Diel vertical Migrations Diel vertical migrations • Primary production confined to upper layers • Visual predators forage most efficiently in upper layers • Both food supply and risks of predation decreases by depth • Vertical distribution depends on size; many small forms (~< 1 mm) in upper waters throughout the diel cycle • Larger forms to upper layers under the cover of darkness, recide in deeper waters during the day • Scale: from a few meters to many hundred meters Other reasons for DVM • Migrations into warm upper waters at night to speed up digestion • Some organisms appear to exploit low oxygen deep-water for predator avoidance – ascent into more oxygenated upper waters at night (we have projects in Bunnefjorden and in the Benguela current off Namibia) The classic DVM: ascent at night; descent at day Note however: • Many small forms in upper layers also at day • Some organisms/(organisms on some occasions) may perform inverse migrations (up at day, down at night) • Asynchornous migrations (e.g. individuals making repetitive excursions between upper and lower layers throughout the night) may be important, yet little documented Inverse diel vertical migrations • If the main predator is a dielly migrating, non-visual predator (invertebrates), then inverse diel migrations may occur, i.e. small plankton stays in upper layer at day, and descend at night Genotypic/phenotypic What is genetically fixed, and how flexible is DVM behavior? Field experiment fresh water • Some Alpine lakes are without fish, some lakes stocked with fish for short periods, and some with fish through centuries • Has natural selection resulted in different vertical distribution and DVM in these lakes? Cyclops sp Vertical distribution of Cyclops sp day and night in a lake without fish (left) and a lake stocked with fish for millenia (right) Vertical distribution of Cyclops in lakes stocked with fish for 12 years, 35 years, and several centuries (note that the copepods may dig into the sediment) Conclusions: • No DVM without predators • More migration with increasing time periods (evolutionary time) exposed for predators • There has been a genetic selction in the population • Apparently a high degree of genetically imprinted DVM in the ocean • However: Phenotypic flexibility? • Genetically invariant behavior is profitable with stable conditions of good growth, but high predation risk in upper layers • It is a waste to descend at day (leave food and high temperature) if the threath of predation is not real • Flexibility has been tested in mesocosms Plankton (copepods) in mesocosms with, and without fish Conclusion • Always DVM in the lagoon outside the mesocosms (fish in the lagoon) • DVM in mesocosm with presence of fish • No DVM without fish If the predator is non-visual • Experiments in mesocosms (freshwater) with Chaoborus as predator and the copepod Diaptomus being prey Diaptomus Vertical distribution of a copepod (shaded) day and night with (left) and without a dielly migrating invertebrate predator. Chla profiles suggest the access to food Vertical distribution of a copepod at night in water from a lake without (left) and with ”predator spiked” water Conclusion • The copepod responds to a chemical from the predator (Chaoberus) • Phenotypically indused reversed DVM related to presence of a non-visual predator • The non-visual predator carries out normal DVM; vulnerable to fish and forage on other prey items than the reversely migrating copepod DVM Mesopelagic fish Diel vertical migration of mesopelagic fauna • Many mesopelagic groups (e.g. fish and shrimps) forage in upper layers at night • Some forage during ”antipredation windows at dusk and dawn”; other forage throughout the night, or carry out asynchronous migrations • Apparently little DVM below 800-1000 m Maurolicus muelleri ”mid-night” sinking and dawn ascent ”Antipredation window” • Visual predators like small fish have their own visual predators (larger fish) • Periods at dusk and dawn may represent sufficient strong light to detect own food, while light is sufficiently low to give relative protection against predators • Planktivores need less light than piscivores since they are feeding on much more numerous prey (i.e. do not have to look as far as piscivores) • In this case, the planktivores stop foraging at night (”mid-night sinking”), which is not always the case Sometimes plankton is safer in mid-waters than in deeper waters Krill, saithe and blue whiting in the Norwegian Sea Implications presence of piscivorous fish • Blue whiting has large eyes and can likely forage efficiently on krill in deep waters • Predation pressure increases if it ascends into more shallow waters ”patrolled” by saithe • Presence of saith makes life safer for krill, and krill is safer in mid-waters than in deeper waters Benthosema glaciale Northern lanternfish/nordlig lysprikkfisk Maurolicus muelleri Laksesild Mesopelagic fish Masfjorden Conclusion mesopelagic fish • Species of mesopelagic fish may show varied and divers DVM-patterns, including normal DVM (ascent in darkness; with and without midnight sinking), reversed DVM (some ascent at day), or no migration. • Normal DVM is likely related to feeding in upper waters; reversed DVM to diurnal feeding on midwater prey, and non-migrating may be satiated, or for other reasons lack ”motivation” to migrate