Ecotypes of NE Atlantic fish species

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Ecotypes of NE Atlantic fish species Ellis, Engelhard & Pinnegar --- Cefas Fish faunas have numerous species, each may respond differently to climate change How to facilitate studying climate impacts on entire fish assemblages? Concept of ecotypes --- groups of species similar in important biological characteristics Life-history, biogeography, feeding guild, etc. We classified all NE Atlantic fishes according to ecotype 1

Ecotype Aspects of ecotypes (a) Biogeographical affinities Boreal Atlantic Lusitanian

Aspects of ecotypes (a) Biogeographical affinities Ecotype Examples Atlantic Eel, blue whiting, tuna, mackerel Boreal Spurdog, herring, cod, saithe, haddock, wolffish, redfish, lemon sole, halibut, witch Lusitanian Many rays, sardine, anchovy, whiting, hake, dory, angler, gurnards, seabass, red mullet, sole

Aspects of ecotypes (b) Reproductive mode Ecotype Planktonic eggs Demersal eggs Egg brooder Nest-forming Viviparous Examples Majority, incl. many clupeids, gadoids, dory, gurnards, (horse-)mackerel, breams, mullets Many rays, herring, many perciforms incl. sandeels, gobies and blennies Seahorses and pipefish Stickleback, wrasses Many sharks and some rays, redfish, eelpout

Ecotype Piscivores Benthivores Aspects of ecotypes (c) Maximum body length (d) Feeding guild and trophic level Feeding on: Preying on smaller fish (and cephalopods) Variety of benthic invertebrates Carcinophages Planktivores Herbivores Detritivores Ectoparasites Specialised on crustaceans (smooth-hounds) Primarily zooplankton Grazing on algae Ingesting sediments (e.g. grey mullets) Feed off other fish (e.g. lampreys)

Aspects of ecotypes (e) Habitat Ecotype Rocky intertidal Estuarine Coastal Contin. shelf Contin. slope Oceanic Examples Few species almost exclusively here, e.g. blenny Grey mullet, flounder, juveniles of many species Many species prevail in shallow coastal zones (but also occur on shelf); nursery areas for many species Majority of North Sea fishes, demersal and small pelagic Northern North Sea and Norwegian Deep: various demersal, bathypelagic, pelagic fishes NW-most North Sea: mainly epi- (-2m) and mesopelagic (2-1m) enter North Sea

Family Species Common name Source Petromyzontidae Lampetra fluviatilis River lamprey Yang (1982) Boreal Fluvial spawner 45 Demersal Coastal 4.5 Ectoparasit Petromyzontidae Petromyzon marinus Sea lamprey Yang (1982) Boreal Fluvial spawner 12 Demersal Shelf 4.4 Ectoparasit Myxinidae Myxine glutinosa Hagfish Yang (1982) Atlantic Demersal eggs 45 Demersal Shelf 3.5 Scavenger Hexanchidae Hexanchus griseus Six-gilled shark Yang (1982) Atlantic Viviparous 5 Bathydemersal Slope 4.3 Piscivore Alopiidae Alopias vulpinus Thresher hark s Yang (1982) Atlantic Viviparous 56 Epipelagic Oceanic 4.5 Piscivore Cetorhinidae Cetorhinus maximus Basking shark Yang (1982) Atlantic Viviparous 98 Epipelagic Shelf 3.2 Planktivore Lamnidae Isurus oxyrinchus Mako shark Yang (1982) Atlantic Viviparous 4 Epipelagic Oceanic 4.5 Piscivore Lamnidae Lamna nasus Porbeagle shark Yang (1982) Atlantic Viviparous 37 Epipelagic Shelf 4.5 Piscivore Scyliorhinidae Galeus melastomus Blackmouthed dogfishyang (1982) Lusitanian Demersal eggs 9 Bathydemersal Slope 4.2 Bentho -piscivor Scyliorhinidae Biogeogr. guild Reprod. guild Scyliorhinus canicula Lesser -spotted dogfish Yang (1982) Lusitanian Demersal eggs 8 Demersal Inner shelf 3.7 Bentho -piscivor Scyliorhinidae Scyliorhinus stellaris Nurse hound Yang (1982) Lusitanian Demersals egg 162 Reef - Inner shelf 4. Piscivore Triakidae Galeorhinus galeus Tope shark Yang (1982) Lusitanian Viviparous 2 Benthopelagic Shelf 4.2 Piscivore Triakidae Mustelus asterias Starry smooth houndyang (1982) Lusitanian Viviparous 14 Demersal Shelf 3.7 Carcinophag Triakidae Mustelus mustelus Smooth hound Yang (1982) Lusitanian Viviparous Demersal Shelf 3.8 Carcinophag Carcharhinidae Carcharhinus longimanus Oceanic whitetip sharkone vagrant 1 Atlantic Viviparous 35 Epipelagic Oceanic 4.2 Piscivore Carcharhinidae Prionace glauca Blue shark Yang (1982) Atlantic Viviparous 383 Epipelagic Oceanic 4.2 Piscivore Sphyrnidae Sphyrna zygaena Common hammerhead Yang (1982) Atlantic Viviparous 4 Epipelagic Oceanic 4.5 Piscivore Dalatiidae Dalatias licha Darkie charlie Yang (1982) Atlantic Viviparous 18 Bathydemersal Slope 4.2 Piscivore Dalatiidae Etmopterus spinax Velvet belly Yang (1982) Atlantic Viviparous 45 Bathydemersal Slope 3.8 Piscivore Dalatiidae Somniosus microcephalusgreenland hark s Yang (1982) Boreal Viviparous 65 Benthopelagic Slope 4.2 Piscivore Squalidae Squalus acanthias Spurdog Yang (1982) Boreal Viviparous Benthopelagic Shelf 4.3 Piscivore Echinorhinidae Echinorhinus brucus Bramble shark Compagno (1984)Atlantic Viviparous 25 Bathydemersal Slope 4.4 Piscivore Squatinidae Squatina squatina Angel shark Yang (1982) Lusitanian Viviparous 25 Demersal Inner shelf 4.1 Piscivore Torpedinidae Torpedo marmorata Marbled electric ray Yang (1982) Lusitanian Viviparous 6 Deme rsal Shelf 4.5 Piscivore Torpedinidae Torpedo nobiliana Common electric rayyang (1982) Lusitanian Viviparous 18 Demersal Shelf 4.5 Piscivore Rajidae Amblyraja radiata Starry ray Yang (1982) Boreal Demersal eggs 9 Demersal Shelf 4. Piscivore Rajidae Dipturus batis Common skate Yang (1982) Boreal Demersal eggs 25 Demersal Shelf 4. Piscivore Rajidae Dipturus linteus Sailray Laverack & Blackler Boreal Demersal eggs 11 Demersal Slope 3.5 Benthivore (1974) Rajidae Dipturus nidarosiensis Black skate Yang (1982) Boreal Demersal eggs 2 Bathydemersal Slope 3.5 Piscivore Rajidae Dipturus oxyrinchus Long-nose skate Yang (1982) Lusitanian Demersal eggs Bathydemersal Slope 3.5 Piscivore Lmax (cm) Habitat Habitat level Trophic Trophic guild

Applying ecosystem concept Climate change and deepening of the North Sea fish assemblage Nick Dulvy and colleagues (Cefas, IMR) From 198-24, fish assemblage has deepened by ~3.6 m decade -1 Also latitudinal response, but more heterogeneous: Northward shift of many Boreal species Southward shift of many warm-tolerant Lusitanian species Aggregate deepening response can be monitored as indicator of warming regional seas 8

Decadal change in depth anomaly in m decade -1 Deeper Shallower pogge scaldfish grey gurnard witch megrim angler hake lemon sole plaice cuckoo ray four-bearded rockling sole lesser weaver bib wolffish solenette cod long rough dab dab haddock whiting ling saithe starry ray lesser spotted dogfish spurdog Norway pout poor cod Significant Not significant -1-5 5 1

Depth anomaly: 4 boreal, 2 lusitanian species 3 3 Depth anomaly (m) -3 3-3 megrim 198 199 2 cod -3 3-3 anglerfish 198 199 2 saithe Shelf Shelf 198 199 2 198 199 2 3-3 sole 3-3 bib Inner shelf 198 199 2 198 199 2

1 5-5 -1 a. boreal 3-3 1 5-5 -1 b. lusitanian 3-3 Depth change by functional group 198 199 2 198 199 2 Depth anomaly (m) 1 5-5 -1 c. Southern boundary 198 199 2 3-3 1 5-5 -1 d. Northern boundary 198 199 2 3-3 1 5-5 -1 e. cold specialists 3-3 1-1 f. warm specialists 3-3 northern boundary species are exception to general pattern 198 199 2 198 199 2

Southward shift northern boundary ecotypes within North Sea: warming and increasing availability shallow habitats in southern North Sea increased inflow warm Atlantic water in northwestern North Sea