1. 行為生態學 Behavioural Ecology 鄭先祐 (Ayo) 臺南大學 環境生態研究所 教授 Japalura@hotmail.com

2. 行為生態學 2 Textbook Krebs, J. R. & N. B. Davies (1997) Behavioural Ecology : An Evolutionary Approach. 4 th ed. Blackwell Science. Caro, Tim (ed.) (1998) Behavioral Ecology and Conservation Biology. Oxford University Press, Inc. Dugatkin, L. A. (ed.) (2001) Model systems in Behavioral Ecology. Princeton University Press. Hall, M. and T. Halliday (eds) (1998) Behaviour and Evolution. The Open University. 尚玉昌 (2003) 行為生態學。五南圖書。 教材： Ayo 台南站 http://mail.nutn.edu.tw/~hycheng/http://mail.nutn.edu.tw/~hycheng/

3. 行為生態學 3 Preface The studies of mechanisms and how they control and constrain the adaptive behaviour of individuals. ( 個體適應行為的機制、如何調控與有何限制？ ) Molecular phylogenies 運用於行為和生活史演化研究。 Ask everyone to provide a review of the main ideas, and empirical data to test them, and to focus on current controversies and unsolved questions. The need for a fifth edition in another 7 years ’ time. 1978 → 1984 → 1991 → 1997 → ？

4. 行為生態學 4 Contents Part I Introduction Part 2 Mechanisms and individual behavior 機制與個體行為 Part 3 From Individual behavior to social systems 從個體行為至社會體系 Part 4 Life histories, Phylogenies and populations 生活史，親緣和族群

5. 行為生態學 5 Contents Part I Introduction Chap.1 The evolution of Behavioural Ecology Part 2 Mechanisms and individual behaviour Chap.2 Sensory systems ( 感覺系統 )and Behaviour Chap.3 The Ecology of Information use( 資訊使用 ) Chap. 4 Recognition systems ( 辨識系統 ) Chap. 5 Managing time and energy ( 時間與能量 ) Chap. 6 Sperm competition and mating systems ( 精 子競爭與交配體系 )

6. 行為生態學 6 Part I Introduction Part 2 Mechanisms and Individual behaviour Part 3 From Individual behaviour to social systems Chap. 7 The evolution of animal signals ( 動物訊息的 演化 ) Chap. 8 Sexual selection ( 性擇 ) and mate choice Chap. 9 Sociality ( 社會性 ) and Kin Selection in insects Chap. 10 Predicting family dynamics ( 家庭動態 ) in social vertebrates Chap. 11 The ecology of relationships ( 關係生態學 ) Chap. 12 The Social gene ( 社會基因 ) Part 4 Life histories, Phylogenies and populations

7. 行為生態學 7 Part 1 Introduction Part 2 Mechanisms and individual behaviour Part 3. From individual behaviour to social systems Part 4. Life histories, phylogenies and populations Chap. 13 Adaptation of Life histories ( 生活史的適應 ) Chap. 14 The phylogenetic foundations ( 親緣基礎 )of Behavioural ecology Chap. 15 Causes and consequences of population structure ( 族群結構的因果 ) Chap. 16 Individual behaviour, populations and conservation ( 個體行為，族群與保育 )

8. 行為生態學 8 Chap. 1 The Evolution ( 演變史 ) of Behavioural Ecology 1.1 Observations and questions ( 觀察與問題 ) 1.2 Tinbergen ’ s four questions ( 四個問題 ) 1.3 Ecology and behaviour 1.4 Economic models( 經濟模型 ) of behaviour 1.5 Evolutionarily stable strategies ( 演化穩定策略 ) 1.6 Kinship, social evolution and breeding systems 1.7 Critical views ( 批評的觀點 ) 1.8 Looking ahead ( 未來方向 )

9. 行為生態學 9 1.1 Observations and questions All natural observations begin with a question. At first our curiosity may be satisfied simply by knowing the species name. Then we may want to discover what it is doing and to understand why it is behaving in a particular way. For students of behavioural ecology, a whole host of questions come to mind as they observe.

10. 行為生態學 10 觀察 a starling bird Searching in the grass for food The starling walks along, pausing every now and then to probe into the ground. Sometimes it succeeds in finding a prey item, such as a beetle larva and eventually, when it has collected several prey, it flies back to the nest to feed its hungry brood.

11. 行為生態學 11 A whole host of questions The first set of questions concern the way the bird feeds. Why has it chosen that particular place( 特定地區 ) to forage? Why is it alone rather than in a flock? What determines its choice of foraging path? Does it collect every item of food it encounters or is it selective for prey type or size: What influences its decision to stop collecting food and fly back to feed its chicks:

12. 行為生態學 12 Another set of questions emerges when we follow the starling back to the nest. Why has it chosen this site? Why this brood size? How do the two parent starlings come to an agreement over how much work each puts into offspring care? Why are the chicks begging so noisily? Are they each simply signaling their own degree of hunger or are they competing for food? What determines how much effort it puts into reproduction versus it own maintenance, the factors influencing the timing of its seasonal activities, its choice of mate, and so on.

13. 行為生態學 13 Behavioural ecology provides a framework for answering these kinds of questions. It combines ideas from evolution, ecology and behaviour and has emerged from five schools of thought, developed primarily in the 1960s and early 1970s. 1.Tinbergen ’ s four questions 2.Ecology and behaviour 3.Economic models of behaviour 4.Evolutionarily stable strategies 5.Kinship, social evolution and breeding systems

14. 行為生態學 14 1.2 Tinbergen ’ s four questions Tinbergen(1963) showed that there are four ways of answering the question ‘ why? ’ in biology. In terms of function ( 功能 ) In terms of causation (cue 因素 ) In terms of development ( 發展 ) In terms of evolutionary history ( 演化歷史 )

15. 行為生態學 15 1.3 Ecology and behaviour To correlate difference between species in behaviour with differences in ecological factors ( 生態因素 ), such as habitat, food and predation ( 棲息地、食物和掠食 ). Cullen(1957), a student of Tinbergen, interpreted the reduced anti-predator behaviour of kittiwake gulls, compared to the ground-nesting gulls, in relation to their safer nest sites on steep cliffs. Winn(1958), linked the reproductive behaviour of 14 species of darter fish to their ecology. Wilson (1959), related the colony size and structure of dacetine ants to their feeding habits.

16. 行為生態學 16 Crook ’ s (1964) study of weaver birds. How differences between species in food and predator pressure affect a whole host of adaptations, including nesting dispersion (colonies vs. territories), feeding behaviour (solitary vs. flock) and mating systems (monogamy vs. polygamy). Crook & Gartlan (1966), primates Crook (1965), Lack (1968), other bird species Jarman (1974), ungulates Krunk (1975), carnivores Fricke (1975), coral reef fish Multivariate statistics ( 多變項分析 ) and phylogenies

17. 行為生態學 17 1.4 Economic models of behaviour The pioneer in the use of mathematical models in ecology to quantify trade-offs was Robert MacArthur, who first applied the idea of optimal choice I the context of foraging behaviour (e.g. MacArthur & Pianka, 1966; MacArthur, 1972). Classic early studies include work by Schoener (1971) and Charnov (1976) on prey choice and patch choice by foragers and Parker ’ s(1970) study of copulation time in the yellow dungfly.

18. 行為生態學 18 1.5 Evolutionarily stable strategies Fisher ’ s (1930) explanation for why parents expend equal rexource on male and female progeny. Hamilton ’ s (1967) analysis of stable sex ratios under local mate competition. Parker ’ s (1970) field study of how male dungflies distribute themselves across different mating sites. Fretwell and Lucas (1970), on habitat choice by birds. Maynard Smith ’ s (1972, 1982) concept of ESS, is now widely accepted as the way of analyzing decision making.

19. 行為生態學 19 1.6 Kinship, social evolution and breeding systems Wynne-Edwards(1962) proposed that social behaviour was an adaptation for regulating animal populations. Many ethologists also used group selection to explain behaviour. Individual benefit vs. group benefit Multiple mating (sperm competition) DNA fingerprinting

20. 行為生態學 20 1.7 critical views 1.7.1 Determinism Genetic determinism 1.7.2 Panglossianism Gould and Lewontin (1979), The Panglossian paradigm, referring to Dr. Pangloss, who took the view that everything was always for the best. 1.7.3 Anthropomorphism Honest signaling

21. 行為生態學 21 1.8 Looking ahead Never prophesy( 預言 ), especially about the future. However, the changes signaled by the new emphases in the Behavioural Ecology coincide with a change in the nature of the subject. A new form of integrated study With powerful techniques, from gene splicing to magnetic resonance imaging

22. 行為生態學 22 http://mail.nutn.edu.tw/~hycheng 問題與討論