How do Long-range Insect Migrants Control their Flight Direction?
Jason W. ChapmanRothamsted Research, UK
Introduction
Class Species Location Numbers Biomass (tons)Insects Darner Dragonfly Argentina 4 – 6 Billion 4000
Monarch Mexico 200 Million 80Desert Locust North Africa 109 – 1011 200,000
Mammals Wildebeest East Africa 1.3 million 280,000Free-tailed Bat New Mexico 20 million 300
Holland, Wikelski & Wilcove (2006) How and why do Insects Migrate? Science
1. Enormous Numbers and Biomass
3. Simple Systems and Model Organisms
2. Crop Pests and Disease Vectors• Orthoptera (e.g. locusts)• Hemiptera (aphids & planthoppers)• Lepidoptera (armyworms, cutworms & bollworms)• Diptera (mosquitoes, midges & blackflies)• Coleoptera (bark beetles)
Biometeorology of Migration
• Weather hugely influential on insect migration patterns• Wind speed & direction
• Airspeeds of large insects about 3 – 6 ms-1
• Airspeeds of ‘micro-insects’ often <1 ms-1
• Wind speeds usually exceed this a few m above the ground
• Recognition of FBL – Taylor (1974)
• Downwind movement most efficient at high-altitudes
• 10 – 20 ms-1 achieved at just a few 100 m
• Most migrants fly above their FBL
• A few well-studied large insects migrate predominantly within it
Taylor (1974) Insect Migration, Flight Periodicity and the Boundary Layer. J. Anim. Ecol.
Seasonal Migration Patterns
• Most insect migration is one-way
• Return movements, if any, carried out by subsequent generations
• Much movement nomadic by nature, but there are general directional trends• In tropical & sub-tropical regions, movements related to rainfall• In temperate regions, there is a tendency for north-south seasonal movements
• Movements over the necessary distances requires favourable winds• Poleward winds in the spring are warm and thus favourable for migration• Equatorward winds are cool and so will tend to suppress migration
• Led to the idea of the ‘Pied Piper’ effect
• Largely discounted now, return movements have been documented in many species
• Natural selection may be expected to act very strongly on the autumn migration
Do Simple ‘Rules’ Guide Migrations?
Selection of Favourable Winds in Green Darner Dragonflies in the Autumn
Wikelski et al (2006) Simple Rules Guide Dragonfly Migration. Biology Letters
Green Darner Dragonflies Anax junius© Christian Ziegler
• Migrate on days following cooler nights• Fly downwind (tend to be northerlies, so southward movement)• Re-orientated when faced with large water crossing • Simple rules (like songbirds)
Monarch – Time-compensated Solar Compass
Danaus plexippus
Mouritsen & Frost (2002) Virtual Migration in Tethered Flying Monarch Butterflies Reveals their Orientation Mechanisms. PNAS
Return Migration of Vanessa cardui
Painted Lady Vanessa cardui © Ian Woiwod
• Annual immigrant to UK from N Africa
• Varying abundance
• ‘Good’ years: 1996, 2000, 2003, 2006
• Evidence for return migrations?
Rebecca Nesbit
Nesbit, Hill, Sivell, Woiwod & Chapman (in prep)
Flight simulators in Gibraltar May 2006 © Ian WoiwodPainted Lady © Ian Woiwod
Flight Simulators
Mouritsen & Frost (2002) PNAS 99: 10162-10166
Orientation of Autumn MigrantsRebecca Nesbit
N = 94 individualsR = 0.32Mean direction = 193°P < 0.001
Clear Skies
N = 27 individualsR = 0.09No mean directionP = 0.813
Overcast Skies
Nesbit, Hill, Sivell, Woiwod & Chapman (in prep)
Time Compensation?Rebecca Nesbit
N = 94 individualsR = 0.32Mean direction = 193°P < 0.001
Control Group
N = 26 individualsR = 0.46Mean direction = 180°P < 0.001
Clock-shifted
Nesbit, Hill, Sivell, Woiwod & Chapman (in prep)
Vertical-Looking Radar (VLR)
Continuous coverage:150 – 1200 m
Individual insects:15 height bands
Parameters:Speed & direction
OrientationMassShape
0 m
600 m
1200 m
5 mg
15 mg
1 mg
500 mg
Chapman et al (2003) Vertical-looking Radar: A New Tool for Monitoring High-altitude Insect Migration. Bioscience
Orientation Mechanisms in High-flying Moths
• Silver Y Autographa gamma
• Annual migrant to N Europe from Med
• Flies at heights of 200 m – 1000 m
• Layers correspond to regions of wind speed maxima
• Can it have any control over its displacement direction?
Chapman et al (2008a) Wind Selection and Drift Compensation Optimize Migratory Pathways in a High-flying Moth. Current Biology
Chapman et al (2008b) A Seasonal Switch in Compass Orientation in a High-flying Migrant Moth. Current Biology
Spring Migration Events
Displacement Directions
N = 83 Events (20,000+ moths)Mean Direction = 354°R = 0.66P < 0.001
Wind Directions
N = 108 eventsMean Direction = 197°R = 0.29P < 0.001
FlightHeadings
N = 78 Events (20,000+ moths)Mean Direction = 18°R = 0.17P < 0.001
Chapman et al (2008a) Current Biology 18: 514-518Chapman et al (2008b) Current Biology 18: R908-909
Autumn Migration EventsDisplacement
Directions
N = 96 Events (38,000+ Moths)Mean Direction = 169°R = 0.39P < 0.001
Wind Directions
N = 121 EventsNo Mean DirectionR = 0.12P = 0.15
Flight Headings
N = 96 Events (38,000+ Moths)Mean Direction = 195°R = 0.24P < 0.001
Chapman et al (2008a) Current Biology 18: 514-518Chapman et al (2008b) Current Biology 18: R908-909
Displacements HeadingsCompass-mediatedOrientation in Other Species
Large Yellow Underwing Noctua pronuba
Chapman et al (in prep)
Concluding Remarks
• Not passive or ‘accidental’ movements
• Central role for behaviour
• Complicated & sophisticated strategies
• Behaviour often inferred from observations of migration patterns
• Need for experimental approaches
Acknowledgements
Ian Woiwod, Alan Smith, Joe Riley, Duncan Sivell, Rebecca Nesbit (Rothamsted)
Don Reynolds (University of Greenwich)
Henrik Mouritsen (University of Oldenburg)
Jane Hill (University of York)
Curtis Wood, Janet Barlow (University of Reading)
Laura Burgin, Peter Clark (Met Office)
Funding (BBSRC)
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