Behavioral ecology studies animal behavior from both proximate and ultimate perspectives. Proximate questions examine the mechanisms and stimuli that trigger behaviors, while ultimate questions address evolutionary significance. Many behaviors have genetic components and are influenced by environment. Behaviors like communication, learning, and mating systems can evolve through natural selection if they influence survival and reproduction.

1. Chapter 51 Behavioral Ecology

2. Overview: Studying Behavior Humans have probably studied animal behavior For as long as we have lived on Earth As hunters Knowledge of animal behavior was essential to human survival

3. Cranes are birds that have captivated people’s interest Possibly because they are large and their behavior is easily observed Figure 51.1

4. The modern scientific discipline of behavioral ecology Extends observations of animal behavior by studying how such behavior is controlled and how it develops, evolves, and contributes to survival and reproductive success

5. Concept 51.1: Behavioral ecologists distinguish between proximate and ultimate causes of behavior The scientific questions that can be asked about behavior can be divided into two classes Those that focus on the immediate stimulus and mechanism for the behavior Those that explore how the behavior contributes to survival and reproduction

6. What Is Behavior? Behavior Is what an animal does and how it does it Includes muscular and nonmuscular activity Figure 51.2 Dorsal fin Anal fin

7. Learning Is also considered a behavioral process

8. Proximate and Ultimate Questions Proximate, or “how,” questions about behavior Focus on the environmental stimuli that trigger a behavior Focus on the genetic, physiological, and anatomical mechanisms underlying a behavioral act

9. Ultimate, or “why,” questions about behavior Address the evolutionary significance of a behavior

10. Ethology Ethology is the scientific study of animal behavior Particularly in natural environments

11. Mid 20th-century ethologists Developed a conceptual framework defined by a set of questions These questions Highlight the complementary nature of proximate and ultimate perspectives

12. Fixed Action Patterns A fixed action pattern (FAP) Is a sequence of unlearned, innate behaviors that is unchangeable Once initiated, is usually carried to completion

13. A FAP is triggered by an external sensory stimulus Known as a sign stimulus

14. In male stickleback fish, the stimulus for attack behavior Is the red underside of an intruder Figure 51.3a (a) A male three-spined stickleback fish shows its red underside.

15. When presented with unrealistic models As long as some red is present, the attack behavior occurs Figure 51.3b (b) The realistic model at the top, without a red underside, produces no aggressive response in a male three-spined stickleback fish. The other models, with red undersides, produce strong responses.

16. Proximate and ultimate causes for the FAP attack behavior in male stickleback fish Figure 51.4 ULTIMATE CAUSE: By chasing away other male sticklebacks, a male decreases the chance that eggs laid in his nesting territory will be fertilized by another male. BEHAVIOR: A male stickleback fish attacks other male sticklebacks that invade its nesting territory. PROXIMATE CAUSE: The red belly of the intruding male acts as a sign stimulus that releases aggression in a male stickleback.

17. Imprinting Imprinting is a type of behavior That includes both learning and innate components and is generally irreversible

18. Imprinting is distinguished from other types of learning by a sensitive period A limited phase in an animal’s development that is the only time when certain behaviors can be learned

19. An example of imprinting is young geese Following their mother

20. Konrad Lorenz showed that When baby geese spent the first few hours of their life with him, they imprinted on him as their parent

21. There are proximate and ultimate causes for this type of behavior Figure 51.5 BEHAVIOR: Young geese follow and imprint on their mother. PROXIMATE CAUSE: During an early, critical developmental stage, the young geese observe their mother moving away from them and calling. ULTIMATE CAUSE: On average, geese that follow and imprint on their mother receive more care and learn necessary skills, and thus have a greater chance of surviving than those that do not follow their mother.

22. Conservation biologists have taken advantage of imprinting In programs to save the whooping crane from extinction Figure 51.6

23. Concept 51.2: Many behaviors have a strong genetic component Biologists study the ways both genes and the environment Influence the development of behavioral phenotypes Behavior that is developmentally fixed Is called innate behavior and is under strong genetic influence

24. Directed Movements Many animal movements Are under substantial genetic influence These types of movements Are called directed movements

25. Kinesis A kinesis Is a simple change in activity or turning rate in response to a stimulus

26. Sow bugs Become more active in dry areas and less active in humid areas Figure 51.7a Dry open area Moist site under leaf (a) Kinesis increases the chance that a sow bug will encounter and stay in a moist environment.

27. Taxis A taxis Is a more or less automatic, oriented movement toward or away from a stimulus

28. Many stream fish exhibit positive rheotaxis Where they automatically swim in an upstream direction Figure 51.7b Direction of river current (b) Positive rheotaxis keeps trout facing into the current, the direction from which most food comes.

29. Migration Many features of migratory behavior in birds Have been found to be genetically programmed Figure 51.8

30. Animal Signals and Communication In behavioral ecology A signal is a behavior that causes a change in another animal’s behavior Communication Is the reception of and response to signals

31. Animals communicate using Visual, auditory, chemical, tactile, and electrical signals The type of signal used to transmit information Is closely related to an animal’s lifestyle and environment

32. Chemical Communication Many animals that communicate through odors Emit chemical substances called pheromones

33. When a minnow or catfish is injured An alarm substance in the fish’s skin disperses in the water, inducing a fright response among fish in the area Figure 51.9a, b (a) Minnows are widely dispersed in an aquarium before an alarm substance is introduced. (b) Within seconds of the alarm substance being introduced, minnows aggregate near the bottom of the aquarium and reduce their movement.

34. Auditory Communication Experiments with various insects Have shown that courtship songs are under genetic control Charles Henry, Lucía Martínez, and ent Holsinger crossed males and females of Chrysoperla plorabunda and Chrysoperla johnsoni, two morphologically identical species of lacewings that sing different courtship songs. EXPERIMENT SONOGRAMS Chrysoperla plorabunda parent Vibration volleys Standard repeating unit Chrysoperla johnsoni parent Volley period crossed with Standard repeating unit The researchers recorded and compared the songs of the male and female parents with those of the hybrid offspring that had been raised in isolation from other lacewings. Volley period

35. The F 1 hybrid offspring sing a song in which the length of the standard repeating unit is similar to that sung by the Chrysoperla plorabunda parent, but the volley period, that is, the interval between vibration volleys, is more similar to that of the Chrysoperla johnsoni parent. RESULTS The results of this experiment indicate that the songs sung by Chrysoperla plorabunda and Chrysoperla johnsoni are under genetic control. CONCLUSION Standard repeating unit Volley period F 1 hybrids, typical phenotype

36. Genetic Influences on Mating and Parental Behavior A variety of mammalian behaviors Are under relatively strong genetic control

37. Research has revealed the genetic and neural basis For the mating and parental behavior of male prairie voles Figure 51.11

38. Concept 51.3: Environment, interacting with an animal’s genetic makeup, influences the development of behaviors Research has revealed That environmental conditions modify many of the same behaviors

39. Dietary Influence on Mate Choice Behavior One example of environmental influence on behavior Is the role of diet in mate selection by Drosophila mojavensis

40. Laboratory experiments have demonstrated That the type of food eaten during larval development influences later mate choice in females Figure 51.12 William Etges raised a D. mojavensis population from Baja California and a D. mojavensis population from Sonora on three different culture media: artificial medium, agria cactus (the Baja host plant), and organ pipe cactus (the Sonoran host plant). From each culture medium, Etges collected 15 male and female Baja D. mojavensis pairs and 15 Sonoran pairs and observed the numbers of matings between males and females from the two populations. EXPERIMENT When D. mojavensis had been raised on artificial medium, females from the Sonoran population showed a strong preference for Sonoran males (a). When D. mojavensis had been raised on cactus medium, the Sonoran females mated with Baja and Sonoran males in approximately equal frequency (b). RESULTS The difference in mate selection shown by females that developed on different diets indicates that mate choice by females of Sonoran populations of D. mojavensis is strongly influenced by the dietary environment in which larvae develop. CONCLUSION 100 75 50 25 0 Artificial Organ pipe cactus Agria cactus Culture medium With Baja males With Sonoran males (b) Proportion of matings by Sonoran females (a)

41. Therese Markow and Eric Toolson proposed That the physiological basis for the observed mate preferences was differences in hydrocarbons in the exoskeletons of the flies Figure 51.13

42. Social Environment and Aggressive Behavior Cross-fostering studies in California mice and white-footed mice Have uncovered an influence of social environment on the aggressive and parental behaviors of these mice

43. Influence of cross-fostering on male mice Table 51.1

44. Learning Learning is the modification of behavior Based on specific experiences Learned behaviors Range from very simple to very complex

45. Habituation Habituation Is a loss of responsiveness to stimuli that convey little or no information

46. Spatial Learning Spatial learning is the modification of behavior Based on experience with the spatial structure of the environment

47. In a classic experiment, Niko Tinbergen Showed how digger wasps use landmarks to find the entrances to their nests After the mother visited the nest and flew away, Tinbergen moved the pinecones a few feet to one side of the nest. Figure 51.14 CONCLUSION A female digger wasp excavates and cares for four or five separate underground nests, flying to each nest daily with food for the single larva in the nest. To test his hypothesis that the wasp uses visual landmarks to locate the nests, Niko Tinbergen marked one nest with a ring of pinecones. EXPERIMENT Nest When the wasp returned, she flew to the center of the pinecone circle instead of to the nearby nest. Repeating the experiment with many wasps, Tinbergen obtained the same results. RESULTS The experiment supported the hypothesis that digger wasps use landmarks to keep track of their nests. Nest No Nest

48. Cognitive Maps A cognitive map Is an internal representation of the spatial relationships between objects in an animal’s surroundings

49. Associative Learning In associative learning Animals associate one feature of their environment with another

50. Classical conditioning is a type of associative learning In which an arbitrary stimulus is associated with a reward or punishment Figure 51.15 Before stimulus Influx of water alone Influx of alarm substances Influx of pike odor Day 1 Day 3 Control group Control group Experimental group Experimental group Relative activity level

51. Operant conditioning is another type of associative learning In which an animal learns to associate one of its behaviors with a reward or punishment Figure 51.16

52. Cognition and Problem Solving Cognition is the ability of an animal’s nervous system To perceive, store, process, and use information gathered by sensory receptors

53. 7 Problem solving can be learned By observing the behavior of other animals Figure 51.17

54. Genetic and Environmental Interaction in Learning Genetics and environment can interact To influence the learning process

55. Concept 51.4: Behavioral traits can evolve by natural selection Because of the influence of genes on behavior Natural selection can result in the evolution of behavioral traits in populations

56. Behavioral Variation in Natural Populations When behavioral variation within a species Corresponds to variation in the environment, it may be evidence of past evolution

57. Variation in Prey Selection Differences in prey selection in populations of garter snakes Are due to prey availability and are evidence of behavioral evolution Figure 51.18a, b (a) A garter snake ( Thamnophis elegans ) (b) A banana slug ( Ariolimus californicus ); not to scale

58. Variation in Aggressive Behavior Funnel spiders living in different habitats Exhibit differing degrees of aggressiveness in defense and foraging behavior Figure 51.19 60 Population 50 40 30 20 10 0 Time to attack (seconds) Field Lab-raised generation 1 Lab-raised generation 2 Desert grassland population Riparian population

59. Experimental Evidence for Behavioral Evolution Laboratory and field experiments Can demonstrate the evolution of behavior

60. Laboratory Studies of Drosophila Foraging Behavior Studies of Drosophila populations raised in high- and low-density conditions Show a clear divergence in behavior linked to specific genes Figure 51.20 14 12 10 8 6 2 0 Average path length (cm) 4 L1 L2 L3 H1 H2 H3 H4 H5 D. Melanogaster lineages Low population density High population density

61. Migratory Patterns in Blackcaps Field and laboratory studies of Blackcap birds Have documented a change in their migratory behavior

62. Birds placed in funnel cages Left marks indicating the direction they were trying to migrate Figure 51.21a (a) Blackcaps placed in a funnel cage left marks indicating the direction in which they were trying to migrate.

63. Migratory orientation of wintering adult birds captured in Britain Was very similar to that of laboratory-raised birds Figure 51.21b (b) Wintering blackcaps captured in Britain and their laboratory-raised offspring had a migratory orientation toward the west, while young birds from Germany were oriented toward the southwest. N E S W Adults from Britain and F 1 offspring of British adults N E S W Young from SW Germany Mediterranean Sea BRITAIN GERMANY

64. Concept 51.5: Natural selection favors behaviors that increase survival and reproductive success The genetic components of behavior Evolve through natural selection Behavior can affect fitness Through its influence on foraging and mate choice

65. Foraging Behavior Optimal foraging theory Views foraging behavior as a compromise between the benefits of nutrition and the costs of obtaining food

66. Energy Costs and Benefits Reto Zach Conducted a cost-benefit analysis of feeding behavior in crows The crows eat molluscs called whelks But must drop them from the air to crack the shells

67. Zach determined that the optimal flight height in foraging behavior Correlated with a fewer number of drops, indicating a trade-off between energy gained (food) and energy expended Figure 51.22 Total flight height (number of drops  drop height) Height of drop (m) 60 50 40 30 20 10 0 Average number of drops 2 3 5 7 15 Average number of drops Drop height preferred by crows 125 100 25 75 50 Total flight height

68. In bluegill sunfish Prey selection behavior is related to prey density Figure 51.23 Low prey density High prey density 33% 33% 33% 32.5% 32.5% 35% 2% 40% 57% 100% 50% 35% 14% 33% 33% 33% Small prey Medium prey Large prey Small prey Medium prey Large prey Small prey Medium prey Large prey Percentage available Predicted percentage in diet Observed percentage in diet Large prey at far distance Small prey at middle distance Small prey at close distance

69. Risk of Predation Research on mule deer populations Has shown that predation risk affects where the deer choose to feed Figure 51.24 70 60 40 30 20 10 0 Predation occurrence (%) 50 Relative deer use Relative deer use Predation risk Open Forest edge Habitat Forest interior 0 5 10 15 20

70. Mating Behavior and Mate Choice Mating behavior Is the product of a form of natural selection call sexual selection

71. Mating Systems and Mate Choice The mating relationship between males and females Varies a great deal from species to species

72. In many species, mating is promiscuous With no strong pair-bonds or lasting relationships

73. In monogamous relationships One male mates with one female Figure 51.25a (a) Since monogamous species, such as these trumpeter swans, are often monomorphic, males and females are difficult to distinguish using external characteristics only.

74. In a system called polygyny One male mates with many females The males are often more showy and larger than the females Figure 51.25b Among polygynous species, such as elk, the male (left) is often highly ornamented. (b)

75. In polyandrous systems One female mates with many males The females are often more showy than the males Figure 51.25c (c) In polyandrous species, such as these Wilson’s phalaropes, females (top) are generally more ornamented than males.

76. The needs of the young Are an important factor constraining the evolution of mating systems The certainty of paternity Influences parental care and mating behavior

77. In species that produce large numbers of offspring Parental care is at least as likely to be carried out by males as females Figure 51.26 Eggs

78. Sexual Selection and Mate Choice In intersexual selection Members of one sex choose mates on the basis of particular characteristics Intrasexual selection Involves competition among members of one sex for mates

79. Mate Choice by Females Male zebra finches Are more ornate than females, a trait that may affect mate choice by the females Figure 51.27

80. Imprinting of female chicks on males with more ornamentation Affects mate selection as adults Figure 51.28 Experimental Groups Control Group Parents not ornamented Both parents ornamented Males ornamented Females ornamented Results Females reared by ornamented parents or ornamented fathers preferred ornamented males as mates. Females reared by ornamented mothers or nonornamented parents showed no preference for either ornamented or nonornamented males. Males reared by all experimental groups showed no preference for either ornamented or nonornamented female mates.

81. The size of eyestalks in stalk-eyed flies Affects which males the females choose to mate with Figure 51.29

82. Male Competition for Mates Male competition for mates Is a source of intrasexual selection that can reduce variation among males

83. Such competition may involve agonistic behavior An often ritualized contest that determines which competitor gains access to a resource Figure 51.30

84. Morphology affects the mating behavior In isopods of the same species that are genetically distinct Figure 51.31 Large Paracerceis  males defend harems of females within intertidal sponges.      Tiny  males are able to invade and live within large harems.  males mimic female morphology and behavior and do not elicit a defensive reponse in  males and so are able to gain access to guarded harems.

85. Applying Game Theory Game theory evaluates alternative behavioral strategies in situations Where the outcome depends on each individual’s strategy and the strategy of other individuals

86. Mating success of male side-blotched lizards Was found to be influenced by male polymorphism and the abundance of different males in a given area Figure 51.32

87. Concept 51.6: The concept of inclusive fitness can account for most altruistic social behavior Many social behaviors are selfish Natural selection favors behavior That maximizes an individual’s survival and reproduction

88. Altruism On occasion, some animals Behave in ways that reduce their individual fitness but increase the fitness of others This kind of behavior Is called altruism, or selflessness

89. In naked mole rat populations Nonreproductive individuals may sacrifice their lives protecting the reproductive individuals from predators Figure 51.33

90. Inclusive Fitness Altruistic behavior can be explained by inclusive fitness The total effect an individual has on proliferating its genes by producing its own offspring and by providing aid that enables close relatives to produce offspring

91. Hamilton’s Rule and Kin Selection Hamilton proposed a quantitative measure For predicting when natural selection would favor altruistic acts among related individuals

92. The three key variables in an altruistic act are The benefit to the recipient The cost to the altruist The coefficient of relatedness

93. The coefficient of relatedness Is the probability that two relatives may share the same genes Figure 51.34 Parent A Parent B  OR Sibling 1 Sibling 2 1 / 2 (0.5) probability 1 / 2 (0.5) probability

94. Natural selection favors altruism when the benefit to the recipient Multiplied by the coefficient of relatedness exceeds the cost to the altruist This inequality Is called Hamilton’s rule

95. Kin selection is the natural selection That favors this kind of altruistic behavior by enhancing reproductive success of relatives

96. An example of kin selection and altruism Is the warning behavior observed in Belding’s ground squirrels Male Female Age (months) Mean distance moved from natal burrow (m) 300 200 100 0 0 2 3 4 12 13 14 15 25 26 Figure 51.35

97. Reciprocal Altruism Altruistic behavior toward unrelated individuals Can be adaptive if the aided individual returns the favor in the future This type of altruism Is called reciprocal altruism

98. Social Learning Social learning Forms the roots of culture Culture can be defined as a system of information transfer through observation or teaching That influences the behavior of individuals in a population

99. Mate Choice Copying Mate choice copying Is a behavior in which individuals in a population copy the mate choice of others

100. This type of behavior Has been extensively studied in the guppy Poecilia reticulata Figure 51.36 Male guppies with varying degrees of coloration Control Sample Female guppies prefer males with more orange coloration. Experimental Sample Female model engaged in courtship with less orange male Female guppies prefer less orange males that are associated with another female.

101. Social Learning of Alarm Calls Vervet monkeys Produce a complex set of alarm calls

102. Infant monkeys give undiscriminating alarm calls at first But learn to fine-tune them by the time they are adults Figure 51.37

103. No other species Comes close to matching the social learning and cultural transmission that occurs among humans Figure 51.38

104. Evolution and Human Culture Human culture Is related to evolutionary theory in the distinct discipline of sociobiology

105. Human behavior, like that of other species Is the result of interactions between genes and environment However, our social and cultural institutions May provide the only feature in which there is no continuum between humans and other animals