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Questions in the study Questions in the study of foraging behavior of foraging behavior ----------------------------- ----------------------------- ow do animals select prey? hen should animals leave one feeding site and go on to another? How should animals divide their time between food-gathering and other activities?

Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

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Page 1: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

Questions in the studyQuestions in the studyof foraging behaviorof foraging behavior

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1. How do animals select prey?

2. When should animals leave one feedingsite and go on to another?

3. How should animals divide their timebetween food-gathering and other activities?

Page 2: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

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LOAD

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TIMETravel Time Searching Time

10

Page 3: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

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LOAD

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TIME

Travel Time Searching Time

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X1

Y1

Page 4: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

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LOAD

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TIME

Travel Time Searching Time

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X2

Y2

Load/Time is maximized at X2,Y2

Page 5: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

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LOAD

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Travel Time Searching Time

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X1 X2

Short optimum

Long optimum

Page 6: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

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Load

Round trip travel time

Page 7: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

STEPS IN ANALYSISSTEPS IN ANALYSIS

1. Determine relevant variables throughobservation

2. Establish (you think) how thesevariables interact

3. Make predictions

4. Test

Page 8: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

Crows feeding on whelks

Select the largest whelks available

Fly roughly 5.5 m high to drop the whelkon the rocks below

If whelk doesn’t break open, select thesame whelk to drop again

Page 9: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and
Page 10: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

Patches

Problem: When to leave a patch?

Page 11: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

Constraints that affect “optimal”foraging behavior

1. Physiological constraints

2. Motivational constraints

3. Ecological constraints

4. Life history constraints

Page 12: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

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Energy constraint

Sodium constraint

Rumenconstraint

Intakeofaquaticplants(gm)

Intake of terrestrial plants (gm)

Page 13: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

Constraints that affect “optimal”foraging behavior

1. Physiological constraints

2. Motivational constraints

3. Ecological constraints

4. Life history constraints

Page 14: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

Food Choice (reward x probability)

reward probability 2 pellets x 1.0 average yield of 2

0 pellets x 0.54 pellets x 0.5 average yield of 2

Page 15: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

Testing Conditions

Starved for 1 hour

Starved for 4 hours

Food Choice (reward x probability)

2 pellets x 1.0 = avg. 2

(0 pellets x 0.5) +(4 pellets x 0.5) = avg. 2

Page 16: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

Testing Conditions

Starved for 1 hour

Starved for 4 hours

Food Choice (reward x probability)

2 pellets x 1.0 = avg. 2

(0 pellets x 0.5) +(4 pellets x 0.5) = avg. 2

Page 17: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

Constraints that affect “optimal”foraging behavior

1. Physiological constraints

2. Motivational constraints

3. Ecological constraints

4. Life history constraints

Page 18: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

Constraints that affect “optimal”foraging behavior

1. Physiological constraints

2. Motivational constraints

3. Ecological constraints

4. Life history constraints

Page 19: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

Life History Evolution

The study of how individuals allocate, throughout life, time and energy to various fundamental activities, such as growth and reproduction

Page 20: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

Life Histories: An inherent trade-off

Investment in any one activity limits an animal’s ability to invest in others.

As applied to reproduction, a parent’s dilemma: investment in any one offspring limits an animal’s ability to invest in others.

Page 21: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

• Growth and Development

• Reproduce Early or Delay

• Clutch Size vs. Clutch Number

• Offspring Size and Offspring Number

• Offspring Size and Parental Care

Components of Life Histories: Where the trade-offs occur

Page 22: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

Life histories: the major questions

Why do organisms age and die?

How many offspring should an individual produce in a given time?

How big should each offspring be?

Page 23: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

Life history traits – characteristics of an individual that influence survival and reproduction

Age at maturity

11 - 20 years 3-6 years

SalmonAfrican elephant

2 months

House Mouse

Page 24: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

Life history traits – characteristics of an individual that influence survival and reproduction

SalmonAfrican elephantHouse Mouse

1 calf every 3-8 years

1,500 to 8,000 eggs

once

5-8 young every month

Number of offspring produced

Page 25: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

Life history traits – characteristics of an individual that influence survival and reproduction

SalmonAfrican elephantHouse Mouse

Number of reproductive events

~3 - 10 1~6-12

Page 26: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

Life history traits – characteristics of an individual that influence survival and reproduction

SalmonAfrican elephantHouse Mouse

Lifespan

60 - 70 years 3-6 years~2 years

Page 27: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

K-strategists:K-strategists:

•long lived•produce few offspring•parental care

Reproductive strategies

Population is controlled by density-dependent

limiting factors - e.g. food

Page 28: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

Gypsy moth caterpillars

r-strategists:•small•short life•no parental care•many offspring

Reproductive strategies

Population is controlled by density-independent limiting factors: weather, pond drying

Page 29: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

2.1m

12-Arm Radial Maze

Page 30: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

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80

40

%Corr.

Delay (hours)

Page 31: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

1

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Page 32: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

Food-storing Birds

Clark’s nutcracker: 33,000 seeds, 7,500 sites

Pinon jays: 22,000 seeds, clumped

Scrub jays: 6,000 seeds

Page 33: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

0 2 4 6 8 10 1250

60

70

80

90

100

% c

orre

ct, 1

st 4

cho

ices

Block (avg of 5 trials)

Nutcracker

Pinyon jay

Scrub jay

Mexican jay

Cache retrieval in corvids

Kamil et al. 1994

Page 34: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

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80%correct

NutcrackersPinyon jaysScrub jaysMexican jays

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Retention interval (min)

Page 35: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

Meadow Vole:Polygynous

Prairie Vole:Monogamous

Femaleranges

Male range

Page 36: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and
Page 37: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

Olson et al. 1995

?

?

Page 38: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

Strict behaviorism:

Any stimulus can, through conditioning, be associated withany response or reinforcer

Learning is a general process phenomenon:All associations are learned equally easilyAll responses are reinforced equally easily

Page 39: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

Biological constraints on learning

Saccharine taste + lights + noise

BECAME SICK SHOCKED

(Garcia & Koelling 1966)

Page 40: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

Biological constraints on learning

Saccharine taste + lights + noise

Avoided saccharine,But no fear of light or noise

Fear reaction to light and noise, but no aversion to saccharine

BECAME SICK SHOCKED

(Garcia & Koelling 1966)

Page 41: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

Summary: Economic decisions

Increasing evidence that animals make “calculations” when foraging

-- make adaptive “choices” among alternative foods-- estimate past rates of return and compare them

with current rates

Page 42: Questions in the study of foraging behavior ----------------------------- 1.How do animals select prey? 2.When should animals leave one feeding site and

Summary: Memory

1. Natural selection has shaped the minds andbehaviors of animals so that they optimize(as near as possible) the exploitation of theirenvironment

-- species differences in memory-- memory of a very specific sort-- species differences in the brain structures

that support memory-- sex differences, too-- differences in the kind of associations that

are formed