1. Migration, Orientation, Movements

2. What is migration? Migration: An annual two-way movement, usually based on seasonality Breeding Area Wintering Area

3. We have learned about migration from many efforts An example of a “watched” migration: the Broad-winged Hawk A medium-sized hawk from Eastern No. America

4. One of first successful attempts to track the migration of a songbird (Swainson’s Thrush) With a radio transmitter From Cochran et al. 1967. Living Bird Tracking Migrating Songbirds

5. How they tracked the migrating thrushes. From Cochran et al. 1967. Living Bird Now use satellite transmitters Photo-sensitive micro data loggers Isotopes

6. Radar Also Provides Much Information Trans Gulf migrants arriving Apr 25, 0031 GMT Trans Gulf migrants leaving Apr 25, 0159 GMT Radar detection of birds arriving near Houston, TX (images from S.A. Gauthreaux, 1999) Use of radar

7. A. Nocturnal exodus of birds from stop-over locations in South Carolina B. Same data superimposed on habitat types from Landsat satellite imagery. Highest bird densities were in hardwood forests, not the predominant pine woodlands. From Gauthreaux & Belser 2003: Auk 120: 266-277.

8. Basic Types of Migration 1.Obligate annual migration Latitudinal migration (neotrops) Elevational 2. Irruptive migration (food highly variable) Rough-legged Hawks Snowy Owls Crossbills Redpolls 3. Partial migration Some individuals of a species migrate while others do not. Adapts to local conditions

9. Resident vs migratory species Life history traits Generally, birds that migrate to the tropics survive the winter better than do those that stay in the temperate zone. TraitTemperate Resident MigrantTropical Resident ProductivityHighModerateLow Adult Survival Low (20- 58% Moderate (50%) High (80- 90 %) Juvenile Survival LowModerateMod - High

10. Differential Migration Sex and age classes may have different migration characteristics Example: Dark-eyed Junco (Slate-colored race) Migrate south to eastern US to Gulf Coast, but migration distances of individual juncos shows different average wintering distributions of males, females, and young. 1. Adult females migrate farthest south 2 young males stay farthest north in Indiana & Ohio. 3. Adult females, young females settle at intermediate latitudes.

11. Fat as fuel for migration Fuels for migration FuelEnergy YieldMetabolic Water (g) Fat38.91.07 Carbohyd.17.60.55 Protein17.20.41

12. Fat as fuel for migration Necessary for long-distance migration Fat yields 2x as much energy & water as carbohydrates & protein Migrants fatten rapidly before migration and can add it quickly while feeding at resting points along the way (called stop-over sites) Stop-over sites are very important for birds crossing vast areas of inhospitable areas (e.g. ocean, large lakes, deserts)

13. Timing of migration Photoperiod Local conditions (food, vegtation changes) Weather patterns (cold, warm fronts)

14. Weather That Prompts Migration In fall, largest flights occur in the southward flow of dry polar air immediately following the passage of a cold front. In spring, major movements occur in warm southerly air flow on the west side of high pressure centers. Once airborne, passerines usually fly downwind regardless of wind direction.

15. From Able, K. (ed.) 1999. Gatherings of Angels

16. When migrating over land, songbirds progress in a series of relatively short flights, up to 200 miles or so, interrupted by 3-5 days of rest. Length of stop-over depends on local weather, how much fat the bird has, and refueling conditions In one October flight, 500,000 ducks flew from eastern Saskatchewan & Manitoba to Lousiana in a little over 1 day! (=1512 miles; mean ground speed 41-50 mph) A flock of Snow Geese flew from James Bay Canada to Louisiana in 60 hours!

17. How Fast Do Migrants Fly? Most songbirds migrate at about 20-30 mph in still air Waterfowl & shorebirds: 30-50 mph Tail winds or head winds can drastically impact bird speed and (potentially) survival.

18. Movements are Remarkable Homing behavior –Homing pigeons that return to their lofts from –Distances of 100’s of miles –Other documented displacements Migration –Latitudinal –Altitudinal

19. Displacement of Selected Seabirds Illustrates Homing Ability

21. Migratory feats A. Arctic shorebirds White-rumped Sandpiper Baird’s Sandpiper

22. Migratory Feats Arctic shorebirds: Stop-over locations

23. Migratory Feats Blackpoll Warbler

24. From Outlaw et al. 2003. Auk 120: 299-310. Long-distance migration in Catharus thrushes

25. Migratory Feats

26. Short-tailed Shearwater

27. Importance of Stopover Sites

28. To Migrate (and move about beyond the range of local cues), Birds: Use a two step process –Consult a map to determine the compass course to their goal –Use a compass to navigate to the goal (Wiltschko and Wiltschko 2009)

29. Use of Map and Compass Compass mechanisms are relatively well known –Magnetic –Sun –Stars –Combinations Maps less so –Geophysical gradients –Landmarks Learned and innate components Recent Review in Wiltschko and Wiltschko (2009)

30. How Do Birds Navigate? Geomagnetism A bar magnet interferes with a pigeon’s ability to return to its loft on overcast days. Solar storms that disrupt the earth’s magnetic field, also disrupted the orientation of Ring-billed Gulls.

31. Birds “see” the inclination of the magnetic field, not its polarity Direction of arrow in He (natural) or H (experimental) does not matter, it is the angle e relative to p that is sensed Birds “see” this by photopigments in the eye. “Radical-pair” model suggests photon absorption leads to formation of a pair of radicals (singlet and triplet) with unpaired electrons. Light triggers this and the ratio of single to triplet states depends on magnetic field. Birds compare singlets to triplets using pigments in eye, which form patterns on the retina.

32. Sun-compass orientation -Not constant moves across sky—requires a clock -Not always available Sun rises and sets rapidly, but is slow to change position around noon --birds use this information --because rate of change varies with location on earth, birds also learn to use this mechanism

33. Polarized Light At sunrise and sunset polarization forms a band at 90° relative to the sun. So, position of sun (and E or W) is noticeable even with clouds.

34. The “Emlen cage” for testing orientation in birds. Star compass

35. Integration of Cues Magnetic field provides a reference system for linking sun position and internal clock with geographic direction Young birds rely on sun compass more than magnetic, older birds adjust misinformation from sun compass (in experiments) by consulting magnetic field

36. More Integration Night migrants set compass with polarized light before star compass is visible Magnetic compass may also be used at this time The magnetic, sun, and star compasses are integrated components of a single complex system that provides directional information

37. Landmarks are a Start to the Map Navigating around the home range, can be expanded to navigate back to a familiar area (homing) –Can be done by route reversal, or better yet by keeping track of direction to home and using compasses to navigate in that direction, then local cues Seems to only be done by very young birds –Different from migration to a new location (first time migrants) –Older birds use a real map, coordinate based, system

38. What about the Map? Gradients used are not known Geomagnetic field Olfaction Gravity Sound (ocean roar)

39. Compass and Map Innate (direction to migrate) and learned components are involved

40. Why do birds migrate ? Temperate Origin: birds escape from the inhospitable climates in the north which negatively impact a bird’s life history. Tropical Origin: migrants aggressively exploit favorable opportunities. Neotropical migrants are essentially tropical species that temporarily exploit the long days & abundant insects of high latitude summers. (rather than as temperate birds that tolerate the tropics to escape the northern winter)

41. Intraspecific, genetic Approach Population founded from South, Likely then developed migration Tropical origin of migration behavior suggested Time of divergence suggests southern populations were original and northern expansion followed retreat of glaciers about 18000 years ago (Mila et al 2006)

42. Migration May Constrain Brain Size Large brains are metabolically expensive Weight counts when you migrate Sedentary birds typically have larger brains than migrants Which came first? Big brain > behavioral flexibility > no need to migrate? Put another way, Small brain > not flexible > need to migrate OR Sedentary behavior > big brains > to enable behaviorally flexible strategies that promote survival (Pravosudov et al. 2007) Compare timing of evolution of migration and brain size within 1 species, the White-crowned sparrow

43. Big brains came after shift to sedentary habit, not as a precursor Recently evolved, sedentary subspecies (Z. i. nuttalli) Older, migratory subspecies Side bar: Migratory subspecies has larger hippocampus and reduced size of remainder of telencephalon --selection works to fashion different parts of the brain

44. Hazards of Migration Birds killed by television transmitter at Madison, Wisconsin Sept. 7-8 and 13-14, 2005. Tower is 1,000 ft tall.

45. Exploring Movements of Residents

46. Nest predators with hypothesized forest edge dependency

47. Estimation of the Home Range

48. Marzluff et al. 2001

49. Univariate Analysis of Land Cover 0.05 Average Use Late Seral Med. Seral Young Seral Clear Cut Human Activity High Use of Human Occupied Areas is Confirmed N=2422122010

50. Utilizes Rempel’s “Patch Analyst” and FRAGSTATS Landscape Pattern is Deduced From Land Cover Land Cover Contrast-weighted Edge

51. Landscape Metrics We selected 4 variables that were minimally correlated –Number of Patches –Contrast-weighted Edge Density –Juxtaposition –Mean Shape Index High Low

52. = + Use is Related to Cover and Pattern Variables Pixel by Pixel Using Multiple Regression

53. Multiple Regression Produces a Resource Utilization Function Relative Use = 1.32 - 0.14 (Mature Forest) - 0.29 (Clear cut) + 0.09 (Number of Patches) + 0.005 (Contrast-weighted Edge) - 0.002 (Patch Juxtaposition) + 0.14 (Patch Shape) Use is a CONTINUOUS MEASURE Use:Availability Can Be Incorporated to Any Degree Desired

54. Relative Importance of Resources at the Population Level (n = 25) Resource AttributeMean Standardize d  95% Confidence Interval P (   # of jays with use significantly associated with attribute + - Number of Patches +0.11* -0.57 – 0.28 0.1914*9 Contrast-weighted Edge +0.06* -0.13 – 0.26 0.5010*9 Mature Forest -0.05 -0.18 – 0.08 0.45128 Clear cut -0.04 -0.17 – 0.09 0.5169 Interspersion – Juxtaposition -0.01* -0.14 – 0.16 0.87118* Patch Shape Index +0.01* -0.11 – 0.14 0.849*12 * Use in direction predicted if jays select for edgy, fragmented areas within their home range Decreasing Importance

55. Relative Importance of Resources at the Population Level (n = 25) Resource AttributeMean Standardize d  95% Confidence Interval P (   # of jays with use significantly associated with attribute + - Number of Patches +0.11* -0.57 – 0.28 0.1914*9 Contrast-weighted Edge +0.06* -0.13 – 0.26 0.5010*9 Mature Forest -0.05 -0.18 – 0.08 0.45128 Clear cut -0.04 -0.17 – 0.09 0.5169 Interspersion – Juxtaposition -0.01* -0.14 – 0.16 0.87118* Patch Shape Index +0.01* -0.11 – 0.14 0.849*12 * Use in direction predicted if jays select for edgy, fragmented areas within their home range Decreasing Importance Greater use of areas with many patches and edge as expected Pattern more important than type of vegetation

56. Relative Importance of Resources at the Population Level (n = 25) Resource AttributeMean Standardize d  95% Confidence Interval P (   # of jays with use significantly associated with attribute + - Number of Patches +0.11* -0.57 – 0.28 0.1914*9 Contrast-weighted Edge +0.06* -0.13 – 0.26 0.5010*9 Mature Forest -0.05 -0.18 – 0.08 0.45128 Clear cut -0.04 -0.17 – 0.09 0.5169 Interspersion – Juxtaposition -0.01* -0.14 – 0.16 0.87118* Patch Shape Index +0.01* -0.11 – 0.14 0.849*12 * Use in direction predicted if jays select for edgy, fragmented areas within their home range Decreasing Importance Population Not Consistent in Use of Resources

57. Correlates of  s Can Indicate Why Effects Are Not Greater Use of edges is related to proximity to human activity (F (1,24) = 5.4; P=0.04) –Anthropogenic food available in these settings –Rate of predation on other birds’ nests is highest closest to such edges in our study area

58. Learning how Steller’s Jays forage Vigallon and Marzluff (2005)

59. Incidental Predation