9.3 Animals migrate in response to changes in the environment Migration – Relatively long-distance two-way movements From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press

Migration and changing resources In the Serengeti, mammals appear to track available biomass of grass and its quality From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press

Figure 8.16 Short-range migration in the three-wattled bellbird

Levey and Stiles 1992

Resource variation and migration in neotropical birds Research question: What factors are important in determining whether a species migrates or not? (Boyle & Conway 2007) Hypothesis: Migration evolved to allow individuals to: (1) take advantage of spatial variation in conditions and (2) avoid seasonal resource depression at different locations Prediction: Species exposed to high degree of fluctuation in environmental conditions and resources will be more likely to migrate than species that live in more stable resource environments From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press Featured Research

From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press

Resource variation and migration in neotropical birds Methods: – Collected previously published data on 300 species in Tyranni – For each species, determined: Migrant or sedentary Insectivore or frugivore Habitat use (tropical forest habitats more stable than nonforest habitats) From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press Featured Research

Resource variation and migration in neotropical birds Results: – Insectivorous species in forest habitats were less likely to be migratory than nonforest species – No relationship between habitat type and migration in frugivorous species Conclusion: – Fluctuations in resource levels and environmental conditions can affect migratory behavior From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press Featured Research

Figure 10-16

Figure Ted Garland

From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press

Figure 8.17 The geographic distribution of Catharus thrushes, a genus that contains both resident and migratory species

Outlaw et al. 2003

Figure 8.18 The long-distance migratory trait overlain on the phylogeny of Catharus thrushes

Figure 8.25 The leapfrog migratory pattern of western North American populations of the fox sparrow

Bell, 1997 fuliginosa

Bell, 1997

Competition and migratory behavior of newts Research question: How do environmental conditions affect migratory behavior? (Grayson & Wilbur 2009) Hypothesis: Competition for resources will result in more individuals migrating Prediction: As density increases, more individuals should migrate From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press Featured Research

Competition and migratory behavior of newts Methods: – Red-spotted newt (Notophthalmus viridescens) – Small enclosures around large pond – Treatments: Density of newts (low, medium, or high) Sex ratio (male or female biased) – Measured migration out of enclosures From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press Featured Research

Competition and migratory behavior of newts Results: – More newts migrated out of high-density enclosures Conclusion: – Population density does affect the migratory behavior of newts From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press Featured Research

Maintenance of polymorphism in migratory behavior Migratory behavior could be a fixed, genetic trait, and frequency-dependent selection maintains the polymorphism Migratory behavior is dependent on individual condition From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press

Competition and migratory behavior of dippers Research question: What maintains resident and migratory behaviors within a population? (Gillis et al. 2008) From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press Featured Research

Competition and migratory behavior of dippers Methods: – American dippers (Cinclus mexicanus) – Uniquely banded individuals – Observed migration behavior over seven years From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press Featured Research

Competition and migratory behavior of dippers Results: – 152 residents and 90 migrants – Residents fledged 2.6 more offspring than migrants Conclusion: – The fitness of residents is higher than that of migrants; migratory behavior appears to be condition-dependent From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press Featured Research

9.4 Animals use multiple compass systems to determine direction Orientation – Determination and maintenance of a proper direction Navigation – Determining a particular location and moving toward it Sun compass – Use of sun for orientation Star compass – Use of stars or constellations to orient Geomagnetic compass – Ability to orient using the earth’s magnetic field From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press

9.4 Animals use multiple compass systems to determine direction From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press

9.4 Animals use multiple compass systems to determine direction From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press

Monarch butterfly migration From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press

Monarch butterfly migration From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press

The magnetic compass in sea turtles Research question: Can sea turtles use a magnetic compass? (Lohmann 1991) From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press Featured Research

The magnetic compass in sea turtles Methods: – Juvenile loggerhead sea turtles (Caretta caretta) – Treatment – altered magnetic fields – Control – normal magnetic fields – Measured movement and orientation From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press Featured Research

The magnetic compass in sea turtles Results: – Only controls moved in correct direction Conclusion: – Loggerhead turtles do use a geomagnetic compass From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press Featured Research

Applying the Concepts 9.2 Citizen scientists track fall migration flyways of monarch butterflies Citizen scientists collect data on migration of butterflies From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press Applying the Concepts

Count of publications presenting new analyses on monarchs (the monarch research category) over time separated by whether they used citizen science data in their analysis. Leslie Ries, and Karen Oberhauser BioScience 2015;65: © The Author(s) Published by Oxford University Press on behalf of the American Institute of Biological Sciences. All rights reserved. For Permissions, please

9.5 Bicoordinate navigation allows individuals to identify their location relative to a goal Bicoordinate navigation – Ability to identify a geographic location using two varying environmental gradients From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press

Bicoordinate navigation and magnetic maps in sea turtles Research question: Do sea turtles use bicoordinate navigation? (Lohmann et al. 2001) From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press Featured Research

Bicoordinate navigation and magnetic maps in sea turtles Methods: – Juvenile loggerhead sea turtles (Caretta caretta) – Exposed to three magnetic fields – Measured orientation and movement From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press Featured Research

Bicoordinate navigation and magnetic maps in sea turtles Results: – All turtles oriented to appropriate direction Conclusion: – Sea turtles can use the earth’s magnetic field to orient in different directions, and this ability is inherited, not learned From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press Featured Research

Bicoordinate navigation in birds Research question: Do birds display bicoordinate navigation? (Chenetsov, Kishkinev, & Mouritse 2008)? From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press Featured Research

Bicoordinate navigation in birds Methods: – Eurasian reed warblers (Acrocephalus scirpaceus) – Migrate between sub-Saharan Africa and Eurasia – Captured birds during spring migration – Displaced half 100 km east From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press Featured Research

Bicoordinate navigation in birds Methods: – Placed in Emlen funnels to measure orientation From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press Featured Research

Bicoordinate navigation in birds Results: – Displaced birds oriented to northwest to compensate for being displaced Conclusion: – Eurasian reed warblers display bicoordinate navigation From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press Featured Research

Homing migration in salmon Salmonids return to natal stream for spawning Use olfaction imprinting and geomagnetic reception From Nordell and Valone, Animal Behavior: Concepts, Methods, and Applications, © 2014 by Oxford University Press