Calanus spp Four species in the northern Atlantic • C. helgolandicus: Mediterranean, Atlantic (central i Bay of Biscay), North Sea (common in the Oslofjord during autumn) • C. finmarchicus: Norwegian Sea • C.hyperboreus: Greenland Sea • C. glacialis: Arctic shelf regions • (Three northernmost species also along the eastern coast of Canada/USA) Relative size C. glacialis, C. hyperboreus (+Metridia sp; left) * C. glacialis and C. finmarchicus Our local hero - Calanus finmarchicus Calanus finmarchicus • Main Calanus-species in the Norwegian Sea • Primarily feed on phytoplankton • Microzooplankton occurs in diet; exact proportion uncertain, partially described as a herbivore, partially as an omnivore • Very important prey organism e.g. fish Large scale distribution C. finmarchicus • Calanus needs access to deep water for overwintering (overwinters at ~500 - 1000 m; somewhat shallower in fjords), thus are confined to deep regions for completing its life cycle • Calanus at the shelf are expatriates, ocean-shelf interactions essential for ”seeding” the shelf • Calanus overwinters in deep fjords, yet probably are expatirates in these environments, dependent on new seeding each year Timing of life history events and number of generations • C. finmarchicus ascends in ~February off western Norway (i.e. well ahead of the spring bloom). Spawning seems to be timed so that their offspring can take benefit of the spring bloom (although the bloom also fuels extra spawning). Ascent/spawning progressively later towards west and north in the Norwegian Sea • C. finmarchicus starts descending for ”overwintering” already in June; one generation being the rule Driving forces life history • Food – spring bloom, length growth season • Predators – timing of life history events, selection of overwintering habitat (horizontally migratory fish/resident mesopelagic fish) • Temperature? (selection overwintering depth) • (advection) Since Calanus are mainly restricted to oceanic regions, major planktivorous fish stocks carry out horizontal feeding migrations Distribution of herring May 2002, blue arrows indicate migrations of herring in April/May and red arrows mirgations in June 2002 Distribution of herring August 2002. Red arrow indicates migrations of herring in July; blue arrows migration August 2002 The herring life history is adapted to Calanus finmarchicus life history (and vice versa?) • Calanus develops progressively later from east to west, and from south to north • Herring is following this progression, exploiting Calanus ascending from overwintering and their progenies • As Calanus descends for overwintering, herring return to their own overwintering areas (fjords in northern Norway). • The herring migrations imply a vast wave of potential mortality – spawning ahead of their arrival might represent a selective advantage • Predation pressure is at a maximum in mid-summer (herring, blue whiting and mackerel peak abundance in the Norwegian Sea). • Driving force for seasonal descent? Copepod life cycles: 6 nauplii stages (NI – NVI) 6 copepodite stages; (CI-CVI), C6 are the adults May diapause in different stages Calanus finmarchcus: * Spawning peak around spring bloom * Diapause in deep water (C5 mostly, C4-C6) * Descent variable, autumn/late summer * Adult about 3.5 mm prosome * 1 – more generations per year Calanus finmarchicus Calanus finmarchicus stn M Differences between Calanus species • Size • Number of yearly generations (although related to habitat) • Timing of life history events • Spawning based on internal (C. hyperboreus), vs primarily external resourses (C. finmarchicus) • Habitat requirements Number of generations • C. finmarchicus normally one generation year-1 in the Norwegian Sea • C. helgolandicus normally 2? generations year-1 (North Sea) • C. glacialis normally needs 2 year to complete the life cycle • C. hyperboreus >2 years to complete life cycle Timing of spawning and habitat requirements • In co-occurring poulations, C. glacialis spawns first, then C. finmarchicus and then C. helgolandicus • Habitat requirements often explained in terms of temperature (e.g. C. glacialis confined to waters colder than 2 ºC) • Overwintering of C. finmarchicus in cold Norwegian Sea water rather than in overlaying warm Atlantic water ”Match-mismatch” theory • Fish spawn at at rather fixed time • Successful recruitment of copepods (nauplii) is related to the spring bloom, which varies with weather conditions • So while fish larvae are present in a rather fixed time window (although temperature affects developmental time), their food occurs during a more variable, and more weather-dependent time window (also related to transport of Calanus onto the shelf). • To find food at time of first feeding (i.e. when the yolk sack is used) is critical for the larvae. • If this coinsides with present of copepod eggs and nauplii, there is a ”match”; if these young copepod stages are scarse during the period for first feeding, there is a ”mis-match”; leading to mass mortality of larvae (according to the theory) Potential food chain impacts of climate change • Changes in temperature, wind and light will impact timing of the spring bloom, and thus timing and magnitude of zooplankton production • C. finmarchcius is very important prey for fish larvae and juvenile fish of most commercially important species (as well as for larger herring and mackerell) • Spring the most critical period for fish larvae/(0group) Possible effects climate change; shifts between C. finmarchicus and C. helgolandicus C. Helgolandicus and C. finmarchicus display opposite climate responses Conclusion: • Substitution of C. finmarchicus with C. helgolandicus will seriously alter timing of the presence of important prey for fish, causing serious ”mis-match” • C. helgolandicus spawns considerably later, and will not be present at time when first-feeding larvae occur. • Increased temperature increases metabolic strain when food is limited