Life History Evolution E-Book
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- Herausgeber: John Wiley & Sons
- Kategorie: Wissenschaft und neue Technologien
- Sprache: Englisch
Provides a timely and authoritative account of Life History Evolution by a multidisciplinary team of scholars and researchers from around the world
Life History Evolution: Traits, Interactions, and Applications presents a cutting-edge synthesis of the mechanisms driving life history strategies that span the breadth of taxa, from bacteria to humans. Integrating classical and contemporary perspectives, this comprehensive volume addresses how organisms evolve traits in response to diverse ecological pressures. Editors Michal Segoli and Eric Wajnberg bring together leading experts to explore the intersection of evolutionary biology, ecology, and applied research, focusing on the evolving complexity of life history traits and their implications.
In-depth yet accessible chapters cover a broad spectrum of life history traits, from classical traits of lifespan and reproduction to more complex interactions like social behaviour, predator-prey dynamics, and human-induced evolutionary processes. The contributing authors explain essential concepts, identify critical knowledge gaps, discuss future research directions, and demonstrate the relevance of life history evolution in addressing climate change, species invasion, pollution, and more.
Providing a well-balanced understanding of life history traits and their implications, Life History Evolution:
- Incorporates recent advances in evolutionary theory, including eco-evolutionary feedback loops and anthropogenic impacts
- Offers diverse perspectives and original research from leading experts in fields such as evolutionary biology, ecology, entomology, zoology, agriculture, and veterinary medicine
- Discusses life history evolution in the context of co-evolved interactions such as predator-prey, parasite-host, plant-herbivore, and endosymbiont-host relationships
- Provides an overview of the foundational theory, recent developments, and current thinking in the field
- Features numerous case studies that highlight real-world applications in biological control, wildlife management, climate change adaptation, and others
Revealing how life history traits shape the evolutionary strategies of organisms, Life History Evolution: Traits, Interactions, and Applications is an essential resource for undergraduate and graduate students, researchers, industry professionals, and policymakers in ecological science. It is an ideal textbook for courses in evolutionary ecology, evolutionary biology, conservation biology, environmental science, and environmental management.
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Table of Contents
Cover
Table of Contents
Title Page
Copyright Page
List of Contributors
Foreword
References
Preface
Part I: Traits
1 Body Size and Timing of Maturation
1.1 Introduction
1.2 Part I
1.3 Part II
1.4 Conclusions
Acknowledgements
References
2 Evolution of Ageing and Lifespan
2.1 Introduction
2.2 Evolutionary Theory of Ageing
2.3 Asynchronous Ageing
2.4 Sex Differences in Ageing
2.5 Williams and Anti‐Williams: Age, Density and Condition‐Dependence of Mortality
2.6 Concluding Remarks
Acknowledgements
References
3 Offspring Size and Life History Theory: What Do We Know?
3.1 Offspring Size Defined
3.2 The Knowns
3.3 The Unknowns
References
4 The Evolution of Insect Egg Loads
4.1 Trade‐Offs Between Early and Late Components of Reproduction
4.2 Time
vs
. Egg Limitation in Insects
4.3 Egg Maturation Patterns
4.4 The Relative Importance of Egg and Time Limitation
4.5 Additional Life History Strategies to Overcome the Risk of Egg Limitation
4.6 Conclusions and Future Directions
Acknowledgements
References
5 Sex‐Specific Life Histories
5.1 Introduction
5.2 Various Unidirectional Effects: Unguarded X, Mother’s Curse and Toxic Y
5.3 Multi‐directionality: Coevolution of Different Traits
5.4 Towards Progress
Acknowledgements
References
6 Parental Care and Life History
6.1 What Is Parental Care and How Does It Relate to Life History?
6.2 Distinguishing Between the Origin and the Maintenance of Parental Care
6.3 Life History and the Origin of Parental Care
6.4 Life History and the Maintenance of Parental Care
6.5 Co‐evolution Between Parental Care, Offspring Traits and Parental Traits
6.6 Sexual Selection, Life History and Sex Differences in Parental Care
6.7 Stochasticity, Environmental Variability, Life History and Parental Care
6.8 Plasticity and the Evolution of Parental Care
6.9 Final Conclusions and Future Directions
Acknowledgements
References
7 Sex Allocation
7.1 Introduction
7.2 Fisher’s Theory
7.3 Interaction with Relatives
7.4 Environmental Condition
7.5 Future Directions
Acknowledgements
References
8 Life History Evolution
8.1 Integration of Metamorphic Development Within the Life Cycle
8.2 The Regulation of Metamorphic Development by Hormones
8.3 Review of Metamorphic Mechanisms Across Taxa with Ecological and Evolutionary Considerations
8.4 Anthropogenic Environmental Impacts and Global Climate Change
References
9 Social Living and Life History Evolution, with a Focus on Ageing and Longevity
9.1 Introduction
9.2 Ultimate Causes of Long Reproductive Lifespans
9.3 Colony Life History in Obligatory Eusocial Insects
9.4 Proximate Mechanisms
9.5 Conclusion
Acknowledgements
References
10 Integrating Dispersal in Life History
10.1 Introduction
10.2 Dispersal as Part of the Life History
10.3 The Theory of Dispersal and Life Histories
10.4 Dispersal‐Life History Co‐variation in Nature
10.5 Concluding Remarks and Outlook: Why Should We Care?
References
11 The Evolution of Human Life Histories
11.1 Introduction
11.2 Life History Trade‐Offs
11.3 The Life Histories of Great Apes
11.4 Variation in Human Life History
11.5 Menopause and the Post‐reproductive Lifespan
11.6 Final Remarks
References
Part II: Interactions
12 Life History Traits in the Context of Predator–Prey Interactions
12.1 Introduction
12.2 Types of Predation
12.3 Theory for Predator‐Driven Life History Evolution
12.4 Empirical Evidence
12.5 Adaptive Plasticity in Life Histories
12.6 Future Directions
Acknowledgements
References
13 Life History Trait Evolution in the Context of Host–Parasite Interactions
13.1 Introduction
13.2 Host Life History Evolution in Response to Parasites
13.3 Parasite Life History Evolution in Response to Hosts: The Case of Virulence
13.4 Concluding Remarks
References
14 How Do Microbial Symbionts Shape the Life Histories of Multicellular Organisms?
14.1 Introduction
14.2 Categories of Microbial Symbiosis
14.3 How Microbial Symbionts Are Involved in Essential Biological Functions of Their Hosts?
14.4 Nutritional Microbial Symbionts
14.5 Reproductive Microbial Symbionts
14.6 Defensive Microbial Endosymbionts
14.7 Diapause and Microbial Symbionts
14.8 Concluding Remarks
References
15 Ecological and Evolutionary Links Between Defences and Life History Traits in Plants
15.1 Evolutionary Ecology of Plant Defences Against Herbivores
15.2 Correlated Evolution of Plant Defences and Life History Traits
15.3 Tripartite Views Shed Insight into the Evolution of Plant Life History Traits
15.4 Challenges for Future Research
References
16 Are you in Synch?
16.1 Generalisation in Pollination Networks
16.2 What Drives Flowering Phenology?
16.3 What Drives Pollinator Phenology?
16.4 Do Interacting Plant–Insect Species Share Similar Reaction Norms to Temperature?
16.5 Species‐Level Phenological Asynchrony and Generalized Pollination: A Case Study
16.6 Community‐Level Phenology and Pollination Specialisation
16.7 Concluding Remarks
References
17 Life Histories in the Context of Mutualism
17.1 Introduction
17.2 Mutualism Benefits and Life History Traits
17.3 Future Directions
References
Part III: Applications
18 Life History and Climate Change
18.1 Introduction
18.2 Effects of ACC on Life History Strategies and Trade‐Offs
18.3 Phenology
18.4 Body Size
18.5 Reproductive Output and Success
18.6 Survival and Senescence
18.7 Population Demography and Extinction Risk
18.8 Genetic or Environmental Responses
18.9 Conclusions and Outlook
Acknowledgements
References
19 Environmental Pollution Effects on Life History
19.1 Introduction
19.2 The Role of Life History Theories in Ecotoxicology
19.3 The Acquisition/Allocation Principle and the Responses of Organisms to Pollution
19.4 Literature Survey on Mechanisms Involved in the Life History Responses to Pollutants
19.5 Case Studies
19.6 Conclusion and Future Directions
References
20 Life History Evolution on Expansion Fronts
20.1 What Are Expansion Fronts and Why Are They Hotspots for Rapid Evolution
20.2 Trade‐Offs Matter
20.3 Other Types of Expansions, How Our Expectations Might Change
20.4 An Applied Case Study: The Cane Toad
20.5 Summary and Future Directions
References
21 Adaptive Evolution of Life History Traits in Urban Environments
21.1 Introduction
21.2 Urban Drivers of Selection on Life History Traits
21.3 Studying Evolution in Urban Areas
21.4 Available Evidence of Adaptive Life History Evolution in Urban Areas
21.5 Synthesis and Perspectives
Acknowledgements
References
22 Life History and Biological Control
22.1 Introduction
22.2 Selecting Among Interspecific Life History Variation
22.3 Managing or Manipulating Intraspecific Life History Variation
22.4 Using Life History to Inform Environmental Management and Agent Release Strategies
22.5 Future Directions and Conclusions
Acknowledgements
References
23 Life History and Exploitative Management of Fish and Wildlife
23.1 Introduction
23.2 Life History Traits, Density‐Dependence and Sustainable Harvest
23.3 Contrasting Life Histories and Harvest Potential
23.4 Ecological Plasticity and Evolutionary Sources of Variability: A Few Ungulate Examples
23.5 How Can Knowledge of Life History Traits Improve Harvest Management?
23.6 Life History and Trophy Hunting
23.7 Life History and Compensatory Population Responses to Harvest
23.8 The Special Case of Sexually Selected Infanticide
23.9 Can Harvest Affect the Evolution of Life History Strategies?
23.10 Conclusion and Future Directions
Acknowledgements
References
24 Life History and the Control of Diseases
24.1 Introduction
24.2 Life History Outcomes: A Classic Theoretical Scaffold to Illustrate Predictions
24.3 Levels of Selection
24.4 The Complexities of Variance and Covariation in Empirical Systems
24.5 Frontiers in Life History Evolution and Pathogen Control
24.6 Conclusions
References
Index
End User License Agreement
List of Tables
Chapter 6
Table 6.1 General life history conditions that are most likely to lead to th...
Chapter 7
Table 7.1 The expected fitness consequence of a mother producing different o...
Table 7.2 The whole reproductive success of a population under different pop...
Chapter 10
Table 10.1 Definitions of the identified cost types of dispersal
Chapter 11
Table 11.1 Life history parameters of extant female great apes from wild pop...
Chapter 13
Table 13.1 Parallels between life history traits in hosts and parasites duri...
Table 13.2 Host life history trait modifications in response to parasitism....
Chapter 14
Table 14.1 Selected examples of nutritional symbioses between eukaryote host...
Table 14.2 Selected examples of defensive symbioses between eukaryote hosts ...
Chapter 15
Table 15.1 A general summary of reviewed mechanisms underlying associations ...
Chapter 19
Table 19.1 Predictions from the two main responses of an organism to a pollu...
Table 19.2 Effects of physical pollutants on life history traits in animal s...
Table 19.3 Effects of chemical pollutants (metals) on life history traits in...
Table 19.4 Effects of chemical pollutants (organic) on life history traits i...
Chapter 21
Table 21.1 Selected case studies of adaptive evolution of life history trait...
Chapter 23
Table 23.1 Key life history traits of native populations of mountain goats (
List of Illustrations
Chapter 1
Figure 1.1 Left panel: conceptual setting for optimality analysis of age and...
Figure 1.2 An ovipositing flightless female of the erebid moth
Orgyia antiqu
...
Figure 1.3 Reaction norms for age and size at maturation. (a) The dependence...
Figure 1.4 A schematic presentation of different shapes of bivariate reactio...
Figure 1.5 (a) The dependence of fecundity on adult body size (expressed as ...
Figure 1.6 A probabilistic reaction norm. Dots indicate particular hypotheti...
Figure 1.7 A schematic presentation of possible plastic and evolutionary cha...
Figure 1.8 Sensitivity of the measures of growth rate to mathematical detail...
Figure 1.9 The reconstructed evolution of body size for northern European re...
Chapter 2
Figure 2.1 The strength of age‐specific selection is maximized during early ...
Figure 2.2 Dillin et al. (2002) provide the most striking example of uncoupl...
Figure 2.3 Illustration of the DTA, or ‘early‐life inertia’, from Lemaitre e...
Figure 2.4 Age‐specific changes in twenty different traits obtained from a w...
Figure 2.5 Experimental evolution of sexual dimorphism and monomorphism in l...
Chapter 3
Figure 3.1 Schematic representation showing two hypothetical species that pr...
Figure 3.2
Bugula neritina
is a colonial marine invertebrate that broods its...
Chapter 4
Figure 4.1 A parasitoid female wasp of the genus
Acroclisoides
ovipositing i...
Figure 4.2 Illustration of egg limitation and time limitation in parasitoids...
Figure 4.3 Results from a model by Segoli and Wajnberg (2020), predicting th...
Figure 4.4 Relationship between relative egg size (egg length/body length) a...
Chapter 5
Figure 5.1 The black‐headed and the red‐headed morph of the Gouldian finch,
Figure 5.2 The results of the model of Gouldian finch morphs differing in ag...
Figure 5.3 A simple model predicting male (blue lines and symbols) and femal...
Chapter 6
Figure 6.1 The complex links between life history, parental care and environ...
Figure 6.2 Parental care is diverse in nature, and one or both sexes provide...
Figure 6.3 Schematic illustration of reaction norms for parental care based ...
Chapter 7
Figure 7.1 Offspring sex ratio (proportion of sons) depending on the number ...
Figure 7.2 Offspring sex ratio after the experimental evolution in the mite
Figure 7.3 Offspring sex ratios in the parasitoid wasp
Melittobia australica
Figure 7.4 Sex allocation (relative investment in male function) increases w...
Figure 7.5 Sex ratio adjustment in response to territory quality in the Seyc...
Figure 7.6 Conceptual model for reproductive success between males and femal...
Figure 7.7 (a) The lifetime reproductive success of sons (solid circle) incr...
Figure 7.8 (a) Observed sex ratio (circles) consistent with predicted sex ra...
Figure 7.9 Predicted sex‐determining systems (a, c) and consequent cohort se...
Chapter 8
Figure 8.1 Metamorphic development with drastic morphological changes and a ...
Figure 8.2 Hormones and peptides are major players in the regulation of meta...
Figure 8.3 Metamorphic life cycles feature important similarities and differ...
Chapter 9
Figure 9.1 Social evolution and the evolution of longevity. High survival fa...
Figure 9.2 Longevity data for termites. Shown are (a) median lifespans acros...
Figure 9.3 The TI‐J‐LiFe network with major results for eusocial animals. Th...
Chapter 10
Figure 10.1 Dispersal costs and the integration life historyThe life cyc...
Figure 10.2 Evolution of dispersal syndromes in relation to spatio‐temporal ...
Figure 10.3 The strength and direction of dispersal syndromes can change whe...
Chapter 11
Figure 11.1 The global distribution of the hunter‐gatherer groups that are r...
Figure 11.2 Phylogeny of 124 mammals showing the proportion of female years ...
Figure 11.3 Hypothetical pathways to the (a) evolution of menopause and (b) ...
Chapter 12
Figure 12.1 Trinidadian guppies and some of their downstream predators in th...
Chapter 13
Figure 13.1 Graphical illustrations of host and parasite life history change...
Figure 13.2 Aquatic model system, used for experimental evolution in the con...
Figure 13.3 Classic model of virulence evolution. Parasite fitness is define...
Chapter 14
Figure 14.1 A sea anemone,
Exaiptasia pallida
, inoculated with algal cells o...
Figure 14.2 A
Trifolium
sp. plant (left) and its root nodules (the whitish b...
Figure 14.3 (a) Transmission electron micrograph showing elongate cells of t...
Chapter 15
Figure 15.1 Trade‐offs between different measures of plant growth and defenc...
Figure 15.2 Secretion of resin (viscous fluid exuded from ducts) from a youn...
Figure 15.3 A honeybee collecting nectar and a caterpillar feeding on flower...
Chapter 16
Figure 16.1
Anemone coronaria
’s distribution in Israel, plotted on the backg...
Figure 16.2
A. coronaria
’s main colour morphs.
Figure 16.3 Proportions of
A. coronaria
blooming reports over the flowering ...
Figure 16.4 The composition of visitor taxa to
A. coronaria
flower arrays at...
Figure 16.5 The composition of visitor taxa to
A. coronaria
flower arrays at...
Chapter 17
Figure 17.1 Hypothetical curves of responses between life history traits and...
Figure 17.2 Examples of mutualisms. (a) Tiny fig wasps carrying pollen swarm...
Chapter 18
Figure 18.1 Examples of species subject to long‐term population studies enab...
Chapter 19
Figure 19.1 Graphical representation of the van Noordwijk and de Jong 1986 m...
Figure 19.2 Hypothetical consequences of a pollutant on resources allocated ...
Figure 19.3 (a) Boxplot of male frequency over time (
i.e
., three‐day transfe...
Chapter 20
Figure 20.1 Evolutionary dynamics on a pulled expansion front. The front is ...
Figure 20.2 The cane toad
Rhinella marina
, a species that has become a textb...
Figure 20.3 Unlike natural invasions, experimental invasions can be replicat...
Figure 20.4 A hypothetical
r, K and D
space showing a possible evolutionary ...
Figure 20.5 Evolutionary dynamics on a pushed front. The density profile of ...
Chapter 21
Figure 21.1 Urban drivers of life history evolution. Five main dimensions of...
Figure 21.2 Evolution of reduced dispersal in the Mediterranean weed
Crepis
...
Figure 21.3 Water fleas of the species
Daphnia magna
have a faster ‘pace of ...
Figure 21.4 Synthetic summary of urban dimensions driving evolution across l...
Chapter 22
Figure 22.1 Types of biological control agents, target species and approache...
Figure 22.2 Biological control agents are often collected from a natural env...
Figure 22.3 The parasitoid wasp
Encarsia formosa
(adult ovipositing into a w...
Figure 22.4 An adult (left) and larva (right) of the northern tamarisk beetl...
Figure 22.5 The parasitoid wasp
Anagrus daanei
is a biological control agent...
Figure 22.6 An example of how understanding the role of life history variati...
Chapter 23
Figure 23.1 The relationship between population size and growth rate (
r
) whe...
Figure 23.2 Mountain goats and white‐tailed deer are similar‐sized ungulates...
Chapter 24
Figure 24.1 Many pathogen life history traits (left box) are shaped by withi...
Figure 24.2 The susceptible‐infected‐recovered model with waning of immunity...
Figure 24.3 Evolution of virulence. The persistence of virulence or the tend...
Figure 24.4 Evolution of virulence: beyond
R
0
maximization. Where higher pat...
Figure 24.5 Immune escape. (a) The phylogeny of the influenza A H3N2 viruses...
Figure 24.6 Levels of selection (a) over the course of an infection, pathoge...
Figure 24.7 Host heterogeneity in pathogen responses. Distinction can be mad...
Figure 24.8 Superspreading and the dynamics of infection. A spectrum of path...
Guide
Cover Page
Table of Contents
Title Page
Copyright Page
List of Contributors
Foreword
Preface
Begin Reading
Index
WILEY END USER LICENSE AGREEMENT
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Life History Evolution
Traits, Interactions, and Applications
Edited by
Michal Segoli
Mitrani Department of Desert EcologyBlaustein Institutes for Desert ResearchBen-Gurion University of the NegevMidreshet Ben-GurionIsrael
Eric Wajnberg
INRAE, Sophia Antipolis, FranceINRIA, Project Hephaistos, Sophia Antipolis, FranceDepartamento de Entomologia e AcarologiaUniversidade de São PauloEscola Superior de Agricultura “Luiz de Queiroz”Piracicaba, São Paulo, Brazil
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Library of Congress Cataloging‐in‐Publication Data
Names: Segoli, Michal, editor. | Wajnberg, E., editor.Title: Life history evolution : traits, interactions, and applications / edited by Michal Segoli, Eric Wajnberg.Description: Hoboken, NJ : Wiley, 2025. | Includes index.Identifiers: LCCN 2024026594 (print) | LCCN 2024026595 (ebook) | ISBN 9781394185726 (hardback) | ISBN 9781394185733 (adobe pdf) | ISBN 9781394185740 (epub)Subjects: LCSH: Evolution (Biology) | Life cycles (Biology)Classification: LCC QH366.2 .L544 2025 (print) | LCC QH366.2 (ebook) | DDC 576.8–dc23/eng20240809LC record available at https://lccn.loc.gov/2024026594LC ebook record available at https://lccn.loc.gov/2024026595
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List of Contributors
Luis Abdala‐RobertsDepartamento de Ecología TropicalCampus de Ciencias Biológicas y AgropecuariasUniversidad Autónoma de YucatánMéridaMéxico
Jun AbeFaculty of ScienceKanagawa UniversityYokohamaJapan
Paul K. AbramAgriculture and Agri‐Food CanadaAgassiz, BCCanada
Megan ArnotDepartment of AnthropologyUniversity College LondonLondonUK
Maud Bernard‐VerdierLeibniz Institute of Freshwater Ecology and InlandFisheries (IGB)BerlinGermany
Institute of BiologyFreie Universität BerlinBerlinGermany
Centre d’Ecologie et des Sciences de laConservation (CESCO)Sorbonne UniversitéParisFrance
Michael B. BonsallDepartment of BiologyUniversity of OxfordOxfordUK
Dries BonteDepartment of BiologyGhent UniversityGentBelgium
Jean‐Marc BonzomInstitut de Radioprotection et de Sûreté Nucléaire (IRSN)PSE‐ENV/ SERPEN/LECO, CadaracheSaint Paul Lez DuranceFrance
Renee M. BorgesCentre for Ecological SciencesIndian Institute of ScienceBangaloreIndia
Elad ChielDepartment of Biology and EnvironmentUniversity of Haifa‐OranimTivonIsrael
Alison B. DuncanISEMUniversity of Montpellier, CNRS, IRD, EPHEMontpellierFrance
Marco Festa‐BianchetDépartement de biologieUniversité de SherbrookeSherbrooke, QuébecCanada
Yuval GottliebKoret School of Veterinary MedicineThe Hebrew University of JerusalemRehovotIsrael
George E. HeimpelDepartment of EntomologyUniversity of MinnesotaSt. Paul, MNUSA
Andreas HeylandIntegrative BiologyUniversity of GuelphGuelph, ONCanada
Yuval ItescuLeibniz Institute of Freshwater Ecology and InlandFisheries (IGB)BerlinGermany
Institute of BiologyFreie Universität BerlinBerlinGermany
Department of Evolutionary and Environmental BiologyUniversity of HaifaHaifaIsrael
Jonathan M. JeschkeLeibniz Institute of Freshwater Ecology and InlandFisheries (IGB)BerlinGermany
Institute of BiologyFreie Universität BerlinBerlinGermany
Oliver KaltzISEMUniversity of Montpellier, CNRS, IRD, EPHEMontpellierFrance
Tamar KeasarDepartment of Biology and EnvironmentUniversity of Haifa‐OranimTivonIsrael
Konstantin KhalturinInstitute of Cellular and Organismic Biology (ICOB)Academia SinicaTaipeiTaiwan
Miriam KishinevskyDepartment of Integrative BiologyUniversity of Wisconsin – MadisonMadison, WIUSA
Hanna KokkoInstitute for Organismic and Molecular EvolutionJohannes Gutenberg University of MainzMainzGermany
Judith KorbEvolutionary Biology & EcologyUniversity of FreiburgFreiburgGermany
Research Institute for the Environment and LivelihoodsCharles Darwin UniversityCasuarina CampusNT0909 DarwinAustralia
Hope KlugBiology, Geology, and Environmental ScienceUniversity of Tennessee at ChattanoogaChattanooga, TNUSA
Tzlil LabinDepartment of Evolutionary and Environmental BiologyUniversity of HaifaHaifaIsrael
Vincent LaudetInstitute of Cellular and Organismic Biology (ICOB)Academia SinicaTaipeiTaiwan
Marine Eco‐Evo‐Devo UnitOkinawa Institute of Science and Technology (OIST)OkinawaJapan
Lei LvSchool of Environmental Science and EngineeringSouthern University of Science and TechnologyShenzhenChina
Division of Ecology and Evolution, Research School ofBiologyAustralian National UniversityCanberra, ACTAustralia
Ruth MaceDepartment of AnthropologyUniversity College LondonLondonUK
Toulouse School of EconomicsInstitute for Advanced Study at ToulouseToulouseFrance
Alexei A. MaklakovSchool of Biological SciencesUniversity of East AngliaNorwichUK
Dustin J. MarshallSchool of Biological SciencesMonash UniversityClayton, VictoriaAustralia
Juha MeriläEcological Genetics Research Unit, Organismal andEvolutionary Biology Research Programme, FacultyBiological & Environmental SciencesUniversity of HelsinkiHelsinkiFinland
Area of Ecology and Biodiversity, School of BiologicalSciencesThe University of Hong KongHong KongHong Kong SAR
Jessica E. MetcalfDepartment of Ecology and Evolutionary BiologyPrinceton UniversityPrinceton, NJUSA
Xoaquín MoreiraMisión Biológica de Galicia (CSIC)“Ecology and Evolution of Plant‐HerbivoreInteractions” GroupPontevedra, GaliciaSpain
Volker NehringEvolutionary Biology & EcologyUniversity of FreiburgFreiburgGermany
Ryan L. PaulDepartment of HorticultureOregon State UniversityCorvalis, ORUSA
Ben L. PhillipsSchool of Molecular and Life SciencesCurtin UniversityPerthAustralia
Loïc QuevarecInstitut de Radioprotection et de Sûreté Nucléaire (IRSN)PSE‐ENV/ SERPEN/LECO, CadaracheSaint Paul Lez DuranceFrance
Denis RéaleDépartement de sciences biologiquesUniversité du Québec à MontréalMontréal, QCCanada
Michal SegoliMitrani Department of Desert EcologyBlaustein Institutes for Desert ResearchBen‐Gurion University of the NegevMidreshet Ben‐GurionIsrael
Stephen C. StearnsDepartment of Ecology and Evolutionary BiologyYale UniversityNew Haven, CTUSA
Justin K. SheenDepartment of Ecology and Evolutionary BiologyPrinceton UniversityPrinceton, NJUSA
Toomas TammaruDepartment of ZoologyInstitute of Ecology and Earth SciencesUniversity of TartuTartuEstonia
Tiit TederDepartment of ZoologyInstitute of Ecology and Earth SciencesUniversity of TartuTartuEstoniaDepartment of EcologyFaculty of Environmental SciencesCzech University of Life Sciences PraguePrague‐SuchdolCzech Republic
Joseph TravisDepartment of Biological ScienceFlorida State UniversityTallahassee, FLUSA
Elodie VerckenBiologie des Populations Introduites, UMR 1355INRAE‐CNRS‐UniCAInstitut Sophia AgrobiotechSophia AntipolisFrance
Eric WajnbergINRAE, Sophia Antipolis, FranceINRIA, Project Hephaistos, SophiaAntipolis, France
Departamento de Entomologia e AcarologiaUniversidade de São PauloEscola Superior de Agricultura “Luiz de Queiroz”Piracicaba, São Paulo, Brazil
Stuart A. WestDepartment of BiologyOxford UniversityOxfordUK
Giacomo ZilioCEFEUniversity of Montpellier, CNRS, EPHE, IRDMontpellierFrance
Foreword
Some issues raised in life history evolution were foreshadowed in the 19th century by Darwin and Weismann and in the early 20th century by Pearl. The impetus they received in the mid‐20th century from Lack, Cole, MacArthur, Wilson, Williams, Gadgil, Charlesworth and others stimulated a body of research that took off in the last quarter of the 20th century. After initial interest in broad comparative patterns explained by analogies to population dynamics (r‐ and K‐selection), the focus shifted to theory couched in terms of age‐ and size‐specific birth and death rates, to experimental evolution in both field and laboratories and to phenotypic plasticity within generations as well as genetic responses across generations. By the early 1990s, the body of work had become significant enough to be summarized in several books. Since then, the insights and intellectual toolkit of life history evolution have penetrated evolutionary and behavioural ecology, and its applications now inform fisheries and game management, biological pest control, conservation biology, gerontology and evolutionary medicine.
Life history evolution explains the evolution of specific traits, but it also raises larger issues. The traits explained include age and size at maturity, the number of reproductive events per lifetime, the number of offspring per reproductive event (clutch or litter size), the rate of ageing, lifespan, sex allocation and all their plastic responses. The larger issues raised include how to formulate a theory of phenotypic evolution, how to connect micro‐ with macro‐evolution, and – the original and continuing concern of life history evolution – how evolution designs phenotypes for reproductive success. At its interface with ecology, life history evolution plays a central role in research on eco‐evolutionary feedbacks. At its interface with comparative biology, it is energized by molecular phylogenetics. Its insights into organismal design are now being woven together with evolutionary physiology and genomics. These enduring contributions to several fields have made life history evolution a very successful intervention.
The present book summarizes the state of play at the end of the first quarter of the 21st century. The hard work done by its editors has paid off in a volume that is comprehensive, stimulating and useful.
This foreword addresses three questions: (1) What did I learn? (2) What issues should be mentioned that are not reviewed here? and (3) What do we not yet know?
I learned that work on the maturation event and its reaction norm should now emphasize theoretical and empirical studies of the coevolution of life histories with physiology and development. In the evolution of ageing, one question that remains unanswered is whether the apparent immortality of some coelenterates might be explained by aspects of their development and maintenance not shared with bilaterians. In marine invertebrates, development rate is more sensitive to temperature than metabolic rate, which implies that small increases in temperature decrease the costs of offspring development, allowing parents to decrease the size of their offspring and increase their fecundity. Insect parasitoids contrast adaptations to spatial vs. temporal stochasticity, allowing an intriguing dissection of risk spreading. Could experimental evolution demonstrate that adaptation to spatial risk maximizes arithmetic mean lifetime reproductive success, whereas adaptation to temporal risk maximizes the geometric mean? Daniel Bernoulli would have been intrigued.
The strikingly long lifespans of termite, ant and naked mole rat queens raise the issue of whether degree of sociality is related to longevity, either as a correlate or as a determinant. Antagonistic pleiotropy between the reproductive performance of queens and the mortality of workers helps to explain the striking difference in their lifespans. The molecular mechanisms of ageing in social insects hold an important lesson: ‘although [they] are highly conserved and shared among nematodes, insects and mammals, they can become uncoupled and re‐wired if ultimate forces select for a long lifespan […] in social organisms’.
Viewing menopause as an adaptation is more defensible than I had thought. One question concerns the implications of stringent quality control through follicular atresia. To extend the reproductive lifespan while maintaining the ancestral level of quality control, one must start life with an ovary so large that it might not fit into a foetus.
The analysis of predator–prey interactions on life history evolution in both parties taught me that the costs of plasticity may be quite small. Here the coupling of theory with experiments done in the field has convincingly revealed alternative explanations for earlier conclusions.
The effects of plant defences on life history traits are quite context‐dependent, varying with the annual‐perennial, monoecious‐dioecious, and herbaceous‐woody distinctions, and often mediated by the tensions between pollination, dispersal and defence. I was surprised to learn that specialist pollinators are not better matched to the phenology of the plants they pollinate than are generalist pollinators, and that Grimes’ description of life history patterns for plants – competitive, stress tolerant and ruderal – can be applied to corals and mycorrhizae as well.
Evidence for the impact of global warming on life history traits is abundant and on population growth is scanty. Poikilothermic organisms do appear to be getting smaller as the world warms, with implications for fish fecundity. Physical and chemical pollution can change the expression and shape the evolution of life history traits in ways that are specific to species and situations, with many different outcomes. That some experiments have found little or no effect of pollution at the population level suggests that variation in individual responses could be masking impacts on populations.
The eco‐evolutionary dynamics of life history evolution on the expansion fronts of invading species reveal the distinction between pushed and pulled expansion fronts and the option of using genetic backburn to slow range expansion. Urban environments challenge organisms that evolved in more natural habitats, creating a mismatch problem like those humans faced as they domesticated themselves and created the diseases of civilization. Work on the biological control of arthropods and weeds has great potential to illuminate life history evolution in general by using experimental evolution to enhance performance and by showing how control agents evolve after they are released into new habitats.
In game and fisheries management, optimal harvesting strategies, based on the elasticity of population growth rates to changes in age‐specific mortality, can be counter‐intuitive. Protecting lactating females in chamois is not the best idea, and where there is sexually selected infanticide, as in African lions and European brown bears, killing a dominant male as a trophy can have large indirect impacts because the remaining males kill juveniles to bring females into oestrus more quickly.
Because life history traits are the central mediators of natural selection, it is not surprising that many other traits connect to evolution through them. The initial simplifying insights of the field fifty years ago have since been complicated in many productive ways. Whether one regrets the loss of simplicity or celebrates the richness of nuanced diversity is probably beside the point: complexity is what progress in our understanding has brought us. We should be grateful for the overarching structure that ties together so many details.
The use of life history thinking in applied fields such as fisheries management, pollution analysis, biological control and conservation ecology has great potential to qualify and even change basic theory, for there the economic incentives are great, funding can be long‐term, and the motivation to dig into details is strong.
While this book is quite comprehensive, it does leave out some issues that may be worth addressing in the future. In 1980 I made the mistake of burying an eco‐evolutionary analysis of the coevolution of seed dispersal, lifespan, reproductive effort and body size in forest trees in an obscure publication (Stearns and Crandall 1981). It suggests an explanation for why there are trees and why there are forests. It has not been followed up and should be revisited. The questions are important; the initial insights need qualification.
The insights of life history evolution also impact evolutionary medicine in several ways, among them are: (1) Trade‐offs help to shape cancer evolution under chemotherapy along the familiar fast‐slow axis, with important implications, and the analysis of phenotypic plasticity helps us to understand the role of stem cells in initiating cancer (Aktipis et al. 2013). (2) The endocrine system of vertebrates is structured to mediate life history trade‐offs. Wang et al. (2019) show how the evolutionary design of organisms for reproductive success embeds principles of systems design into their physiology, thereby suggesting a general approach to evolutionary physiology. Similar work is needed on plants and insects. (3) There are interesting parallels between urban evolution and the mismatch theme in evolutionary medicine: can organisms in cities be models for human mismatch to the environments that we have created? That deserves systematic exploration.
Acknowledging ignorance is a first step towards future research. Here are three things about life histories that we do not yet understand: (1) Why are bilaterians not able to remodel their tissues as do some coelenterates and thereby extend their lifespan, perhaps indefinitely? (2) How do complex life cycles – like those found in algae, jellies and parasites with multiple hosts – evolve? What governs the number of stages and the duration of each? (3) How can we explain major phylogenetic events, such as internal fertilization, endothermy or clutch sizes fixed at one, and their consequences for the subsequent evolution of life history traits?
Stephen C. StearnsMay 2024
References
Aktipis, C.A., Boddy, A.M., Gatenby, R.A., Brown, J.S., and Maley, C. (2013). Life history trade‐offs in cancer evolution. Nature Reviews Cancer 13: 883–892.
Stearns, S.C., and Crandall, R.E. (1981). Bet‐hedging and persistence as adaptations of colonizers. In: Evolution today. Proceedings of thew second international conference of systematic and evolutionary biology (eds. Scudder, G.G.E., and Reveal, J.L.), 371–383. Hunt Institute for Biological Documents.
Wang, A., Luan, H.H., and Medzhitov. R. (2019). An evolutionary perspective on immunometabolism. Science 363: 140.
Preface
Life history traits are defined as those related to the timing and magnitude of major events in the life of an organism (e.g., birth, maturation, reproduction and death). Life history theory aims to explain the huge variation in these trait values and their combinations that we see in nature. Hence, life history strategies are at the heart of evolutionary biology and ecology, illuminating how organisms are structured and how they allocate resources, behave and evolve, in response to different environmental pressures.
Throughout the last decade, the study of life history evolution has experienced some major advances: First, while classical life history theory relied on optimization approaches and had some major empirical successes, more advanced approaches involving frequency‐ and density‐dependency, explicit population dynamics and evolutionary game theory, have further promoted our understanding in the field. Second, classical theory often emerged from observations of life history patterns in a specific group of animals (e.g., clutch size in birds). Later work extended and further developed these principles to the study of various organisms, including vertebrates, invertebrates (in particular arthropods), plants, fungi, bacteria, and even phages. At the same time, concepts from life history evolution were used to promote our understanding of human evolutionary ecology. Third, classical life history theory often focused on a few fundamental life history traits, most commonly including body size, lifespan, investment in reproduction and sex ratio. Later works extended our view to consider additional traits such as parental care, life cycles, sociality and dispersal, either as life history traits in their own right or in the context of their interaction with major life history traits. Fourth, while classical life history theory often addressed the evolution of each organism independently, later advances extended this approach to consider life history evolution also in the context of interspecific and even community‐level co‐evolutionary interactions, including predator–prey, parasite–host, plant–herbivore and mutualistic interactions. Fifth, with the advances in molecular, genetic and physiological methods, we can better understand the underlying proximate mechanisms of life history patterns. Sixth, while theory often focuses on evolutionary responses to environmental conditions, we now have accumulating evidence that evolutionary processes may occur at a short enough time scale to allow eco‐evolutionary feedback. Finally, there is an increasing awareness that life history evolution responds to anthropogenic‐induced changes such as climate change, pollution and urbanization and that insights from these fields could be incorporated into how we manage natural resources globally.
Several foundational books have laid the groundwork for the analysis of life history evolution. However, a comprehensive, up‐to‐date volume summarizing recent advances in the field – as highlighted above – and presenting current examples and applied aspects was lacking. The current volume aims to fill this gap by presenting current ideas, analyses, and case studies in life history evolution, spanning a wide range of taxa (from bacteria to insects to humans), traits and applications.
This volume starts with a foreword by Stephen C. Stearns, Yale University, one of the leading scientists in this field and the author of several books on life history evolution theory. The volume is then organized into three parts. The first addresses different traits that are often considered, and a few that are less commonly considered, in the study of life history evolution. The second part focuses on life history evolution in the context of interspecific interactions, and the third part discusses anthropogenic impacts and possible applications. In the first part, the first three chapters present recent developments in the study of classical life history traits, namely, body size and the timing of maturation, lifespan and offspring size and number. The fourth chapter discusses optimized egg loads focusing on parasitoid insects. Two chapters then address the question of sex‐specific life histories and parental care, and an additional chapter focuses on the complexity of life cycles. Finally, three additional chapters discuss life history in social groups, in relation to dispersal, and the particular case of humans focusing on menopause. The second part then starts with three chapters presenting recent advances in life history evolution in the context of predator–prey, host–parasite and host–endosymbiont interactions. Three more chapters discuss life history evolution in the context of plant–herbivore, plant–pollinator, and mutualistic interactions. Finally, the last part starts with two chapters dealing with the effects of human‐induced stresses such as climate change and pollution on the evolution of life history traits of different organisms. Two more chapters address the case of species range expansions and urban environments, followed by three final chapters discussing applications in biological control programmes against crop pests, wildlife management and disease management.
Throughout the volume, the goal of each chapter is to present the main current ideas and developments in the field and to give the readers the main information required for further investigation. Each chapter ends with a brief description of remaining knowledge gaps, listing several open questions to be addressed in the future.
We hope that this volume will broaden the scope of previous publications by discussing life history evolution of both classical life history traits, and in relation to behavioural, morphological and physiological traits (Part I), in the context of specific interactions (Part II), and its applications for resource and disease management (Part III).
We express our gratitude to all authors for their excellent contributions and their collaboration throughout the edition of this volume. A special thanks to Stephen C. Stearns for writing the foreword section for this book. All chapters underwent a reviewing process to increase their relevance and foster links between them. In this respect, we thank the referees for their time and insightful comments. These include Carlos Barata, Javier Belles, Judith Bronstein, Julien Cote, Damian K. Dowling, Anja Felmy, Jessica Forrest, Jean‐Michel Gaillard, Megan Greischar, Ian Hardy, George Heimpel, Ruth Hufbauer, Richard Karban, Boris Krasnov, Kevin Lafferty, Yael Lubin, Allen Moore, Michael Moore, Michael Poulsen, John Reynolds, Jay Rosenheim, Locke Rowe, Inon Scharf, Rebecca Sear and Elsa Youngsteadt.
Lastly, we also express our thanks to the editors of Wiley Publishing for their continuous help and guidance to the IIAS (Hebrew University, Jerusalem) which hosted both of us for five month during the initial phase, and to Fapesp (São Paulo State, Brazil) which hosted one of us (Eric Wajnberg) for eight months during the final phase of the preparation of this book (Process # 2022/10870‐1).
Michal Segoli and Eric WajnbergMay 2024
Part ITraits
1Body Size and Timing of Maturation
Toomas Tammaru1 and Tiit Teder 1,2
1 Department of Zoology, Institute of Ecology and Earth Sciences, University of Tartu, Tartu, Estonia
2 Department of Ecology, Faculty of Environmental Sciences, Czech University of Life Sciences Prague, Prague-Suchdol, Czech Republic
1.1 Introduction
Body size is perhaps the most apparent trait of any organism, at least from the perspective of visually oriented observers, including us, humans. However, the implications of body size reach far beyond the limits of visual appearance as basic physical principles determine where and how an organism of a certain size can or cannot live. Just as a few examples, the circulation of vital substances within the body is critically dependent on the size of the organism, as are the ways an organism can or cannot move around in its environment. From a more ecological perspective, size largely determines the range of items one can include in its diet, as well as the range of the consumers one should be afraid of. Proceeding to more derived traits, body size sets the limits for permissible colouration patterns and adequate behavioural repertoire, in the context of both inter‐ and intraspecific interactions.
Since the dawn of life history theory as a separate discipline, the evolutionary explanation of adult body size has been viewed in the framework of optimality analysis of age and size at maturation. It is indeed straightforward to ask when it pays to start investing in reproduction instead of continuing the investment in one’s own somatic growth (Figure 1.1). Such an approach allows us to conveniently break down the question about the optimal body size into the analysis of costs and benefits of early and late maturation and costs and benefits of large and small size per se.
In the present chapter, we briefly review the knowledge about such costs and benefits, and how the optimal balance between them may depend on environmental conditions. We conclude that, while we can often provide satisfactory evolutionary explanations for the differences in size, e.g., the differences between biological sexes, among populations or among related species, we may not know enough to explain absolute values. To be able to do so, we may need a tighter integration of the ecologically based optimality approach with the study of physiological and ontogenetic phenomena, often termed constraints in the research tradition of evolutionary ecology. Furthermore, it is crucial to learn how fast body size and associated traits evolve.
The chapter consists of two parts. We first provide an overview of past and current developments in the field (Part I), and thereafter focus on a few selected topics related to our research interests (Part II). We illustrate a number of key points with examples from our own work, which we happen to know best. More generally, due to the ‘ontogenetic constraints’ of the authors, our approach will be biased towards insects. This should not be a big problem as multicellular species other than insects form just a minority among the known biodiversity. We will nevertheless try not to ignore them.
1.2 Part I
1.2.1 Benefits of Large Body Size
For many organisms, the benefits of large size are straightforward to understand. A frequent fitness benefit of large size in females is the so‐called fecundity advantage. This implies that larger‐bodied females are expected to produce more offspring than smaller conspecifics (see Lim et al. (2014) for a meta‐analysis, and Pincheira‐Donoso and Hunt (2017) for a critical discussion of the concept of fecundity advantage). This can primarily result from large females having more resources to be converted into offspring or because they have better capabilities to obtain such resources. The pattern is clear and straightforward at least for many insects (Honěk 1993), some other invertebrates (reviewed in Roff 2002) and fish (Barneche et al. 2018).
Figure 1.1 Left panel: conceptual setting for optimality analysis of age and size at maturation. We can ask when a juvenile should stop growing and switch to reproduction. Right panel: a simple model for optimal age and size at maturation for an organism with determinate growth. Size is assumed to increase proportionally with growing time (not shown). Fecundity is a linear function of size at maturation with a negative intercept. Probability of survival is a negative exponential function of time. Fitness is the product of survival to maturation and fecundity, the function attains its maximal value at some intermediate (optimal) value of time at maturation.
The strength of the size–fecundity relationship naturally differs between different organisms and not in a random manner. One aspect to consider is the relative role of having more resources vs. the capability of obtaining them. This distinction can be illustrated by the contrast between capital‐ and income‐breeding species (Drent and Daan 1980, Jönsson 1997). As applied to insects (Davis et al. 2016), the contrast is between females, which primarily depend on larvally derived resources (capital) in their reproduction, as opposed to females obtaining such resources through adult foraging (income). In capital‐breeding insects, the correlation between body size and potential fecundity is often remarkably strong (e.g., Tammaru et al. 1996b, 2002). This is because the mass of a newly eclosed female directly determines the amount of resources she can use for reproduction.
In females of capital‐breeding insects, there is no need to move around in search of food. Amplified by the trade‐off between fecundity and mobility, this has frequently led to the evolution of short adult life spans and overall behavioural simplification (Tammaru and Haukioja 1996, Davis et al. 2016). In such females (Figure 1.2), realized fecundity is largely determined by potential fecundity, as factors related to adult behaviour have little effect on the number of eggs laid. In contrast, in income‐breeding insects with active long‐living adults, the number of eggs laid should primarily depend on foraging success, efficiency in finding suitable oviposition substrates and survival rates. These fitness correlates may not need to depend strongly on body size, which can lead to substantial weakening of the fecundity advantage in income‐breeding insects (Gotthard et al. 2007).
Figure 1.2 An ovipositing flightless female of the erebid moth Orgyia antiqua. The adult female does not move away from its pupal case and lives just for a few days. No costs of large adult size were found in this species, and realized fecundity is proportional to body mass (Tammaru et al. 2002). With such a reduced behavioural repertoire, there is little ‘space’ for costs of large adult size to operate.
Source: © entomart.
In an attempt to generalize from the comparison between capital‐ and income‐breeding insects, we propose that the strong fecundity advantage should primarily be characteristic of animals in which females have simple behavioural repertoires. This suggestion is supported by the observations that the fecundity advantage of large female size may be weak, non‐existent or even reversed in groups known for complex behavioural patterns, such as birds (Blums and Clark 2004, Magrath et al. 2009) and mammals (Isaac 2005), including humans (Valge et al. 2022). Plants represent the other extreme of behavioural complexity, and accordingly, there is typically a strong correlation between the size of a plant individual and its seed set (Shaanker et al. 1988, Weiner et al. 2009). Notably, however, in plants, the amount of resources readily available for reproduction and the ability to obtain further resources are both directly size‐dependent.
Obviously, the fecundity advantage of large size is not the only source of the positive correlation between body size and fitness in females. Larger females may also produce offspring of better quality, partly through providing better‐quality maternal care (Clutton‐Brock et al. 1985, 1988; see also Chapter 6). In their meta‐analysis, Lim et al. (2014) found evidence of a positive intraspecific correlation between maternal size and offspring size, broadly consistent across major taxa and environments. Such a relationship appears, however, not to be typical of insects (Fox and Czesak 2000). Large body size may also contribute to fitness through increased longevity in animals as different as flies (Reim et al. 2006) and primates (Blomquist et al. 2011; see also Chapter 2). Additionally, in some animal groups, males may actively prefer large females (Bonduriansky 2001).
For males, the advantages of large body size are frequently less clear, also being taxon‐specific to a greater extent. This is primarily due to variations in the behavioural contexts of mate location and mate choice (see Choe and Crespi 1997, for the high diversity of mating systems in insects alone). Most typically, the higher reproductive output of larger males can be due to greater success in male‐to‐male competition or female choice (Shine 1989, Roff 1992, Andersson 1994, Dale et al. 2007). Opposite to the situation in females, we can therefore expect a stronger correlation between male size and fitness in species with more complex behavioural repertoires. In capital‐breeding lepidopterans with simplified adult behaviours (Figure 1.2), for example, there may be no male‐to‐male interaction or female choice involved in the process of forming mating pairs (Tammaru et al. 1996a, van Dongen et al. 1998). This contrasts with the often much more complex behavioural interactions in butterflies, the most well‐known income‐breeding lepidopterans (Wiklund and Kaitala 1995, Kemp and Wiklund 2001), and various income‐breeding moths (Phelan and Baker 1990, Svensson 1996). This difference may have left an imprint in the evolution of sex‐specific body size: capital‐breeding insects are generally characterized by strong female‐biased sexual size dimorphism (SSD, females larger than males; Davis et al. 2016), while in butterflies, males can even be the larger sex (Wiklund and Forsberg 1991, Teder 2014). Such contrasts can also be made at higher taxonomic levels as insects in general tend to have female‐biased SSD (Teder and Tammaru 2005) compared to the tendency towards males being larger than females in birds and mammals (Lindenfors et al. 2008). It appears likely that the opposite effect of behavioural complexity on fitness consequences of body size in the two sexes is largely responsible for this broad‐scale pattern (see also Dale et al. 2007, Dial et al. 2008).
1.2.2 Costs of Large Body Size
The straightforward and often strong positive correlation between body size and major components of fitness raises the question about selective pressures counterbalancing the fitness advantage of large body size. Without such costs, we should expect a continuous evolutionary increase in body size. Such trends are sometimes indeed observed on the scale of tens of millions of years (known as Cope’s rule: Kingsolver and Pfennig 2004, Hone and Benton 2005, Roy et al. 2024). Nevertheless, respective macro‐evolutionary changes are still slower by many orders of magnitude than what could be expected on the basis of the nearly proportional relationship between body size and the number of eggs laid in capital‐breeding insects, for example. In other words, the evolutionary stability of body size calls for an explanation (Hansen and Houle 2004, Rollinson and Rowe 2015).
The costs of large body size are usually much more challenging to see than the advantages. The available evidence of such costs was reviewed and synthesized by Blanckenhorn (2000). First, life history models usually assume that it takes more time to grow larger (see further in the chapter for a discussion of this assumption). This being the case, juveniles attempting to attain larger adult sizes are subjected to higher cumulative mortality risk and have lower chances of achieving maturity (see Teder and Kaasik 2023 for a consistent pattern). Because mortality is never zero, this principle cannot perhaps be questioned in its general form. As juvenile mortality rates are frequently dramatically high, for example, in fish (Moyle and Cech 2004) and herbivorous insects (Cornell and Hawkins 1995, Remmel et al. 2011), the survival costs of attaining large size must correspondingly be substantial. There is also empirical evidence of high predation rates causing evolution towards smaller body sizes (see, e.g., Edley and Law 1988, Reznick et al. 1996, Stearns et al. 2000 for experimental studies, and the voluminous literature on selective effects of fishing, discussed further in the chapter; see also Chapters 12 and 23).
Besides the costs of attaining a large size, there may also be costs of being a large adult (Kozlowski 1991, Blanckenhorn 2000
