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Zebra Stripes

Coming Soon

By: Tim Caro (Author)

320 pages, 33 plates with colour photos; 28 b/w photos, 55 tables

University of Chicago Press

Hardback | 2017 | #230391 | ISBN-13: 9780226411019
Available for pre-order: Due Jan 2017 Details
NHBS Price: £31.50 $39/€36 approx

About this book

From eminent biologists like Alfred Russel Wallace and Charles Darwin to famous authors such as Rudyard Kipling in his Just So Stories, many people have asked, "Why do zebras have stripes?" There are many explanations, but until now hardly any have been seriously addressed or even tested. In Zebra Stripes, Tim Caro takes readers through a decade of painstaking fieldwork examining the significance of black-and-white striping and, after systematically dismissing every hypothesis for these markings with new data, he arrives at a surprising conclusion: zebra's markings are nature's defense against biting fly annoyance.

Popular explanations for stripes range from camouflage to confusion of predators, social facilitation, and even temperature regulation. It is a challenge to test these proposals on large animals living in the wild, but using a combination of careful observations, simple field experiments, comparative information, and logic, Caro is able to weigh up, scientifically, the pros and cons of each idea. Eventually – driven by experiments showing that biting flies avoid landing on striped surfaces, observations that striping is most intense where biting flies are abundant, and by his knowledge of zebras' susceptibility to biting flies and vulnerability to the diseases that flies carry – Caro concludes that black-and-white stripes are an adaptation to thwart biting fly attack. Not just a tale of one scientist's quest to solve a classic mystery of biology, Zebra Stripes is also a testament to the tremendous value of longitudinal research in behavioral ecology, demonstrating how observation, experiment, and comparative research can reshape our understanding of the natural world.

"A tour de force that characterizes the biology, morphology, physiology, and behavior of the equid genus as a starting point for examining in detail the unique features of its striped members. Caro clearly and succinctly presents the competing hypotheses put forward over the last one-hundred-fifty years (some even earlier) for why zebras are striped. The mix of methods with facts and interpretation is compelling, making what could be a complex and tedious presentation one that is clear, engaging, and to the point. Zebra Stripes is an easy read, full of authoritative documentation from the literature bolstered by clever experiments (with Caro putting himself literally in a zebra's shoes – or actually a pelt), and constructed in a didactic, hypothetical, deductive way that gives it credibility. Its completeness and attention to detail will make it a must read."
– Daniel I. Rubenstein, Princeton University

"Zebra Stripes is an attempt to answer the mysterious, longstanding, and popular question of why zebra have strips. A personal journey in addition to a scientific appraisal of the evidence, it is highly original in its discussion of stripe hypotheses and its personal aspect; I am not aware of any work of comparable synthesis in depth or scope. It is comprehensive, up-to-date, and authoritative, and also contains a wealth of unpublished work and observations. Well written in a very readable style, interesting, and clear, Zebra Stripes will unquestionably be of importance to any specialist in the field and of much interest to biologists more broadly, especially those interested in animal coloration, behavior, and evolution. Highly successful."
– Martin Stevens, University of Exeter, author of Cheats and Deceits: How Animals and Plants Exploit and Mislead



Preface and acknowledgments

Chapter 1. Stripes and equids
1.1. The question of stripes
1.2. Hypotheses for striping in equids
     1.2.a. Antipredator hypotheses
     1.2.b. Antiparasite hypotheses
     1.2.c. Communication hypotheses
     1.2.d. Thermoregulation hypothesis
1.3. Equid evolution
     1.3.a. Plains zebra
     1.3.b. Mountain zebra
     1.3.c. Grevy’s zebra
     1.3.d. African wild ass
     1.3.e. Asiatic wild ass
     1.3.f. Kiang
     1.3.g. Przewalski’s horse
     1.3.h. Other equids
1.4. Zebra hair
1.5. Conclusion

Chapter 2. Predation and crypsis
2.1. Background matching
     2.1.a. Initial discomfort with the idea
     2.1.b. Detecting zebras
2.2. Disruptive coloration
     2.2.a. Predictions
     2.2.b. Sightings at dusk and dawn
2.3. Countershading
2.4. Zebras as seen by nonhumans
2.5. Conclusions

Chapter 3. Predation and aposematism
3.1. Aposematism in mammals
3.2. Signaling component of aposematism
     3.2.a. Visibility
     3.2.b. Noisy behavior
3.3. Defense component of aposematism
     3.3.a. Response to predators
3.4. Conclusion

Chapter 4. Predation and confusion
4.1. Confusion
4.2. Miscounting numbers of prey individuals
4.3. Striping obscuring outlines of fleeing prey
     4.3.a. Lines of stripes shown to humans
     4.3.b. Lines of stripes in dangerous situations
4.4. Striping preventing a single prey individual being followed
4.5. Dazzle effect
4.6. Motion dazzle
4.7. Misjudging the size of prey
     4.7.a. Subjective and estimated heights and girths
     4.7.b. Subjective heights and girths and degree of striping
4.8. Quality advertisement
4.9. Conclusion
4.10. Difficulties with the predation hypothesis

Chapter 5. Ectoparasites
5.1. Biting flies
5.2. Behavioral indices of fly infestation in Katavi
5.3. Behavioral indices of fly infestation in Berlin
5.4. Tsetse fly traps
     5.4.a. Biconical traps
     5.4.b. Cloth traps
5.5. Tabanid traps
     5.5.a. Canopy traps
     5.5.b. Pelt canopy traps
5.6. Moving objects
     5.6.a. Walking in suits
     5.6.b. Walking in pelts
     5.6.c. Driving with pelts
5.7. Conclusions
5.8. Polarized light
     5.8.a. Reflected light
     5.8.b. Horvath’s work
     5.8.c. Polarization signatures of wild zebras

Chapter 6. Intraspecific communication
6.1. Intraspecific signaling
6.2. Species recognition
6.3. Stripes as a facilitator of mutual grooming and social bonding
     6.3.a. Allogrooming
     6.3.b. Social bonding
6.4. Stripes as a means of individual recognition
6.5. Stripes as an indicator of quality
6.6. Conclusion

Chapter 7. Temperature regulation
7.1. Black and white surfaces
7.2. Heat measurements in the field
7.3. Heat management
7.4. Conclusions

Chapter 8. Multifactorial analyses
8.1. Comparing hypotheses simultaneously
8.2. The interspecific comparison
     8.2.a. Comparative methodology
     8.2.b. Overall striping
     8.2.c. Striping on different parts of the body
     8.2.d. Evaluating the hypotheses
8.3. Conclusions
8.4. The intraspecific comparison
8.5. Concordance on multifactorial analyses

Chapter 9. The case for biting flies
9.1. Last man standing
9.2. Host choice
9.3. Ectoparasite population sizes
9.4. Host seeking
9.5. Parasites and diseases transmitted by bloodsucking diptera
9.6. Mechanistic studies
9.7. Further outstanding issues
     9.7.a. Multiple functions
     9.7.b. Loose ends
9.8. Conclusion

Appendix 1. Scientific names of vertebrates mentioned in the text
Appendix 2. Nature of wounding seen in African ungulates in Katavi National Park
Appendix 3. Families of insects identified in each type of biconical trap color
Appendix 4. Families of insects identified in each type of cloth trap color
Appendix 5. Photographic sources for comparative analyses
Appendix 6. Derivation of zebra phylogenies
Appendix 7. Phylogenetic analyses


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Tim Caro is professor of wildlife biology at the University of California, Davis. He is also the author of Cheetahs of the Serengeti Plains: Group Living in an Asocial Species and Antipredator Defenses in Birds and Mammals, both published by the University of Chicago Press, as well as Conservation by Proxy: Indicator, Umbrella, Keystone, Flagship, and Other Surrogate Species.

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