Ecology and Evolution of Dung Beetles

Leigh Simmons was born and educated in the UK, and is currantly Professor of Evolutionary Biology at the University of Western Australia. His research interests lie in all aspects of reproductive biology, from the evolution of sperm form and function, to mate searching and courtship behaviour. James Ridsdill-Smith was born and educated in the UK but carried out all his research in Australia working for CSIRO Entomology. He has been developing biological and ecological solutions to various pest problems and 15 years involved in the biological control of dung with scarabaeine dung beetles.

• 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 141.4.1 Niche expansion 151.4.2 Regional distribution and seasonal activity 171.4.3 Community dynamics 181.5 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 222.2.1 Dichotomiini and Coprini 242.2.2 Canthonini 252.2.3 Eucraniini 252.2.4 Phanaeini 252.2.5 Phanaeini + Eucraniini 262.2.6 Scarabaeini 262.2.7 Gymnopleurini 262.2.8 Eurysternini 262.2.9 Sisyphini 262.2.10 Onitini 272.2.11 Oniticellini 272.2.12 Onthophagini 272.3 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 78Box 4.1 Indirect genetic benefits of cryptic female choice in Onthophagus taurus 814.8 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 915.3.1 Morphology of pheromone-producing and -dispersing structures 935.3.2 Pheromone-dispersing behaviour 945.4 Chemical composition of Kheper pheromones 955.4.1 Electroantennographic detection 985.4.2 Comparison of the responses of beetle species to attractant compounds 985.4.3 The pheromone-disseminating carrier material 1025.5 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 1096.2.1 Does the development of a horn dimorphism in male dung beetles occur in a manner consistent with the assumptions of the ET model? 1106.3 Applying the threshold model 1186.3.1 Predicting the mean switchpoint of a population 1186.3.2 Estimating the selection on thresholds using the ET model 1196.3.3 Estimating selection under positive allometry 1206.4 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 128Box 7.1 Developmental genetic tools available in Onthophagus beetles: utility and limitations 1297.4 The origin and diversification of novel traits 1327.4.1 Dung beetle horns as novel traits 1337.4.2 How horns develop 1347.4.3 The developmental genetics of horn growth 1357.4.4 The developmental genetics of pupal remodelling 1377.4.5 The origin of adult thoracic horns through exaptation 1387.5 The regulation and evolution of scaling 1407.5.1 Onthophagine scaling relationships: the roles of nutrition and hormones 1427.5.2 Onthophagine scaling relationships: the role of trade-offs during development and evolution 1437.5.3 Onthophagine scaling relationships: developmental decoupling versus common developmental programme 1447.5.4 Onthophagine scaling relationships: the developmental genetics of size and shape 1477.6 The development, evolution, and consequences of phenotypic plasticity 1487.6.1 Developmental mechanisms and the evolutionary consequences of plasticity 1497.7 Conclusion 1518 The evolution of parental care in the onthophagine dung beetles 152John Hunt and Clarissa House8.1 Introduction 1528.2 Parental care theory 1548.2.1 A conventional view of parental care theory 1548.2.2 More recent developments in parental care theory 1568.3 Testing parental care theory using onthophagine dung beetles 1578.3.1 Parental care in onthophagine dung beetles 1588.3.2 The costs and benefits of parental care in onthophagine dung beetles 1608.3.3 Behavioural dynamics of the sexes during biparental care 1638.3.4 Confidence of paternity and paternal care 1668.3.5 Do parents optimize the care they provide? 1698.3.6 Evolutionary quantitative genetics of parental care 1738.4 Conclusions and future directions 1749 The visual ecology of dung beetles 177Marcus Byrne and Marie Dacke9.1 Introduction 1779.2 Insect eye structure 1799.2.1 The apposition eye 1799.2.2 The superposition eye 1799.3 Eye limitations 1819.4 Dung beetle vision 1829.4.1 Dim light vision 1829.4.2 The tapetum and enlarged rhabdoms 1859.4.3 The canthus 1869.5 Visual ecology of flight activity 1879.5.1 Diel flight activity 1879.5.2 Crepuscular flight activity 1889.5.3 Endothermy and vision 1889.5.4 Body size and flight activity 1899.6 Sexual selection and eyes 1909.7 Ball-rolling 1929.7.1 Orientation by ball-rolling beetles 1929.7.2 The polarization compass 1949.7.3 Polarization vision 1949.7.4 Polarization vision in dim light 1969.8 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 22111.2.1 The Finnish cow pat 22211.2.2 The Malagasy lemur pellet 22311.3 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 24612.2.1 Dung burial and nutrient cycling 24612.2.2 Control of dung-breeding flies 24712.2.3 Control of parasites 25012.3 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 25712.6.1 Summer rainfall climate area of Queensland 25812.6.2 Mediterranean climate area of south Western Australia 26012.6.3 Long-term population trends 26112.7 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 27013.3.1 Dung beetles as a viable candidate for biodiversity research 27113.3.2 Dung beetles as reliable indicators of environmental change 27213.3.3 Interpreting disturbance response patterns: application of a trait-based framework for ecological research 27613.3.4 Dung beetles as ecological disturbance indicator taxa: applied examples 28613.4 Dung beetle conservation 28613.5 Some ways forward 290References 293Subject index 340Taxonomic index 343

“The result is a complete book, highlighting however that there is still much to learn from these creatures.”  (Ethology Ecology & Evolution, 1 May 2013)“This book clearly demonstrates the value of dung beetles in a multitude of research disciplines and deserves a place on the bookshelf any scientist or natural historian.”  (J Insect Conserv, 9 February 2012)“Whilst this is already an interesting read, my overwhelming feeling is that there is an opportunity to greatly extend work on the ecology of dung beetles, with a real hope that they will provide genuine insight into common ecological problems. Perhaps this feeling is just what the authors hoped would be produced by their work.”  (British Ecological Society Bulletin, 1 March 2012)"I have however no doubt that there will be a second edition in five or so years time. And if any book deserves a second edition (with more in-situ photographs), it truly is this one." (The Amateur Naturalist magazine, 2011)