 1998).
6–9 yr (Valentinsson et Gonochorisitc, ovigerous Annual (inferred from N. Low mobility, no pelagic
al.1999 from B.
mini-adults (Fetter &
antiqua).
larval stage.
undatum) Females, 10 yr Graham 1962).
(Graham 1953 from N.
antiqua).
2–5 (inferred from other 1 (inferred from other
Gonochoristic,
Annual (inferred from
Pelagic larvae.
decapod crustaceans of decapod crustaceans of ovigerous, planktonic
longevity).
similar size).
similar size).
larvae.
10–11 yr (Fish & Fish 1.5–2 (Allen 1960).
Gonochoristic (Fish & Annual, usually 2 cycles Free swimming veliger
1996).
Fish 1996).
(Allen 1960).
larvae (Fish & Fish
1996) Migratory
populations move
between inshore and
offshore waters (Allen
1960).
At least 20 yr in Alaska Between 1–5 (inferred Sexual, spawner, larvae Annual (Lambert 1981). Some evidence for
(Carlson & Pfister
from longevity).
prlagic (Carlson &
migrations (Carlson &
1999).
Pfister 1999).
Pfister 1999).
3–4 (Al-Adhub &
2 yr (Al-Adhub &
Sexual, pelagic larvae Annual (Smaldon 1979) Pelagic larvae, possible
Bowers 1977, Pike
Bowers 1977).
(Pike 1954).
migration inshore in
1954).
autumn (Pike 1954).
2 yr (Medernach et
1 yr (Medernach et
Sexual, larvae pelagic. Annual protracted, 2
Pelagic phase 6 wk
al.2000).
al.2000).
spawning peaks/female Medernach et al.2000.
(Medernach et al.2000)
E.cordatum10–20 yr
E.cordatum 3 yr
Sexual, spawner
E.cordatum annual, off E.cordatum pelagic
(Buchanan 1966,
(Buchanan 1966) off(inferred from
shore populations do not larvae.
Hayward et al.1996)
shore populations don't E.cordatum).
breed (Buchanan 1967).
15 yr (Buchanan 1966, breed (Buchanan 1967).
Buchanan 1967).
10
Echinocardium
flavescens
Echinus esculentus
51.4
Ensis ensis
76.5
Flustra foliacea
n/a
Hairy
hydroids/bryozoans
mixes
Liocarcinus holsatus
n/a
Luidai cilaris
229.5
Luidia sarsi
32.3
Macropodia (Genus)
3.9
Nereis fucata
5.6
Necora puber
Nephrops norvegicus
76.5
25.4
10–20 (inferred from E.
cordatum.)
8–10 yr (Nichols 1979
Nichols & Bishop 1985)
16 yr (Gage 1992a & b).
10–20 yr (Hill 2000)
2–3 (inferred from E.
cordatum).
1–3 yr.
Like E. cordatum a
spawner.
Spawner (Fish & Fish
1996).
3 yr (Henderson &
Richardson 1994)
Annual (inferred from E. Pelagic larval stage.
cordatum).
Annual, single cycle
Larvae pelagic for >30 d
(Tyler-Walters 2003).
(Tyler-Walters 2003)
Gonochoristic
planktotrophic (Hill
2000)
Regularly reach 6 yr,
Zooids fertile at 1–2 yr Fertilisation internal,
12 yr old individuals
of age (Eggleston 1972). short-lived pelagic
have been reported
larvae released (Tyler(Stebbing 1971, Ryland
Walters & Ballerstedt
1976).
2002).
Life history
Life history characters Life history characters
characteristics not used not used in the analysis. not used in the analysis.
in the analysis.
2–10 possibly older
I yr (from L. depurator Sexual, pelagic larvae
(inferred from size).
Mori & Zunino 1987). (Choy 1991).
Annual, single cycle
(Fish & Fish 1996)
At least 3.5 yr (Fenchel 1 yr (Fenchel 1965 using Sexual, spawner
1965 from congener L. congener L. sarsi).
(Inferred from other
sarsi).
Echinoderms).
At least 3.5 yr (Fenchel 1 yr (Fenchel 1965).
Sexual, spawner
1965).
(inferred from other
Echinoderms).
3 (inferred from size)
1 possibly 2–5 (inferred Sexual, pelagic larvae
from size).
(Hartnoll 1965)
Annual, (inferred from
other echinoderms).
Annual (Eggleston
1972).
Adults permanently
attached, larvae pelagic
for one day only (but live
in areas of high water
flow).
Life history characters Life history characters
not used in the analysis. not used in analysis.
More than 1 annually
(Choy 1988:1991).
Pelagic larvae, adult
migration >10 km
possible (Choy 1988,
Venema & Creutzberg
1973).
Pelagic larvae.
Annual (inferred from
other Echinoderms).
Long pelagic life as
larvae (Thorson 1961).
3 yr (Hartnoll 1965).
Low mobility, (Hartnoll
1960) pelagic larvae
(inferred).
Pelagic larvae.
Sexual, epitokous (Fish Semelparous (Fish &
& Fish 1996).
Fish 1996).
49.9
3–4 yr (inferred from
Sexual epitokous (Fish
other polychaetes of
& Fish 1996).
similar size).
10 yr (Philippart 1998). 1 yr (Choy 1988).
94
5–10 yr (Sabatini & Hill 2–3 yr (Howard 1989,
Sexual and
Sexual, pelagic larvae
(Choy 1988).
Larval settling time
1–1.5 mo (Hill 2000).
More than 1 annually
(Choy 1988:1991).
Females migrate 10–
100s of m (Norman &
Jones 1993). Little long
distance migration
(Kinnear & Mason 1987,
Norman 1989).
Annual (Sabatini & Hill Larval settling time 11–
11
2004).
Bailey et al.1986).
parthenogenesis,
(Jorgensen 1925).
Females, 10 yr. Males
mature younger at
50–60 mm (Graham
1953).
1 yr (Davis & Wilson
1985).
Sexual, oviparous, mini
adults emerge from egg
mass (Fish & Fish 1996,
Pearce &Thorson 1967).
Spawner, pelagic larvae
(Davis & Wilson 1985).
Neptunea antiqua
337.8
10 yr + (Pearce &
Thorson 1967), 50
(Philippart 1998).
Nucula turgida
3.5
Ophiura ophiura
10.4
Opthiothrix fragilis
5.5
7 yr (Wilson 1988)
5–6 yr (Davis Wilson
1985, Rachor 1976 using
congener N. nitidosa).
5 (Fish & Fish 1996),
Between 1–5 most
5–6 (Gage 1990).
likely, possibly 2–5
(inferred from
longevity).
10 (Fish & Fish 1996), 6–10 mo (Davoult et
10 Gage et al.1990; 9 mo al.1990).
Davoult et al.1990.
Pagurus bernhardus
39.7
Pagurus prideauxi
16.7
Pandalus borealis
9
Pecten maximus
226.8
Pennatula phosphorea
3
2004).
60 mo dispersal gen. low
(Chapman 1980,
Chapman & Bailey 1987
quoted in Tuck et
al.1994). Adults not
migratory Jensen 1965)
Annual, but females may Adults mobile but no
breed once and die
pelagic life stage.
(Pearce & Thorson
1967).
Annual (Davis & Wilson Pelagic larvae.
1985).
Sexual, spawner. (Fish & Annual (Hughes 1998b). Pelagic larvae (Fish &
Fish 1996)
Fish 1996).
Sexual, spawner. (Ball et Annual episodic (Ball et Pelagic larvae
al.1995).
al.1995).
dissemination >10 km
(Jackson 1999) 11–30 d
settling time.
3 yr (Fish & Fish 1996). 1 yr (Fish & Fish 1996). Sexual females carry
Annual, 1 in younger
Several weeks as pelagic
eggs, pelagic larvae
females, more in older larvae (Fish & Fish
(Hazlett 1981)
(Fish & Fish 1996) 1-2 1996) long-range
Lancaster 1990
transport as larvae
(Hazlett 1981)
3 yr from P. bernhardus 1 yr from P. bernhardus. Sexual, pelagic larvae. Most adults produce 2 Pelagic larvae.
broods (Fish & Fish
1996).
3 yr Howard 1983,
18 mo (Howard 1983). Sexual, larvae pelagic Annual 1 (Allen 1959). Planktonic larvae.
7 yr Sokolov 2002,
(Howard 1983).
3–8.5 yr (Rasmussen
1953 quoted in
Shumway et al.1985).
20 yr (Fish & Fish
2 yr (Fish & Fish 1996). Hermaphrodite, sexual, Annual from (Fish &
Larvae planktonic for
1996).
spawner (Fish & Fish
Fish 1996).
3-4 wk (Fish & Fish
1996).
1996).
15 yr (from congener
5–6 yr (Birkeland 1974). Sexual, spawner (Hill & Annual (Hill & Wilson Pelagic larvae, (Hill &
Ptilosaraus guerneyi
Wilson 2000).
2001).
Wilson 2001).
Birkeland 1974).
12
Polinices (Genus)
6.8
Porania pulvillus
116.2
Porifera (Phylum)
92.2
Processa canaliculata
2
Psammechinus miliaris 39.7
Pseudamussium
septemradiatum
Spatangus purpureus
17
Turritella communis
3
85.6
At least 4 (Edwards &
Huebner 1977 from P.
duplicata).
E. catena 22 mo (Ansell Sexual, pelagic larvae E. catena - year round
1982).
(Kingsley-Smith et
>10 broods (juveniles
al.2003 from P.
emerge from eggs as
pulchellus).
mini adults).
5–10 (inferred from
Between 0–5 (inferred Spawner (inferred from Annual 1 (inferred from
other echinoderms)
from other echinoderms) other echinoderms)
other Asteroidea)
Sponges have varying Sexual maturity reached Hermaphrodite, spawner Annual, more than once
lifespans, scored evenly between 1–3 yr (inferred and pelagic larvae
due to variety of
across categories.
from other life history produced, also asexual reproduction strategies.
characterisitcs).
regeneration, budding
(Fish & Fish 1996).
2 yr (Smaldon 1979).
1 yr (Smaldon 1979).
Protandrous
Annual (Smaldon 1979).
hermaphrodite,
fertilisation internal.
(Smaldon 1979).
10+ yr (Fish & Fish
1–2 (Fish & Fish 1996); Sexual, spawner (Kelly Annual (Kelly 2000)
1996, Philippart 1998); 6 1 yr (Gage 1991, Kelly & Cook 2001).
from Kelly & Cook
(Bull 1939) 8 (Jensen
2001) 2 (Jensen 1969).
2001).
1969) 10–12 (Alain
1978).
5–10 yr (from A.
1 yr (from A.
Spawner (from other
Annual (from A.
opercularis).
opercularis).
bivalves).
opercularis).
10–20 yr (Ursin 1960). Between 1–5 (from B. Spawner (from other
Between 1–5 (from B.
Lyrifera and E.
echinoderms).
Lyrifera and E.
cordatum.
cordatum).
‘Fairly old' Thorson
2 (inferred).
Sperm released by males Annual (Thorson 1965).
1965, 5–10 (inferred).
'caught' by females
(Yonge & Thompson
1976).
Low mobility, no pelagic
larval stage.
Low mobility, pelagic
larvae (inferred)
Pelagic larvae.
Pelagic larvae (inferred
from other Decapoda).
Larvae pelagic for 20 d
to 3 mo (Kelly & Cook
2001, Bruce et al.1963).
Pelagic larvae (from A.
opercularis).
Pelagic larvae (from
other echinoderms).
Pelagic larvae (Lebour
1933, Thorson 1946)
Mobilty low (Yonge &
Thompson 1976).
13
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