1. Chap. 13 Habitat selection, territoriality and migration 鄭先祐 (Ayo) 教授 國立台南大學 環境與生態學院 生態科學與技術學系 環境生態研究所 + 生態旅遊研究所

2. Ayo 2010 Ethology2 Habitat selection, territoriality, and migration  Models of habitat choice  The ideal free distribution model(IFD) and habitat choice  The IFD model and foraging success  Territoriality  migration

3. Ayo 2010 Ethology3 Habitat selection, territoriality and migration  Habitat selection  The decision-making process is complicated. Mates, food, and predators  Home range  Nomads ( 遊牧民 ), constantly wandering and never returning to the same place with any regularity.  Territory: an area occupied and defended by the owner. To keep intruders outside of this area.

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5. 5 Model of habitat choice  The ideal free distribution model (IFD) and habitat choice  Habitat 1(H1) with resource 1(R1)  Habitat 2 (H2) with resource 2 (R2) At equilibrium, R1/N1 = R2/N2 Resource matching rule  The IFD model and foraging success  Stickleback fish  Mallard ducks

6. Ayo 2010 Ethology6 The matching resource rule in stickleback fish  Experiments ： Each tank had two feeders that distributed food at opposite ends of a tank.  Two treatments food (water fleas) were released from the two feeders in a 5:1 ratio the ratio was 2:1.  Results: the fish distributed themselves under feeders in a ratio similar to the resource matching rule (Fig. 13.2)

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9. 9 In mallard ducks  Two observers who were stationed 20 meters apart throw bread (food) into the pond.  When equal amounts of food were thrown into patches by both observers, ducks quickly distributed themselves in a 1:1 ratio (Fig. 13.4).  When one patch had twice as much bread as the other, the ducks distributed themselves in a 2:1 ratio.  But not all individuals were receiving the same amount of food across patches.

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11. Ayo 2010 Ethology11 The two foraging patches created when two individuals threw bread into a pond from different locations had equal profitability. The dashed line represents the predicted number of ducks at site 1.

12. Ayo 2010 Ethology12 Group Territoriality in chimpanzee  Group territoriality, are defended by a group of unrelated individuals.  Between-group raiding ( 侵襲 )  To be War-like  Larger group eradicating( 消滅 ) a smaller group of chimps.

13. Ayo 2010 Ethology13 Territoriality and learning in Anolis lizards  材料： Anolis aeneus lizards (Stamps, 2001)  Despite numerous experiments manipulating food availability, she did not uncover a clear-cut effect of food availability on territory formation.  Rather, she found that safety from predators and suitable temperature appeared to be the most important attributes of a desirable territory.  How do juvenile lizards determine which territories are suitable with respect to temperature and predation pressure?  Do lizards learn what areas are best from their interaction with other lizards?

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15. Ayo 2010 Ethology15 (A) Juveniles not only spent more time on experimental (E) versus control (C) homesites, but they also arrived at experimental homesites more quickly

16. Ayo 2010 Ethology16 (B) Juvenile lizards were drawn to experimentally manipulated homesites (E) over control (C) homesites.

17. Ayo 2010 Ethology17 Territoriality and learning in Anolis lizards  Conspecific cueing hypothesis  A juvenile would be allowed to observe two very similar territories, one that was currently occupied and one that was vacant.  When given the choice between these two areas, with the territory owner now removed, juveniles showed a strong preference for the previously inhabited area.  The other juveniles displayed no preference for the previously occupied territory.  A strong visual component to conspecific cueing in A. aeneus.

18. Ayo 2010 Ethology18 Territory owners and satellites in pied wagtails  Territory ownership often requires constant vigilance ( 警戒 ) against both predators and conspecific intruders.  Owners, satellites, and territory defense in pied wagtails.

19. Ayo 2010 Ethology19 Fig. 13.8 pied wagtails and food search.  Pied wagtails systemically search for food on their territories along riverbanks  (A) a single bird can complete a circuit of the riverbank in 40 minutes and gets all the food it finds  (B) when a territory is shared by two birds, however, the circuit is divided up as well, and so each bird primarily gets the food that it finds in its 20 minutes of walking the riverbank.

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21. Ayo 2010 Ethology21 Owners often allow “satellite” individuals to forage on territories. Some territorial defense is provided by the satellites.

22. Ayo 2010 Ethology22 How to keep a territory in the family  Fig. 13.9 Scrub jays and territories.  Territories are “inherited” across generations, leading to the establishment of “family dynasties”.  Territory size is increased through a process called budding.

23. Ayo 2010 Ethology23 Migration and navigation  Navigation  The sun compass  By the stars  The Earth ’ s magnetic field  The heritability of migratory restlessness  Defense against parasites  Phylogeny

24. Ayo 2010 Ethology24 (A)Here we see migration in geese (B) gnu.

25. Ayo 2010 Ethology25 Migration and the sun compass  Monarch butterflies migrating each year fro North America in the mountain ranges of central Mexico.  During their annual migration, the branches of trees have been known to collapse from the weight of too many butterflies (Fig. 13.11).  Monarch butterflies traverse up to 6,000 miles on their migratory trip, and they almost always navigate successfully without getting lost, even on their first migration.  To examine the role of solar navigation in the Monarch butterflies.  ran a classic “ clock-shift ” experiment

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27. Ayo 2010 Ethology27 Monarch butterflies  Raised two group of monarch butterflies in a laboratory  One group was slowly shifted the butterflies ’ body clocks bake six hours.  The second group, were not shifted  During the period of autumn migration, released them  The control butterflies headed south  The clock-shifted butterflies, however, flew almost due west. (Fig.12.12)

28. Ayo 2010 Ethology28 Clock-shifted butterflies fly west Laboratory raised butterflies, who were raised on a normal light-dark cycle Butterflies from natural populations fly south

29. Ayo 2010 Ethology29 The pipe at the bottom of the simulator directed a constant flow of air up toward the butterfly so that it could fly; a video camera was connected to the bottom of the simulator; an encoder was attached to the butterfly from the top of the simulator and was connected to a computer that kept a timed record of all the butterfly’s movements. With this setup, researchers could track the direction the butterfly was orienting toward and whether it was actively flying or gliding.

30. Ayo 2010 Ethology30 Indigo buntings and navigation by the stars  Emlen began by creating funnel-shaped test cages for buntings, at the bottom of which he placed an ink pad.  The cages were constructed such that each time a bunting tried to fly out, the location of its footprint was marked by ink, and so its orientation pattern was easily recorded (Fig. 13.15)

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33. Ayo 2010 Ethology33 The Earth’s magnetic field  Evidence that the magnetic field of the earth is important in migration has been found in a wide diversity of animals, including birds, amphibians, reptiles, and insects.  Bobolink (rice bird) has one of the longest round-trip annual migrations of any animal– 12,400 miles.  These birds spend the summer months in the northern US and Canada and then undertake a migration to South America (primarily Brazil, Paraguay, and Argentina) before they return to the Northern Hemisphere.  Do they use the magnetic field of the earth?

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35. Ayo 2010 Ethology35 Magnetic fields and bobolinks  Bringing birds into a planetarium and projecting the star patterns, also to manipulate the magnetic polarity.  When the visual cues and magnetic polarity provided the same information, the birds oriented in the correct southern direction.  A visual cues were correct, but the magnetic polarity was reversed, birds headed toward the magnetic south. Magnetic cues were critical in the annual round-trip migration. High levels of an iron-rich, magnetically sensitive substance in bobolinks.

36. Ayo 2010 Ethology36 The heritability of migratory restlessness  Compared the onset of migratory activity between laboratory and wild birds in 18 different species.  A strong correlation between onset of migratory activity in both groups of birds, suggesting that the timing of departure for migration may be under genetic control.  Migratory restlessness during the autumn migratory season.  Collected 40 blackcaps from the field  The 10 birds with the latest dates for migratory restlessness were selected and allowed to mate. They produced a total of 26 offspring, and from that group 4 pairs with late-onset migratory birds.  In only two generations, migratory restlessness was delayed for an average of 7.65 days (Fig. 13.17)

37. Ayo 2010 Ethology37 In each generation, German blackcap birds with the latest onset of migratory activity were chosen to breed. In just two generations, the onset of migratory activity was set back more than a week.

38. Ayo 2010 Ethology38 Defense against parasites  The energy expended during long migration can reduce immune responsiveness, making animals more susceptible to disease.  In addition, long-distance migrants also face new parasites and diseases upon arrival at their migratory end point.  Migratory birds should therefore invest more heavily in immune function compared to related resident relatives. (hypothesis)  Test: the bursa of Fabricus was larger in birds from the migratory species, while in nine of thirteen pairwise comparisons. The spleen was larger in birds from the migratory species.

39. Ayo 2010 Ethology39 Phylogeny and migratory behavior  In the avian family Motacillidae  Evolutionary precursor model of migration  Migration will be associated with species that live in open or edge habitats (so-called non buffered areas) rather than species that live in forests (buffered areas). The pied wagtail The golden pipitThe yellow-throated longclaw

40. Ayo 2010 Ethology40 Evolutionary precursor model  49 spp. In the Motacillidae family  As either “ migratory ” or “ sedentary ”, and their habitat as either open/edge or forest.  They found that  there was no association between habitat in terms of open/edge versus forest  Species that were associated with open/edge habitats were no more likely to migrate than were species that lived in the forest.  Species that lived at higher altitudes were much more likely to migrate than species that live at low altitudes.

41. Ayo 2010 Ethology41 Interview with Dr. Judy Stamps  Of all the systems you could have chosen to study territoriality, how did you end up working with small lizards?  Dr. Judy Stamps is a professor at the University of California at Davis. Her long-term work on lizards and territoriality has produced fundamentally new ideas on both territoriality and the role of learning in territory formation.

42. Ayo NUTN website: http://myweb.nutn.edu.tw/~hycheng/ 問題與討論