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Spider Webs

Behavior, Function, and Evolution

In this lavishly illustrated, first-ever book on how spider webs are built, function, and evolved, William Eberhard provides a comprehensive overview of spider functional morphology and behavior related to web building, and of the surprising physical agility and mental abilities of orb weavers. For instance, one spider spins more than three precisely spaced, morphologically complex spiral attachments per second for up to fifteen minutes at a time. Spiders even adjust the mechanical properties of their famously strong silken lines to different parts of their webs and different environments, and make dramatic modifications in orb designs to adapt to available spaces. This extensive adaptive flexibility, involving decisions influenced by up to sixteen different cues, is unexpected in such small, supposedly simple animals.

As Eberhard reveals, the extraordinary diversity of webs includes ingenious solutions to gain access to prey in esoteric habitats, from blazing hot and shifting sand dunes (to capture ants) to the surfaces of tropical lakes (to capture water striders). Some webs are nets that are cast onto prey, while others form baskets into which the spider flicks prey. Some aerial webs are tramways used by spiders searching for chemical cues from their prey below, while others feature landing sites for flying insects and spiders where the spider then stalks its prey. In some webs, long trip lines are delicately sustained just above the ground by tiny rigid silk poles.

Stemming from the author’s more than five decades observing spider webs, this book will be the definitive reference for years to come.

See additional online materials for the book.

816 pages | 195 halftones, 97 line drawings | 8-1/2 x 11 | © 2020

Biological Sciences: Behavioral Biology, Biology--Systematics, Ecology, Evolutionary Biology


"The big book on this exciting subject, lavishly illustrated and based on over five decades of research. Eberhard answers any query about the origins, morphology and utilization of spicier webs, as well as their weavers' surprising mental and physical capabilities." 


"You would think that after centuries of studying spider webs we have a pretty good grasp of them. Yet a thorough, book-length review of their construction, function, and evolution has been missing. Eberhard has taken on that colossal task, based on his nearly fifty years of observing spiders and their webs. Some works go on to define their discipline. Spider Webs has all the trappings of becoming the arachnological benchmark. . . . It is one of those rare books that sets a discipline’s research agenda for decades to come and cannot be ignored. This book has set the bar and has set it high. For professional arachnologists this is a vital and invaluable reference work. For serious spider enthusiasts it is an absolute treasure trove. Beyond the spider community, however, there is much here of interest to ethologists and evolutionary biologists."

Leon Vlieger | Inquisitive Biologist

“In summary, what does one get by buying the book? Spider silk is often compared to steel, weight by weight and materially speaking this book is excellent value for money at just under 2 kg weight and just under 10 cents per A4 page. Intellectually speaking, Eberhard’s book not only provides the reader with an exceptionally comprehensive, generous, and intuitive text supported by well explained illustrations that lighten the text, and overall aided by an outstanding index that allows for flitting between subjects and chapters. As importantly, there is plenty of intellectual stimulation for both spider and non-spider experts. And the honest discussion of the many open questions will provide generations of students interested in behavioral ecology with topics for research. . . . The editor and team [at the University of Chicago Press] must be congratulated not only for publishing this true labor of love but also for the superb production nowadays so rare for an academic book of this topic and scope.”

Fritz Vollrath | ISBE Newsletter

“Based on half a century of study, Spider Webs is an unprecedented and very large book on [spiders’] biology, evolution, and diversity.”

Natural History Book Service

“This is a wonderful book. In it Eberhard explores the mechanical and evolutionary aspects of spider webs from all sides. The book is lavishly illustrated with photographs of webs appearing throughout. Each chapter has a down-to-earth summary that repeats the main points of the chapter. Eberhard is not afraid to rail against the many errors that infect previous work on spiders and webs, including adultophilia (the tendency to ignore the webs of juvenile spiders), typology (the eagerness to describe ‘the’ spider and its web when in reality there are many variations in the webs of any one species), and many more shortcomings. But by far the strength of the book is Eberhard’s ability to tell his readers what is needed on topics A to Z if we are to test hypotheses further. This willingness to lay out predictions that are derived from particular hypotheses on puzzling behaviors that have not been checked makes the book a gold mine for students of spider webs and indeed for all animal behaviorists.”

John Alcock, Arizona State University, author of “Animal Behavior: An Evolutionary Approach”

“There is only one comparable book on this topic that was published over thirty years ago, and it is not comprehensive. Eberhard’s book is an extensive, encyclopedic review of what is known (and unknown) about spider webs today. It also provides many original observations from Eberhard himself.”

Rainer F. Foelix, author of “Biology of Spiders”

“A comprehensive book on all aspects of spider webs is long overdue, and Eberhard is one of the few biologists—if not the only one—who has the knowledge, passion, and expertise to write such a book. Very thorough and detailed, Spider Webs is of great importance to specialists in its field, especially since it not only summarizes previously published studies, but also presents new ideas and views on structure, construction, function, and evolution of spider webs and their (potential) relevance to many different fields of biology.”

Samuel Zschokke, University of Basel, Switzerland

Spider Webs provides both an authoritative and unique synthesis of research and still-to-be-addressed questions on spider web biology. There is no one better equipped to write a book on this subject than Eberhard. It will be a major contribution to the fields of ecology, evolution, and behavior, providing new information, new syntheses, and new insights on a broad range of topics.”

Brent Opell, Virginia Tech

Table of Contents

Chapter 1. Introduction
1.1 Introduction
1.2 A foreign world: life tied to silk lines
1.3 A brief history of spider web studies
1.4 Emphasis on behavior
1.5 The scope of this book and tactics in presentation
1.6 Evolutionary history and phylogeny
1.7 Terminology and other procedural matters
1.8 Acknowledgments

Chapter 2. The “hardware” of web-building spiders: morphology, silk, and behavior
2.1. Introduction
2.2 Silk glands and silk
2.3 Spinnerets as high-precision instruments
2.4 Leg morphology and behavior: grasping lines precisely and securely
2.5 Cutting lines and recycling silk
2.6 How spiders avoid adhering to their own webs: a mystery partly solved
2.7 Central nervous system basis for web construction
2.8 Summary

Chapter 3. Functions of orb web designs
3.1 Introduction
3.2 Correcting common misconceptions about orb webs
3.3 How orbs function
3.4 Summary

Chapter 4. Putting pieces together: tradeoffs and remaining puzzles
4.1 Introduction
4.2 “Optimal” orb designs: tradeoffs between functions are difficult to measure
4.3 “Multiple trap” design: a new way to view orb webs
4.4 Tensions and stresses
4.5 Relative numbers of radii and sticky spiral loops
4.6 Testing visibility and stopping functions: the extreme case of trunk orbs
4.7 Correlations between spider size and orb design?
4.8 Spider positions, attack behavior, and up-down asymmetries in orbs
4.9 Remaining puzzles
4.10 Non-orb webs
4.11 Evolutionary responses by insects? A neglected aspect of prey capture
4.12 Summary (including part of chapter 3)

Chapter 5. The building behavior of non-orb weavers
5.1 Introduction
5.2 Order of lines and other higher-level patterns
5.3 Lower-level patterns: leg movements and manipulation of lines
5.4 Stereotyped behavior in non-orb construction
5.5 Adjustments to substrate-imposed constraints
5.6 Managing swaths of fine lines
5.7 Summary
Box 5.1 The funnel web diplurid Linothele macrothelifera

Chapter 6. The building  behavior of orb-weavers
6.1 Introduction
6.2 Simplifications for smoother reading
6.3 Behavior of two araneids
6.4 Senility in orb construction: a new frontier?
6.5 Detailed movements
6.6 General patterns
6.7 Summary

Chapter 7. Cues directing web construction behavior
7.1 Introduction
7.2 Classifying the cues
7.3 Cues for sticky spiral construction
7.4 Temporary spiral
7.5 Hub
7.6 Stabilimentum construction
7.7 Radii, frames, and anchor lines
7.8 Early radii, and frames and anchor lines: determining web size, shape, and design
7.9 To build or not to build: triggering orb construction and destruction
7.10 Cues that trigger transitions between stages of orb construction
7.11 Other stimuli that spiders can sense but that are not (yet) known to guide orb construction
7.12 Hints of abilities: follow circular paths and sense radius lengths
7.13 Effects of psychotropic drugs on orb construction
7.14 Coordinating different adjustments to different cues
7.15 The (limited) role of simulations in understanding orb construction behavior
7.16 A missing link: translating cues into attachment sites
7.17 Summarizing the behavioral challenges met by orb weavers
7.18 Independence (?) of the spider’s responses
7.19 Changes in responses to cues: learning and maturation
7.20 Cues guiding the construction of non-orbs
7.21 Summary
Chapter 8. Web ecology and website selection
8.1 Introduction: what is and is not included
8.2 Webs and ecological foraging theories
8.3 What is enough? “Fast lane” and “slow lane” spiders
8.4 Processes that produce habitat biases
8.5 A general correlation between website selectivity and web design flexibility?
8.6 Website tenacity, web durability, and recycling
8.7 Web durability
8.8 Limited by websites? Possible competition for prey and websites
8.9 Problems in attempts to study cues that guide website choices
8.10 Time of day: day webs vs. night webs
8.11 Summary

Chapter 9. Evolutionary patterns: an ancient success that produced high diversity and rampant convergence
9.1 Introduction
9.2 Patterns in the diversity of webs
9.3 Consequences of the failure of the prey specialist hypothesis for understanding diversity and convergence
9.4 What is a sheet web? Problems inherited from previous imprecision
9.5 Mygalomorphs: similar patterns of diversity and rampant convergence in a different world
9.6 Diversity of relations with insects
9.7 Lack of miniaturization effects
9.8 Paths not followed: alternative web forms in other animals
9.9 Summary and a new synthesis
Box 9.1 The most spectacular convergence of all: Fecenia
Box 9.2 The most spectacular divergence of all: Theridiidae
Box 9.3 Sand castles: extreme modifications of Seothyra henscheli webs to shifting sand
Box 9.4 Relation between web design and silk properties: stiff silk in Uroctea durandi

Chapter 10. Ontogeny, modularity, and the evolution of web building
10.1 Introduction
10.2 Web ontogeny and evolution
10.3 Early web evolution
10.4 The behavior patterns used to build early webs
10.5 Evolution of later non-orb webs
10.6 Inconsistent evolutionary trends in non-orb webs
10.7 Diversity in non-orbs that results from behavioral stability
10.8 The (probably) monophyletic origin of orb webs
10.9 Evolutionary changes in orb designs
10.10 “Post-orb” web evolution in Orbiculariae
10.11 Coevolution between attack behavior and web design (and its lack)
10.12 What didn’t happen, possible synapomorphies, and further puzzles
10.13 Modularity and adaptive flexibility
10.14 Modules and evolutionary transitions in web-building behavior
10.15 Summary


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