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Preface Ecology and biomechanics: A mechanical approach to the ecology of animals and plants. When we decided upon this title there were two elements we wanted to emphasize: first, that biomechanical approaches have a lot to offer to ecological questions and second, that the approaches included in this book are independent of the organism being studied. Indeed, the present collection of state-of-the-art papers beautifully highlights how biomechanics can provide novel insights into long standing ecological and evolutionary questions (e.g. chapters 4, 14). As illustrated, for example, in the chapter by Wendy Griffiths (chapter 5) on grazing in ruminants there is tremendous scope for applying engineering principles for understanding the foraging strategies used by animals. Although most of the examples included in the book emphasize distinct organism-environment relationships, it is foreseeable that in the longer-term these kinds of approaches will span larger temporal and spatial scales to achieve wider application across ecosystems. The chapter by Karen Christensen-Dalsgaard (chapter 13) nicely illustrates this, and highlights how microbial ecosystems can be understood from the mechanics, morphology and motile responses of the individual organisms. The range of topics covered clearly demonstrates that increasing numbers of workers have begun to combine biomechanics and ecology to generate novel insights into questions of an ecological nature. We hope that this book will highlight the important cross fertilization that can occur by combining approaches from two ¿ at first sight ¿ very disparate sub-disciplines within the general field of biology and will stimulate other researchers to follow these kinds of approaches. The subjects covered include research based on both plants (chapters 1-4) and animals (chapters 10-12) as well as the interaction between plants and animals (chapters 5-9). By highlighting both theoretical concepts and practical approaches we hope that this book will be an important reference to students and researchers alike. Most of the chapters included in this book were originally presented at a symposium entitled ¿Ecology and biomechanics¿ held at the annual meeting of the Society for Experimental Biology in Edinburgh, U.K. in April 2004. Generous support by the Biomechanics Group of the Society for Experimental Biology enabled us to invite many outstanding speakers, most of which decided to contribute to the present volume. Anthony Herrel is a postdoctoral researcher of the Fund for Scientific Research, Flanders -Belgium (FWO-Vl). Thomas Speck is Professor for Functional Morphology and Director of the Botanic Garden of the Albert-Ludwigs-Universität Freiburg (Germany). Nick Rowe is Chargé de Recherche (CNRS) at the Botanique and Bioinformatique Research Institute, Montpellier, France. Acknowledgements We would like to acknowledge the much appreciated help of Bieke Vanhooydonck and Katleen Huyghe with the final editing and formatting of the chapters presented in this book. Additionally, we would like to thank the following referees for critical comments on the chapters and the many helpful suggestions for improvement: P. Aerts (U. Antwerp, Belgium), B. Borrell (U.C. Berkeley, U.S.A.), T. Buckley (Australian National University, Canberra, Australia), A. Davy (U. East Anglia, Norwich, U.K.), R. Dudley (U.C. Berkeley, U.S.A.), M. Edmunds (U. Central Lancashire, Preston, U.K.), S. Eigenbrode (U. Idaho, Moscow, U.S.A.), T. Fenchel (U. Copenhagen, Denmark), F. Gallenmüller (U. Freiburg, Germany), A. Goodman (U. Lincoln, U.K.), D. Irschick (Tulane University, New Orleans, U.S.A.), G. Jeronimidis (U. Reading, U.K.), M. Jervis (Cardiff University, U.K.), R. Jetter (U. British Columbia, Vancouver, Canada), J. Losos (Washington University, St. Louis, USA), K. Lunau (U. Duesseldorf, Germany), B. Moon (U. Louisiana, Lafayette, USA), C. Neinhuis (T.U. Dresden, Germany), J. Pilarski (Northern Arizona University, Flagstaff, USA), B. Vanhooydonck (U. Antwerp, Belgium), J. Vincent (U. Bath, U.K.). Finally we would like to thank David Fausel and John Sulzycki at CRC for all their help with the practical aspects of putting together the book. Preface Ecology and biomechanics: A mechanical approach to the ecology of animals and plants. When we decided upon this title there were two elements we wanted to emphasize: first, that biomechanical approaches have a lot to offer to ecological questions and second, that the approaches included in this book are independent of the organism being studied. Indeed, the present collection of state-of-the-art papers beautifully highlights how biomechanics can provide novel insights into long standing ecological and evolutionary questions (e.g. chapters 4, 14). As illustrated, for example, in the chapter by Wendy Griffiths (chapter 5) on grazing in ruminants there is tremendous scope for applying engineering principles for understanding the foraging strategies used by animals. Although most of the examples included in the book emphasize distinct organism-environment relationships, it is foreseeable that in the longer-term these kinds of approaches will span larger temporal and spatial scales to achieve wider application across ecosystems. The chapter by Karen Christensen-Dalsgaard (chapter 13) nicely illustrates this, and highlights how microbial ecosystems can be understood from the mechanics, morphology and motile responses of the individual organisms. The range of topics covered clearly demonstrates that increasing numbers of workers have begun to combine biomechanics and ecology to generate novel insights into questions of an ecological nature. Chapter 1: Tree Biomechanics and growth strategies in the context of forest functional ecology 1. INTRODUCTION 2. SOME BIOMECHANICAL CHARACTERISTICS OF TREES 2.1 WOOD AS LIGHTWEIGHT? CELLULAR AND FIBER REINFORCED MATERIAL 2.2 WOOD VARIABILITY 2.3 MECHANICS OF SECONDARY GROWTH 3. BIOMECHANICAL AND ECOLOGICAL SIGNIFICANCE OF HEIGHT 3.1 BIOMECHANICAL AND ENVIRONMENTAL CONSTRAINTS ON TREE HEIGHT AND THEIR ECOLOGICAL SIGNIFICANCE 3.1.1 Safety factor 3.1.2 Analysis of successive shapes occurring during growth due to the continuous increase of supported loads 3.2 BIOMECHANICAL FUNCTIONAL TRAITS DEFINED FROM RISK ASSESSMENT 3.2.1 Buckling or breakage of stems 3.2.2 Root anchorage 3.3 BIOMECHANICAL FUNCTIONAL TRAITS AND PROCESSES INVOLVED IN HEIGHT GROWTH STRATEGY 4. THE GROWTH PROCESSES THAT CONTROL THE MECHANICAL STABILITY OF SLENDER TREE STEMS 4.1 THE MECHANICAL CONTROL OF GROWTH 4.2 THE CONTROL OF STEM ORIENTATION TO MAINTAIN OR RESTORE THE TREE FORM, AND ALLOW VERTICAL GROWTH 4.3 THE CONTROL OF ROOT GROWTH TO SECURE ANCHORAGE 5. A PRACTICAL APPLICATION OF TREE BIOMECHANICS IN ECOLOGY 6. CONCLUSION 7. REFERENCES Chapter 2: Diversity of mechanical architectures in climbing plants: an ecological perspective 1. INTRODUCTION 1.1 IMPORTANCE OF CLIMBERS 1.2 MECHANICAL STRUCTURE AND DEVELOPMENT OF CLIMBERS 1.3 ATTACHMENT MODES OF CLIMBERS 1.4 MECHANICAL CONSTRAINTS AND TYPES OF ATTACHMENT 2. METHODS AND MATERIALS 2.1 EXPERIMENTAL PROTOCOLS 2.2 CALCULATION OF STRUCTURAL YOUNG¿S MODULUS 2.3 SAMPLING 3. RESULTS 3.1 MECHANICAL PROPERTIES AND TYPES OF ATTACHMENT 3.1.1 Twining climbers 3.1.2 Tendril climbers 3.1.3 Hook climbers 3.1.4 Branch angle climbers 3.1.5 Leaning climbers 4. DISCUSSION 4.1 MECHANICAL PROPERTIES AND ATTACHMENT OF DICOTYLEDONOUS CLIMBERS 4.2 CLIMBING GROWTH STRATEGIES IN MONOCOTYLEDONS AND PLANTS WITHOUT SECONDARY GROWTH 4.3 ECOLOGICAL DIVERSITY OF CLIMBERS AMONG DIFFERENT GROUPS 5. CONCLUSIONS 6. REFERENCES Chapter 3: The role of blade buoyancy and reconfiguration in the mechanical adaptation of the southern bullkelp Durvillaea 1. INTRODUCTION 1.1 THE INTERTIDAL ZONE 1.2 THE SOUTHERN BULLKELPS DURVILLAE ANTARCTICA AND D. WILLANA 1.3 DRAG AND STREAMLINING 1.3.1 Vogel number 1.4 OBJECTIVES 2. MATERIAL AND METHODS 2.1 TESTED SEAWEEDS 2.2 DRAG FORCES 2.3 SHORTENING EXPERIMENTS 2.4 DRAG COEFFICIENTS AND RECONFIGURATION 2.5 BUOYANCY 2.6 FIELD STUDIES 2.7 MORPHOLOGICAL SURVEY 2.8 STATISTICAL ANALYSIS 3. RESULTS 3.1 DRAG FORCES 3.2 SHORTENING EXPERIMENTS 3.3 DRAG COEFFICIENTS AND RECONFIGURATION 3.4 VOGEL NUMBER 3.5 BUOYANCY 3.6 FIELD STUDIES 3.7 MORPHOLOGICAL SURVEY 4. DISCUSSION 4.1 DRAG FORCES 4.2 DRAG COEFFICIENTS, RECONFIGURATION AND THE VOGEL NUMBER 4.3 BUOYANCY AND FIELD STUDIES 4.4 MORPHOLOGICAL SURVEY 5. CONCLUSION 6. ACKNOWLEDGEMENTS 7. REFERENCES Chapter 4: Murray¿s law and the vascular architecture of plants 1. INTRODUCTION 2. MURRAY¿S LAW 3. APPLYING MURRAY¿S LAW TO XYLEM 4. IMPORTANCE OF THE CONDUIT FURCATION NUMBER (F) 5. DOES XYLEM FOLLOW MURRAY¿S LAW? 6. DOES TREE WOOD NOT FOLLOW MURRAY¿S LAW? 7. NATURE OF THE MECHANICAL CONSTRAINT ON HYDRAULIC EFFICIENCY 8. DA VINCI¿S RULE 9. DEVELOPMENTAL AND PHYSIOLOGICAL CONSTRAINTS ON TRANSPORT EFFICIENCY 10. COMPARATIVE EFFICIENCY OF CONIFER VS. ANGIOSPERM TREE WOOD 11. CONCLUSIONS 12. ACKNOWLEDGEMENTS 13. REFERENCES Chapter 5: Plant-animal mechanics and bite procurement in grazing ruminants 1. INTRODUCTION 2. RUMINANT SPECIES 3. HARVESTING APPARATUS 4. BITE PROCUREMENT 5. PLANT FORM AND FRACTURE MECHANICS AT THE PLANT LEVEL 6. INSTRUMENTATION FOR MEASURING PLANT FRACTURE MECHANICS UNDER TENSION 7. APPLICATION OF PLANT FRACTURE MECHANICS TO FORAGING STRATEGIES 8. INSTRUMENTATION FOR MEASURING BITE FORCE AT THE ANIMAL LEVEL 8.1 PREDICTION OF BITE FORCE ASSESSMENT OF PLANT FRACTURE PROPERTIES 8.2 BIOMECHANICAL FORCE INSTRUMENTS 9. BITING EFFORT 10. CONCLUSION 11. ACKNOWLEDGMENTS 12. REFERENCES Chapter 6: Biomechanics of Salvia flowers: the role of lever and flower tube in specialization on pollinators 1. INTRODCUTION 1.1 BIOMECHANICS AND BEE POLLINATION 1.2 A CASE STUDY: THE STANDARD LEVER MECHANISM IN SALVIA 2. MATERIALS AND METHODS 2.1 MATERIALS 2.2 FORCES OF FLOWER-VISITING BEES 2.3 FORCE MEASUREMENTS ON SALVIA FLOWERS AND STAMINAL LEVERS 3. RESULTS 3.1 FORCES EXERTED BY B. TERRESTRIS AND A. MELLIFERA 3.2 FORCES AND FLOWER VISITORS OF SALVIA 4. DISCUSSION 4.1 INSECT FORCES 4.2 OBSERVED FLOWER VISITORS 4.3 FORCES MEASURED IN SALVIA FLOWERS 4.3.1 Critical discussion of the applied methods 4.3.2 Comparison of levers and internal barriers in flowers 4.4 COMPARING INSECT FORCES TO THE BARRIERS IN FLOWERS 4.5 PROBOSCIS LENGTH, FLOWER-TUBE LENGTH AND FORCES EXERTED BY VISITING BEES 5. CONCLUSION 6. ACKNOWLEDGEMENTS 7. REFERENCES Chapter 7: Do plant waxes make insect attachment structures dirty? Experimental evidences for the contamination-hypothesis 1. INTRODUCTION 2. MATERIAL AND METHODS 2.1 PLANT SURFACES AND OTHER SUBSTRATES 2.2 MODEL INSECT SPECIES AND EXPERIMENTS 3. RESULTS 3.1 PRUINOSE PLANT SURFACES 3.2 ADHESIVE PADS OF THE BEETLE CHRYSOLINA FASTUOSA 3.3 PAD CONTAMINATION 4. DISCUSSION 4.1 CONTAMINATING EFFECT OF CRYSTALLINE EPICUTICULAR WAXES ON INSECT ATTACHMENT DEVICES 4.2 DEPENDENCE OF PAD CONTAMINATION ON THE WAX MICROMORPHOLOGY 5. ACKNOWLEDGEMENTS 6. REFERENCES Chapter 8: Ecology and biomechanics of slippery wax barriers and waxrunning to Macaranga-ant mutualisms 1. INTRODUCTION 2. ECOLOGY AND EVOLUTION OF WAX BARRIERS IN THE ANT-PLANT GENUS MACARANGA 2.1 PROTECTION OF SPECIFIC ANT-PARTNERS AGAINST GENERALIST ANTS 2.2 EFFECT OF WAX BARRIERS ON HOST SPECIFICITY 2.3 EVOLUTION OF MACARANGA WAX BARRIERS 2.4 ADAPTIVE SYNDROMES OF ANT-ASSOCIATIONS IN WAXY AND NON-WAXY MACARANGA ANT-PLANTS 3. BIOMECHANICS OF WAXRUNNING IN CREMATOGASTER (DECACREMA) ANTS 3.1 TARSAL ATTACHMENT DEVICES IN ANTS 3.2 MECHANISMS OF SLIPPERINESS 3.3 MECHANISMS OF WAXRUNNING 3.3.1 Attachment force vs. climbing performance: is waxrunning capacity based on greater attachment or superior locomotion? 3.3.2 Comparative morphometry of waxrunners and non-waxrunners 3.3.3 Mechanical benefit of long legs for climbing ants 3.3.4 Kinematics of climbing in Crematogaster (Decacrema) waxrunners and non-waxrunners 4. CONCLUSIONS 5. ACKNOWLEDGEMENTS 6. REFERENCES Chapter 9: Nectar feeding in long-proboscid insects 1. INTRODUCTION 2. FUNCTIONAL DIVERSITY OF LONG MOUTHPARTS 2.1 EVOLUTION OF SUCTION FEEDING 2.2 ANATOMICAL CONSIDERATIONS 2.2.1 Proboscis sealing mechanisms 2.2.2 Tip region 2.2.3 Fluid pumps 3. FEEDING MECHANICS AND FORAGING ECOLOGY 3.1 PROBOSCIS MOBILITY AND FLORAL HANDLING 3.2 FACTORS INFLUENCING FLUID HANDLING 3.3 ENVIRONMENTAL INFLUENCES ON FLORAL NECTAR CONSTITUENTS 3.4 HAVE NECTAR SUGAR CONCENTRATIONS EVOLVED TO MATCH POLLINATOR PREFERENCES? 3.5 TEMPERATURE AND OPTIMAL NECTAR FORAGING 4. CONCLUDING REMARKS 5. ACKNOWLEDGEMENTS 6. REFERENCES Chapter 10: Biomechanics and behavioural mimicry in insects 1. INTRODUCTION 1.1 BATESIAN MIMICRY 1.2 MÜLLERIAN MIMICRY 2. MORPHOLOGICAL MIMICRY 3. BEHAVIOURAL MIMICRY 3.1 BEHAVIOURAL MIMICRY IN INSECTS 3.2 MIMICRY IN TERRESTRIAL LOCOMOTION 3.3 FLIGHT MIMICRY 3.3.1 The mimetic flight behaviour of butterflies 3.3.2 The Mimetic flight behaviour of hoverflies 4. CONCLUSION 5. REFERENCES Chapter 11: Inter-individual variation in the muscle physiology of vertebrate ectotherms: consequences for behavioural and ecological performance. 1. INTRODUCTION 2. EVOLUTIONARY IMPLICATIONS OF INDIVIDUAL VARIATION IN BEHAVIOURAL PERFORMANCE AND MUSCLE PHYSIOLOGY 3. SCALING EFFECTS ON VERTEBRATE ECTOTHERM MUSCLE AND WHOLE BODY PERFORMANCE 4. RELATIONSHIPS BETWEEN MUSCLE SPECIALIZATION AND THE INDIVIDUAL BEHAVIOURAL PERFORMANCE OF VERTEBRATE ECTOTHERMS 5. TRADE-OFFS IN WHOLE-MUSCLE FUNCTION AND ITS ECOLOGICAL IMPORTANCE 6. CONCLUSION 7. REFERENCES Chapter 12: Power generation during locomotion in Anolis lizards: an ecomorphological approach. 1. INTRODUCTION 2. MATERIAL AND METHODS 2.1 ANIMALS 2.2 MORPHOLOGY 2.3 RUNNING TRIALS 2.4 JUMPING TRIALS 2.5 CONFIGURATION OF HIND LIMBS 2.6 ECOLOGY 2.7 STATISTICAL ANALYSIS 3. RESULTS 4. DISCUSSION 4.1 ECOLOGICAL CORRELATES 4.2 INTERSPECIFIC VARIATION 4.3 POWER OUTPUT DURING RUNNING AND JUMPING: A TWO-SPECIES COMPARISON 5. ACKNOWLEDGEMENTS 6. REFERENCES Chapter 13: Implications of microbial motility on the water column ecosystems. 1. INTRODUCTION 1.1 MICROBIAL ECOLOGY IN A LARGER CONTEXT 2. GENERATING MOTION WITH CILIA OR FLAGELLA 2.1 SMOOTH FLAGELLA 2.2 HISPID FLAGELLA 2.3 CILIA 3. ENERGETICS OF MOTION 4. FEEDING MECHANISM 4.1 THE COEXISTENCE OF FILTER FEEDERS 4.2 ATTACHING TO PARTICLES WHILE FEEDING 5. ORIENTATION TO STIMULI 6. ACKNOWLEDGEMENTS 7. REFERENCES Chapter 14: The Biomechanics of Ecological speciation. 1. INTRODUCTION 2. MODES OF SPECIATION 3. BIOMECHANICS AND ECOLOGICAL SPECIATION 3.1 MATING DISPLAYS AND BODY SIZE 3.2 MATING DISPLAYS AND LOCOMOTION 3.3 MATING DISPLAYS AND FEEDING 4. ECOLOGICAL (IN)DEPENDENCE AND THE EVOLUTION OF ISOLATING BARRIERS 5. POSITIVE FEEDBACK LOOPS 6. DUAL FITNESS CONSEQUENCES FOR ECOLOGICAL SPECIATION 7. PERFORMANCE AND MATING DISPLAY PRODUCTION 8. CONCLUSION 9. REFERENCES
Library of Congress Subject Headings for this publication:
Ecology.
Biomechanics.