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Essentials of Ecology 鄭先祐 (Ayo) 國立臺南大學 環境與生態學院 生物科技學系 生態學 (2008) 教科書 • Essentials of Ecology, 3rd Edition • Townsend, Begon, Harper • March 2008, Wiley-Blackwell • ISBN: 978-1-4051-5658-5 Website: http://www.blackwellpublishing.com/townsend/ 2 About the Authors • Colin R. Townsend is the Director of the Ecology, Conservation and Biodiversity Research Group at the University of Otago. He has published over 150 ecology papers and books, including the bestselling and award-winning Ecology, 4th Edition, with co-authors Michael Begon and John L. Harper. He is also co-editor of the international journal Freshwater Biology. • Michael Begon (University of Liverpool) (animal population ecology) • John L. Harper (University of Exeter) (plant population biology) 3 • Colin R. Townsend (University of Otago) (conservation biologist) • Michael Begon is Professor of Ecology in the School of Biological Sciences at the University of Liverpool. He has authored several books in addition to Ecology and around 150 research articles. His current research interests are focused on the population ecology of pathogens in wildlife populations. • John L. Harper is an award-winning biologist and a leading figure in plant population biology. He has served as a council member of the Fellowship of the Royal Society and received the Darwin Arward in 1990. He is also the author of Population Biology of Plants and a co-author of Ecology. 4 Contents • Preface • Acknowledgments • Part I: Introduction: (簡介) • Part II: Conditions and Resources: (情境與資源) • Part III: Individuals, Populations, Communities and Ecosystems: (個體、族群、群落與生態體系) • Part IV: Applied Issues in Ecology: (應用生態學) • References • Index 5 Contents • Part I: Introduction: – 1. Ecology and how to do it (簡介) – 2. Ecology‘s evolutionary backdrop (演化背景) • Part II: Conditions and Resources: – 3. Physical conditions and the availability of resources (物理情境與資源的供應力) – 4. Conditions, resources and the world‘s communities (情境、資源和世界群落) • Part III: Individuals, Populations, Communities and Ecosystems: • Part IV: Applied Issues in Ecology: 6 Contents • Part I: Introduction: • Part II: Conditions and Resources: • Part III: Individuals, Populations, Communities and Ecosystems: – – – – – – – 5. Birth, death and movement (出生、死王和遷移) 6. Interspecific competition (種間競爭) 7. Predation, grazing and disease(掠食、草食和疾病) 8. Evolutionary ecology (演化生態學) 9. From populations to communities (族群到群落) 10. Patterns in species richness (物種豐富度) 11. The flux of energy and matter through ecosystems (生態体系的能量與物質流動) • Part IV: Applied Issues in Ecology: 7 Contents • • • • • Preface Acknowledgments Part I: Introduction: Part II: Conditions and Resources: Part III: Individuals, Populations, Communities and Ecosystems: • Part IV: Applied Issues in Ecology: – 12. Sustainability (可持續力) – 13. Habitat degradation (棲地衰退) – 14. Conservation (保育) • References • Index 8 Ecology: concepts & Applications Molles, M. C. Jr. (2008) 4th ed. McGraw-Hill Co., Inc. • • • • Introduction: What Is Ecology? (chap.1) Section I Natural History (chap. 2, 3) Section II Individuals (chap. 4,5,6,7) Section III Population Ecology (chap. 8.9.10.11.12) • Section IV Interactions (chap.13,14,15) • Section V Communities and Ecosystems (chap.16,17,18,19,20) • Section VI Large-Scale Ecology (chap. 21,22,23) 9 Fundamentals of Ecology By Odum, E. P. and G. W. Barrett (2005) 1. The Scope of Ecology 2. The Ecosystem 3. Energy in Ecological systems 4. Biogeochemical Cycles 5. Limiting and Regulatory Factors 6. Population Ecology 7. Community Ecology 10 Fundamentals of Ecology 8. Ecosystem Development 9. Landscape Ecology 10. Regional Ecology: Major Ecosystem Types and Biomes 11. Global Ecology 12. Statistical Thinking for Students of Ecology • Glossary • References • Credits • index 11 Odum, E. P. (1997) Ecology: A bridge between Science and Society. Sinauer Associates, Inc. Brief Contents • • • • • • • • • • 緒言:Apollo 13號的航行 第一章 維生環境 第二章 組織的層次 第三章 生態體系 第四章 能量學 第五章 生存的物質循環和物理環境 第六章 族群與群區生態學 第七章 發展與演化 第八章 世界主要生態體系的類型 結語:從年輕至成熟的轉變 12 Preface • 這本書的目的: – 分享自然複雜世界的美妙 – 面對受傷害的自然,我們大家都需要有生態學 識,才有能力為保護自然辯護,以及貢獻於化 解生態問題。 • 這本書的適用: – A semester-long beginners course • 完整的生態學:可參閱 – Ecology: from individuals to ecosystems (Begon, Townsend & Harper, 4th ed. 2005) 13 參考書 • Ecology: From Individuals to Ecosystems, 4th Edition • Begon, Townsend, Harper • • • • ISBN: 978-1-4051-1117-1 Paperback 752 pages August 2005, Wiley-Blackwell 14 Essentials of Ecology 3rd. Ed. Chap. 01 Ecology and how to do it 鄭先祐 (Ayo) 國立臺南大學 環境與生態學院 生物科技學系 生態學 (2008) Contents Part I: Introduction: • Chap.1. Ecology and how to do it – 1.1 Introduction – 1.2 Scales, diversity and rigor (尺度、多樣性、精確) – 1.3 Ecology in practice (生態學的研究) • Chap.2. Ecology's evolutionary backdrop Part II: Conditions and Resources: Part III: Individuals, Populations, Communities and Ecosystems: Part IV: Applied Issues in Ecology: 16 Key concepts 1. Define ecology (an applied and a pure science) 2. To describe and understand, and on the basis of their understanding, to predict, manage, and control 3. Spatial and temporal scales(時空尺度). Patterns may be evident only at particular scales 4. By means of observations, field and laboratory experiments, and mathematical models. 5. Relies on truly scientific evidence (and the application of statistics) 17 1.1 introduction • Ecology (originally in German, Oekologie), was first defined in 1866 by Ernst Haeckel. – Comprehensive science of the relationship of the organism to the environment. – The science which concerns itself with the external relations of plants and animals to each other and to the past and present conditions of their existence, to be contrasted with physiology (internal relations) and morphology (structure). (Burdon-Sanderson, 1893) 18 「生態學」的定位(起源) 1866-1891 Haeckel, Ernst: 第一個使用 ecology (1866)。 – Ecology as a part of physiology (生理學)。 – total relationships of the animal both to its inorganic and its organic environment. – complex interrelations referred to by Darwin as the conditions for the struggle for existence. • eco-logy (the study of household) – 字源取自 eco-nomics (household + management) – the economy of nature (自然的經濟) 19 「生態學」的定位( by 1920s) • 1893 J. S. Burdon-Sanderson: President, British Ass. for Advancement of Science. – 將 Ecology 成為「生物學」中三大部之一。 – ecology, physiology and morphology – 生態學 => philosophy of living nature. • 1900s Clements & Cowles → plant ecology – 生態學 = 生理學 • 1910s Adam (1917) & Elton (1927) → animal ecologists – 生態學= new natural history (Adam, 1917) = scientific natural history (Elton, 1927) ┤ 20 「生態學」的定位( by 1950s) • 1949 Allee et al. principles of animal ecology – lack of knowledge of ecology among scholars and philosophers • 1. 由於內在的多元性 → 廣泛 distortion of 內容 • 2. 缺乏 historical studies 21 「生態學」的定位( 1960-80s) • 生態學 = – 生物的分佈和豐富量之研究 (Andrewartha, 1961) – 自然的結構和功能之研究 (Odum, 1963) – 「生物與其生活環境」的互動關係之研究 (Leo Smith, 1986) – 自然科學和社會科學的橋樑 (Odum, 1983) 22 「生態學」的定位( 1990s) • 生態學 = – 五部份:演化生態+行為生態+族群生態+群落 生態+應用生態 (Stiling, 1992) – 主要兩個根源:植物地理學 和 自然歷史學 (Leo Smith, 1992) – the Science of the universe(宇宙) (Hutchinson) (Colinvaux, 1993) • • • • 1. 体系的龐雜和歧異之維持和來源之瞭解 2. 生物一起生活的原則 3. 生命如何受天擇影響 4. 自然的經濟學 23 「生態學」的定位( by 2000) • 生態學 = – 生命與維生環境之關係 (Odum, 1993) – 決定「生物的分佈和豐富量」之互動的研究: Where, How many, Why (Krebs, 1994) – 「生物與其生活環境」的互動關係之研究 (Brewer, R., 1994) – 生物與生活環境間的互動關係。 (Stiling, P. D.,1996 – 自然的經濟 (Ricklefs, R. E.,1997; Smith and Smith, 1998) – 研習自然世界的次序 (order) (Ricklefs and Miller,2000) 24 Charles Elton (1900-1991), Animal Ecology (1927)→ Essentials of Ecology (3rd. Ed.) • The scientific study of the distribution and abundance of organisms and the interactions that determine distribution and abundance. (Townsend, Begon and Happer, 2008) • The primitive humans must have been ecologists of sorts – driven by the need to understand where and when their food and their enemies were to be found. (applied ecologists). 25 Fundamental problems • To what extent does competitions for food determine which species can coexist in a habitat? • What role does disease play in the dynamics of populations? • Why are there more species in the tropics than at the poles? • What is the relationship between soil productivity and plant community structure? • Why are some species more vulnerable to extinction than others? 26 Essentials of Ecology (3rd. Ed.) • The beauty of ecology is that it challenges us to develop an understanding of very basic and apparent problems – in a way that recognizes the uniqueness and complexity of all aspects of nature – but seeks patterns and predictions within this complexity rather than being swamped by it. 27 Essentials of Ecology (3rd. Ed.) • Two different classes of explanation in biology – Proximate explanation – Ultimate explanation • 學習研究的階段 – 描述、機制(功能)、模式、預測 (參閱實習手冊) • This book is about: 1. How ecological understand is achieved. 2. What we do understand. 3. How that understanding can help us predict, manage and control. 28 「生態學」的定位 生態學 = Eco + logy = 生活科學 = 自然的經濟 (Economy of Nature) 經濟學(Economics) =Eco + nomics = 生活的經營管理 • 生態學 (Ecology) – 生活科學:研究生物生活的學 理 (生存與發展之道) • 生物:個體、族群、群落,以及 生態體系 • 生活:存活 + 繁殖 (生生不息) 29 經濟學 vs.生態學 Odum, E. P. and G. W. Barrett (2005) Fundamentals of Ecology. 30 1.2 Scales, diversity and rigor (尺度、多樣性、精確) Three points 1. Ecological phenomena occur at a variety of scales 2. Ecological evidence comes from a variety of different sources 3. Ecology relies on truly scientific evidence and the application of statistics. 31 Chap. 1 Ecology and how to do it • 1.1 introduction • 1.2 Scales, diversity and rigor – 1.2.1 Questions of scale – 1.2.2 the diversity of ecological evidence – 1.2.3 statistics and scientific rigor • 1.3 Ecology in practice – 1.3.1 Brown trout in New Zealand – 1.3.2 Successions on old fields – 1.3.3 Hubbard Brook: a long-term – 1.3.4 a modeling study: vultures extinction 32 組成的位階 分類的位階 Biosphere Ecosystem Clade Community 複雜 Species Population Deme Organism Organism Organ System 1.2.1 Questions of scale Organ Tissue 圖a-1. 生物的組成與 分類的位階。clade 包 含:種以上的各個分 類位階。原資料摘取 自Kohn (1989), p.1097。 Cell Organelle Macromolecule-gene Molecule 簡單 33 8 6 地球:生物界 生物學 台灣生態 4 空 1km 2 間 的 生物体 1m 0 小型生物 量 度 大型生物 -2 無脊椎動物 1mm 原生物 -4 胞器 1um -6 細胞 病毒 蛋白質 -8 小分子 1A -10 -16 -14 -12 -10 1 微秒 -8 -6 1分 -4 -2 1天 0 2 1年 4 6 8 10 1 百萬年 時 間 的 量 度 圖a-2. 生物學的時間與空間的範圍。中間方形區是指「生物個体」 的生物學。時間量度以年為單位;空間則以公尺為單位。基本資料 34 取自Kohn (1989), p.1096。 1.2.2 the diversity of ecological evidence 1. Observations and field experiments 2. Laboratory experiments – Simple laboratory systems – Mathematical models 35 1.2.3 statistics and scientific rigor 1. Ecology: a search for conclusions in which we can be confident 2. Ecologists must think ahead 3. Ecology relies on representative samples 36 Species 分 陸域 類 位 海域 階 Phylum 0% 100% 百分率 (%) 圖a-3. 於各個分類位階,陸域與海域生態体系的龐雜 度之比較。資料取自Ray (1985) Amer. Zool., 24:452468。 37 圖a-4. 一般生命的延續 過程。「生命」起自 「基因型」(genotype, Phenotype space 受精卵, G1),經過t1時 t2 間,發展成「個体」(表 P1 現型,phenotype,P1); 再經過t2時間發展成為 t1 「成熟的個體」(P2), 經過t3時間產生「配子」 (精子或卵子,G2);再 Genotype space 經過t4時間結合成「受 精卵」(G‘1),另一個新 生命的開始,進入另一 G1 個循環。 P‘2 P2 t‘2 P‘1 t3 t‘1 t’3 t“1 G‘2 t‘4 G2 t4 G‘1 G“1 38 整合的生物學 --生態体系生物學 --群落(社區)生物學 真菌界 原核類 植物界 病毒類 動物界 原生物 族群生物學 個体生物學 生理與解剖學 組織生物學 細胞生物學 分子生物學 系統分類學 圖a-5. 生物學的結構。 親緣遺傳學 生 物 學 的 根 基 演化生物學 39 • Fig. 1-4 Compared with the strong setpoint controls at the organism level and below, organization and function at the population level and above. 40 Fig. 1-2. Ecological levels-of-organization spectrum emphasizing the interaction of living and nonliving components. 41 1.3 Ecology in practice • 1.3.1 Brown trout in New Zealand: effects on individuals, populations, communities and ecosystems • 1.3.2 Successions on old fields in Minnesota: a study in time and space • 1.3.3 Hubbard Brook: a long-term commitment of large-scale significance 42 1.3.1 Brown trout in New Zealand • (a) a brown trout (外來種):視覺覓食 • (b) a Galaxias fish (本土種):動態感應覓食 • 食物:Mayfly nymphs (浮游蛹) 43 白天量少 白天量少 • Fig. 1.7 (a) Mean number (+SE) of mayfly nymphs collected either from a trout stream or a Galaxias stream. (b) mean number (+SE) of mayfly nymphs observed on the upper surfaces of cobbles during late afternoon . 44 45 藻類量多 無脊椎動 物量少 • Fig. 1.8 (a) total invertebrate biomass • (b) algal biomass (chlorophyll a) • T, trout present; G, Galaxias present; N, no fish. 46 • Fig. 1.9 47 1.3.1 Brown trout in New Zealand • Individual level: trout → Mayfly 白天活動量 少 • Population level: trout 影響 Galaxias fish的 族群分布 • Communities (ecosystem) level: • Brown trout ← invertebrates ← algae – Energy flow – Cause a cascade of effects 48 1.3.2 Successions on old fields: a study in time and space • Ecological succession • (a) invader species decreased • (b) native prairie species increased • (c) annual species decreased • (d) perennial species increased • (e) soil nitrogen increased 49 Nitrogen effects • Do patches receiving different supply rates of nitrogen become less similar in species composition over time? • Do patches receiving similar supply rates of nitrogen become more similar in species composition over time? 50 • Fig. 1.11 (a) between 1982 and 1992, plots receiving 17g of nitrogen/m2/yr in the two fields became increasingly similar in composition. • (b) with only 1 g of nitrogen/m2/yr. 51 1.3.3 Hubbard Brook: a long-term commitment • Fig. 1.12 the Hubbard Brook experimental forest. • The value of large-scale studies and longterm data records 52 • The output of chemicals was greater than their input from rain, sleet and snow, except of nitrogen. 53 • Fig. 1.13 deforested 和 control 比較 54 • Fig. 1.14 Longterm changes in concentrations • Clean Air Act in 1970, emissions of SO2 were reduced. • 但是Ca+也下降。 • Acid rain may be responsible • Forest growth decline. 55 1.3.4 A modeling study: vultures extinction • In 1997, vultures in India and Pakistan began dropping from their perches. • Population surveys from 2000 to 2003 confirmed alarming rates of decline. • =0.50-0.52 Oriental white-backed vulture Gyps bengalensis 56 • Vultures 族群下降,野外腐肉增多,蒼蠅、野狗、 鼠類等增多,傳染病增多。 • 信仰,鳥葬 (a special tower, dakhma) • It took a few years to find, vultures 死因是累積尿 酸在體內,最後腎臟衰竭。 • 這是因為吃食到 diclofenac 藥物。 • 抗發炎的非類固醇藥物,1970年代使用於人體,後 來於Pakistan和印度廣泛使用於牲畜。 • 但這是唯一的因素嗎? 57 Fig. 1.14 58 59 6 about models • A model is a formulation that mimics a real-world phenomenon. • Computer-simulated models permit one to predict probable outcomes as parameters in the model are changed. • Symbols: – – – – E= energy source (forcing function) P= state variables F= flow pathways I=interaction functions 60 F= flow pathways L= feedback loop Fig. 1-6 Compartment diagram showing the five basic components of primary interest in modeling ecological systems. 61 Fig. 1-6 Compartment model with a feedback or control loop that transforms a linear system into a partially cyclical one. 62 隔離 Fig. 1-7 Interaction of positive and negative feedbacks in the relationships of atmospheric CO2, climate warming, soil respiration, and carbon sequestration. 63 Fig 1-8. The H. T. Odum energy language symbols used in model diagrams. 64 Fig. 1-9 Ecosystem model using energy language symbols and including estimated rates of energy flow for a Florida pine forest. 65 Boxes • 1.1 historical landmarks • 1.2 quantitative aspects: interpreting probabilities • 1.3 quantitative aspects: attaching confidence to results • 1.4 quantitative aspects: estimation: sampling, accuracy and precision • 1.5 Topical ECOncerns: invasions and homogenization of the biota: does it matter? 66 Questions 1. Search the library for a variety of definitions of ecology: which do you think is most appropriate and why? 2. Discuss the pros and cons of descriptive studies as opposed to laboratory studies of the same ecological phenomenon. 67 問題與討論 [email protected] Ayo 台南站 http://mail.nutn.edu.tw/~hycheng/ 國立臺南大學 環境與生態學院 Ayo 院長的個人網站