Martha Stone Wiske is lecturer at the Harvard Graduate School of Education where she co-directed the Educational Technology Center.  Her research is concerned with the integration of new technologies and the incorporation of learner-centered teaching for understanding. She is coeditor of Teaching for Understanding: Linking Research with Practice. Kristi Rennebohm Franz is an award-winning Washington State teacher who is known for her innovative use of new technologies in the classroom. Her classroom teaching has been filmed and featured in the PBS documentary Digital Divide.Lisa Breit develops professional development programs to help K-12 teachers design and implement curriculum with new technologies, and consults with school leaders on how to cultivate leadership and provide institutional support as teachers and students gain proficiency.

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• Preface xiiiAcknowledgements xvContributing authors xvii1 Reproductive competition and its impact on the evolution and ecology of dung beetles 1Leigh W. Simmons and T. James Ridsdill-Smith1.1 Introduction 11.2 Competition for mates and the evolution of morphological diversity 21.3 Competition for resources and the evolution of breeding strategies 91.4 Ecological consequences of intraspecific and interspecific competition 141.5 Conservation 191.6 Concluding remarks 202 The evolutionary history and diversification of dung beetles 21T. Keith Philips2.1 Introduction 212.2 Scarabaeinae diversity and tribal classification issues 222.3 Scarabaeine dung beetle phylogenies 272.4 The sister clade to the Scarabaeinae 312.5 The origin of the dung beetles 332.6 The oldest lineages and their geographical origin 342.7 Evolution of activity period 362.8 Evolution of feeding habits 362.9 Evolution of derived alternative lifestyles 372.10 Evolution of nidification: dung manipulation strategies 402.11 Evolution of nidification: nesting behaviour and subsocial care 422.12 Conclusions 442.13 Future work/gaps in knowledge 453 Male contest competition and the evolution of weapons 47Robert Knell3.1 Introduction 473.2 Dung beetle horns as weapons 493.3 Functional morphology of horns 503.4 Horns as predictors of victory 533.5 Are beetle horns simply tools? 553.6 The evolution of horns: rollers vs. tunnellers 563.7 The evolution of horns: population density 593.8 The evolution of horns: sex ratio 633.9 Future work 644 Sexual selection after mating: the evolutionary consequences of sperm competition and cryptic female choice in onthophagines 66Leigh W. Simmons4.1 Introduction 664.2 Sperm competition theory 684.3 Evolution of ejaculate expenditure in the genus Onthophagus 714.4 Evolutionary consequences of variation in ejaculate expenditure 724.5 Theoretical models of female choice 754.6 Quantitative genetics of ejaculate traits 764.7 Empirical evidence for adaptive cryptic female choice in Onthophagus taurus 784.8 Conclusions and future directions 834.9 Dedication and acknowledgement 865 Olfactory ecology 87G.D. Tribe and B.V. Burger5.1 Introduction 875.2 Orientation to dung and other resources 875.3 Olfactory cues used in mate attraction and mate recognition 915.4 Chemical composition of Kheper pheromones 955.5 Kairomones 1035.6 Defensive secretions 1045.7 Conclusions and future directions 1056 Explaining phenotypic diversity: the conditional strategy and threshold trait expression 107Joseph Tomkins and Wade Hazel6.1 Introduction 1076.2 The environmental threshold model 1096.3 Applying the threshold model 1186.4 Future directions 1237 Evolution and development: Onthophagus beetles and the evolutionary development genetics of innovation, allometry and plasticity 126Armin Moczek7.1 Introduction 1267.2 Evo-devo and eco-devo – a brief introduction 1277.3 Onthophagus beetles as an emerging model system in evo-devo and eco-devo 1287.4 The origin and diversification of novel traits 1327.5 The regulation and evolution of scaling 1407.6 The development, evolution, and consequences of phenotypic plasticity 1487.7 Conclusion 1518 The evolution of parental care in the onthophagine dung beetles 152John Hunt and Clarissa House8.1 Introduction 1528.2 Parental care theory 1548.3 Testing parental care theory using onthophagine dung beetles 1578.4 Conclusions and future directions 1749 The visual ecology of dung beetles 177Marcus Byrne and Marie Dacke9.1 Introduction 1779.2 Insect eye structure 1799.3 Eye limitations 1819.4 Dung beetle vision 1829.5 Visual ecology of flight activity 1879.6 Sexual selection and eyes 1909.7 Ball-rolling 1929.8 Conclusions 19810 The ecological implications of physiological diversity in dung beetles 200Steven L. Chown and C. Jaco Klok10.1 Introduction 20010.2 Thermoregulation 20110.3 Thermal tolerance 20710.4 Water balance 20810.5 Gas exchange and metabolic rate 21510.6 Conclusion and prospectus 21811 Dung beetle populations: structure and consequences 220Tomas Roslin and Heidi Viljanen11.1 Introduction 22011.2 Study systems 22111.3 Range size 22411.4 Habitat and resource selection 22711.5 Dung beetle movement 23011.6 The genetic structure of dung beetle populations 23511.7 Consequences: spatial population structures and responses to habitat loss 23811.8 Perspectives 24312 Biological control: ecosystem functions provided by dung beetles 245T. James Ridsdill-Smith and Penny B. Edwards12.1 Introduction 24512.2 Functions of dung beetles in ecosystems 24612.3 Dung beetles in pasture habitats 25012.4 Seasonal occurrence and abundance of native dung beetles in Australia 25112.5 Distribution and seasonal occurrence of introduced dung beetles in Australia 25412.6 Long-term studies of establishment and abundance 25712.7 Competitive exclusion 26212.8 Optimizing the benefits of biological control 26413 Dung beetles as a candidate study taxon in applied biodiversity conservation research 267Elizabeth S. Nichols and Toby A. Gardner13.1 Introduction 26713.2 Satisfying data needs to inform conservation practice 26813.3 The role of dung beetles in applied biodiversity research in human-modified landscapes 27013.4 Dung beetle conservation 28613.5 Some ways forward 290References 293Subject index 340Taxonomic index 343