diel vertical migration - soest · karen e. selph, ocn 621 spring 2009 diel vertical migration:...
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Diel Vertical Migration
OCN 621
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Karen E. Selph, OCN 621 Spring 2009
Outline
Definition
Who does it? How fast?
Migration cues
Why?
Variations: seasonal, ontogenic, reverse
Biogeochemical implications
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Karen E. Selph, OCN 621 Spring 2009
Diel Vertical Migration: Definitions
Usually involves migration into food-filledshallow water at night, and descent into relativelyfood-depleted depths during the day. NormalDVM is when migration occurs at dawn and dusk.
2 general patterns: Nocturnal migration (as above): most common Reverse migration: surface rise during the day, night-
time descent to a maximum depth.
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Karen E. Selph, OCN 621 Spring 2009
How widespread?
Freshwater and marine systems
Occurs in all major groups ofzooplankton
Extent/occurrence varies with life stageand/or sex, season, geographic location,and general weather conditions
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Karen E. Selph, OCN 621 Spring 2009
Examples of Diel Vertical Migrators
These phyla have representatives that DVM, but not allspecies within these phyla migrate:
Crustacea: dominant group: copepods, especially the calanoids,and euphausiids (krill)
Siphonophores Chaetognaths Squid Fish Protists: Reverse DVM (down at dusk, up at dawn)
Ciliates (e.g., autotrophic Mesodinium rubrum) Dinoflagellates
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Karen E. Selph, OCN 621 Spring 2009
Amplitude and extent of migration
Amplitude (depth range): 10s of cm to 100sof meters, and at all depths down to theabyssopelagic zone
Occurs ocean-wide, but especially in highlyproductive tropical areas.
Occurs over diel, seasonal and ontogenictime scales
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Karen E. Selph, OCN 621 Spring 2009
How fast do they do it?
Speeds ranging from 3.24 m/h (0.9 mm/s) inthe small copepod Paracalanus parvus to215 m/h (60.2 mm/s) in the krillMeganyctiphanes norvegica (i.e., slower insmaller swimmers than larger ones).
Downward speeds are often faster thanupward speeds, probably due to gravity.
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Karen E. Selph, OCN 621 Spring 2009
Deep Scattering Layers Moving deep scattering layers are due to diel vertical
migrators
Look like false sea bottoms on echograms
Formed by larger crustaceans, small fish with swimbladders, and occasionally copepods and heteropods
from Lalli & Parsons 1997
Day-time deepscattering layersin Saanich Inlet,BC, Canada
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Karen E. Selph, OCN 621 Spring 2009
Example: ADCP, NE Atlantic
Wade & Heywood 2001
Main scattering layer: ascent at 3-4 cm/s, descent at 10 cm/s
increasing scattering
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Karen E. Selph, OCN 621 Spring 2009
ADCP &MOCNESS
Courtesy of M. Zhou
Antarctica
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Karen E. Selph, OCN 621 Spring 2009
MOCNESS
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Karen E. Selph, OCN 621 Spring 2009
Euphausiasuperba
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Karen E. Selph, OCN 621 Spring 2009
Vertical Migrators at BATS(Sargasso Sea, Atlantic Ocean)
Steinberg et al. 2000
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Karen E. Selph, OCN 621 Spring 2009
Proximate Cue
Light the major cue: in particular theintensity of downwelling irradiance islinked to timing of migration and amplitude.
Is the absolute magnitude of light or the rateof change of light that zooplankton use ascues? Rate of change hypothesis morewidely supported by evidence and scientificcommunity.
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Karen E. Selph, OCN 621 Spring 2009
Total Solar Eclipse, 30 June 1973
Solid Line: Vertical position of the 0.2 µW cm-2 nm-1 isolume
DSL: Vertical position of the sonic scattering layer during the eclipse.
The gap in the record was caused by ship repositioning for betterobservation.
From Kampa 1975 Deep-Sea Res 22:417-423.
isolu
me
DSL
SR
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Karen E. Selph, OCN 621 Spring 2009
Light cues: Longer residence time at thesurface during winter months
Hays 2003
North Atlantic copepods:size also impacted length ofstay at the surface
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Karen E. Selph, OCN 621 Spring 2009
Why do it? Most favor the hypothesis that DVM allows zooplankton to
avoid visual predation in the surface layers by takingtemporary refuge in the darker depths during the day
Cost vs. Benefit: reduced feeding period vs. reducingprobability of predation (“better hungry than dead”)
If this hypothesis true, then expect ascent at dusk and descent at dawn, and DVM will be more pronounced in more conspicuous individuals, and the amplitude of migrations will vary with the abundance and activity
of planktivorous fish
In fact, Reverse DVM also could be explained by this, as thepredators of those doing the R-DVM are themselves doingDVM (invertebrate predators).
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Karen E. Selph, OCN 621 Spring 2009
Visibility & Relative Predation Risk
Vulnerability to visualpredation varies withlight intensity & preysize
It should also dependupon opticaltransparency of thewater (i.e., particleload).
DeRobertis 2002
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Karen E. Selph, OCN 621 Spring 2009
Calanus pacificus:adult females
• Trawls for larval planktivorous fish• Stomach contents of larval fish examined for presence/absence of C. pacificus• Also looked at night/day distributions of C. pacificus (± 75 m)• Found that when significant numbers of fish, DVM behavior strong and vice versa
Bollens & Frost 1989
V = index ofvertical migrationF = fish abundance
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Karen E. Selph, OCN 621 Spring 2009
Egg-bearing females more attractive to predators...
Photo by K.E. Fischer
When presented with egg & non-egg bearingfemales, Pacific herring ingested 4.67 timesas many ovigerous females.
Bollens & Frost 1991
Euchaeta elongata
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Karen E. Selph, OCN 621 Spring 2009
Seasonal/Ontogenic Vertical MigrationLalli & Parsons 1997
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Karen E. Selph, OCN 621 Spring 2009
Calanus pacificus
lipid storessufficient for winter
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Karen E. Selph, OCN 621 Spring 2009
Some fish and sharks migrate,too...
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Karen E. Selph, OCN 621 Spring 2009
Example of vertical migration to maintain position
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Karen E. Selph, OCN 621 Spring 2009
Reverse Diel Vertical Migration
This is the oppositeof DVM: organismcome into surfacewaters during theday to feed, andsink to unlit depthsat night
Pseudocalanus sp. in Dabob Bay, WA
Ohman et al. 1983
Top: Vertical distributionof copepod during day (white)and night (black).
Bottom: Distributions of potential invertebrate predatorsof Pseudocalanus.
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Karen E. Selph, OCN 621 Spring 2009
Seasonalvariation
Ohman 1990
P. newmani femalesPseudocalanus newmani
• 2 stations (shallow & deep)
• displaying diel vertical migration, reverse DVM and no migration
• migration or absence thereof correlated with predator presence
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Karen E. Selph, OCN 621 Spring 2009
Salp migrations inthe Subarctic Pacific
Cyclosalpa bakeri
Top: Day and night distributions ofCyclosalpa bakeri at Stn. P in Aug. 1988
Bottom: Gut pigment contents as afunction of time of day.
Purcell & Madin 1991
Primary predators not visualMigration enables reproduction
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Karen E. Selph, OCN 621 Spring 2009 Jarvis 2003
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Karen E. Selph, OCN 621 Spring 2009
Contribution of Migrants to C & N flux
Al-Mutairi & Landry 2001note: these are minimum flux estimates, as they don’t include deathof migrants at depth, under-sampled micro-nekton, dissolved organic excretion or inter-zonal fecal transport.
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Karen E. Selph, OCN 621 Spring 2009
Steinberg et al. 2000
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Karen E. Selph, OCN 621 Spring 2009
Active area of research...
Deep-Sea Research I, 2009, 1777-1791
Concluded that DVM behavior resulted in 15% of theannual total POC flux at 150 m, mainly through
respiration at depth
One reference of many:
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Karen E. Selph, OCN 621 Spring 2009
Kobari et al. 2003 -- Neocalanus, W. Subarctic Pacific
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Karen E. Selph, OCN 621 Spring 2009
Biogeochemical Implications Active transport of material from euphotic zone to
deep (i.e. food in zp gut is metabolized at depth).
Numerous studies to date have drawn five mainconclusions(1) inter-zonal migrants can significantly enhance oceanic
export fluxes via their DVM behavior,
(2) the relative importance of the active flux is highlydependent on the biomass of the migrating community,
(3) the relative importance of the active versus the passiveflux increases with increasing depth,
(4) inter-zonal migrants may provide a steady source ofnutrients to deep-sea microbial communities, and
(5) we should be including the active flux caused by DVMwhen modeling the cycling of biogeochemically importantelements.