Table Of ContentFascinating Life Sciences
Robert J. Marquis
Suzanne Koptur Editors
Caterpillars
in the Middle
Tritrophic Interactions
in a Changing World
Fascinating Life Sciences
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Robert J. Marquis • Suzanne Koptur
Editors
Caterpillars in the Middle
Tritrophic Interactions in a Changing World
Editors
Robert J. Marquis Suzanne Koptur
Department of Biology Department of Biological Sciences
Whitney R. Harris World Ecology Center International Center for Tropical Botany
University of Missouri–St. Louis Florida International University
St Louis, MO, USA Miami, FL, USA
ISSN 2509-6745 ISSN 2509-6753 (electronic)
Fascinating Life Sciences
ISBN 978-3-030-86687-7 ISBN 978-3-030-86688-4 (eBook)
https://doi.org/10.1007/978-3-030-86688-4
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Chapter “The Natural History of Caterpillar-Ant Associations” is licensed under the terms of the Creative
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Cover image: We thank Sam Jaffe of the Caterpillar Lab ([email protected]) for his cover
illustrations depicting the biology of Synchlora aerata (Geometridae), which cuts plant parts and attaches
them with silk to its dorsal surface. Five Synchlora caterpillars, variously adorned with petals and leaf
fragments, are shown across the first two rows. A fresh brown pupa, a developed and ready to eclose
green pupa, and an adult Synchlora moth are pictured to the bottom left. In the third row, two unique
parasitoid wasps that are closely associated with Synchlora appear: a specialist camplopegine
ichneumonid, and a chalcid hyperparasitoid. The campoplegine’s cocoon, constructed from its host
caterpillar’s cadaver, is above the wasps, at the end of the second row. Finally, to the bottom right a
Synchlora caterpillar is shown actively feeding on a daisy fleabane blossom after decorating itself with
the available petals.
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To our families
Carol, Emma, and Zoe
John and JJ
We love you even more than caterpillars
Foreword
The caterpillar is a poster child for insect holometaboly—a developmental style
that involves a change from a sedentary feeding stage to a resting pupa in which
massive internal changes occur, finally leading to the emergence of an adult. An
important part of the story is that females must make the momentous decisions
on where to place their eggs.
In a caterpillar, growth rate is maximized at the cost of food processing effi-
ciency, and growth is largely independent from the tissue differentiation required to
generate organs for dispersal and reproduction. Its function is to eat, its gut is the
principal organ, and its stretchable cuticular protein is efficiently reused at each
molt. Although the caterpillar life stage emphasizes feeding at the expense of finesse
at fleeing from peril or ability to find scarce food, these predicaments are clearly
offset in the evolution of the holometabolous insect life history. Holometabolous
development is relatively more abundant than the ancestral hemimetabolous pattern.
But risks exist, and the study of caterpillars inevitably involves research on
the multitude of inimitable traits that enhance survival. Caterpillars face threats
from above from numerous species of natural enemies. The dangers are almost
infinite, and their selection pressure has influenced every aspect of caterpillar
biology. Lepidopteran larvae masquerade as any possible non-caterpillar object
in the environment, and in many cases, their defenses against enemies depend
on adaptations to specific host plants. Diverse behavioral traits provide endless
ways to deceive parasites and predators, such as dropping down from the host
plant on silk, flinging frass away from give-away feeding sites, or hiding in
homemade houses. They must feed fast to avoid being noticed and conse-
quently demonstrate mandibular adaptations uniquely suited to the particular
food. Dietary specialists may become masters of chemical protection by
sequestering chemicals that are deterrent or toxic to predators and parasites.
Other specialist species excel at host-specific visual or chemical crypsis, mak-
ing them harder to detect by their enemies. Generalists have found diverse
vii
viii Foreword
mixtures of traits to contend with enemies—opportunistic sequestration of
plant poisons, nimble behaviors, mimicking predators, and even making their
own noxious chemicals. Different species have particular habits of food choice
that enhance protection from particular groups of natural enemies that domi-
nate in their microhabitats.
But caterpillars are “in the middle” in a changing tritrophic world. There are also
hazards from below—the physical and chemical plant defenses. First are the jeop-
ardies associated with physical structures that plants produce to make it difficult to
maintain a purchase, or prevent easy walking, or make biting and chewing too chal-
lenging. There are secretions that are sticky or gluey, such as latex or resin, which
can glue up their mouthparts. A remarkable evolutionary feat of the Lepidoptera has
been the adoption of countermeasures to deal with each of the defenses plants have
come up with.
The all-important plant chemistry has been studied in great depth, with the dis-
covery of innumerable compounds that interfere with caterpillar growth and sur-
vival. These must be avoided or overcome physiologically one way or another. In
addition, the details of sequestration of noxious plant chemicals have been eluci-
dated in numerous species, different chemical structures being of particular value
against different groups of natural enemies.
Plants present a food source that is typically low in protein and variable in nutri-
ent quality. Plants also provide diverse physical challenges and an immense variety
of potentially toxic metabolites that often increase with damage. Environmental
changes involving climate and chemicals from human activity challenge not only
the plants and their availability, but also the caterpillars that feed on them, and the
mothers who must locate their foods.
Unlike many herbivores, caterpillars have the task of ingesting such vast amounts
that the gut content may be up to twice the weight of its own tissues. An alimentary
throughput time of just hours means there is virtually no role for symbiotic micro-
organisms to aid digestion. And the frenzy of feeding involves big doses of potential
toxins. The first parts of this book address some of these problems and the remark-
able ways that caterpillars have dealt with them.
Finally, there are gambles associated with availability of hosts; variation in qual-
ity of the host, genetically or as a result of the herbivore damage; the sensing of
nearby herbivore damage; or the acquisition of a plant disease.
Presented here are updates about impact of the natural enemies and diseases on
the ecology and evolution of caterpillars and the multiple interactive effects among
the three trophic levels—plants, herbivores, and herbivore enemies. The exceptional
diversity of anatomical, physiological, and behavioral traits to deal with the number
and variety of problems provides a classic arena for the study of evolution.
Research on caterpillar behavior and ecology continues to be important in natu-
ral ecosystems for understanding the impact of climate change and loss of habitat
on all three trophic levels, and the complexity of all those tangled webs. How will
herbivorous Lepidoptera be affected by changing distribution of hostplants in rela-
tion to changes in climate? Can citizen scientists provide the quantity of information
to monitor changes in each of the three trophic levels around the world?
Foreword ix
Continued study of caterpillars is also needed to understand best practices for
wild lands and forest management, and to handle the devastating impacts of both
invasive species and native outbreaking species. The combined research from all
these interlocking research elements, most of them addressed in this book, also aim
to elucidate the essential balance that is ultimately important for vertebrates includ-
ing humans.
Regents Professor Emerita Elizabeth Bernays
University of Arizona,
Tucson, AZ, USA
Acknowledgments
We thank the following persons for reviewing chapters. These reviewers made sub-
stantial contributions to the quality of the material contained within this book
through their careful reading and resulting constructive criticism and suggestions:
Annette Aiello, Inge Armbrecht, Christina Baer, Alex Baranowski, Karina Boege,
Deane Bowers, Laura Braga, Sue Carnahan, Phyllis Coley, Rex Cocroft, Tatiana
Cornelissen, Jaret Daniels, Ek del-Val, David Dussourd, Lee Dyer, Marianne
Espeland, James Fordyce, Jessica Forrest, Andres Freitas, Rieta Gols, David Heckel,
Owen Lewis, John Lill, Eric LoPresti, Tanner Matson, Kailen Mooney, Xoaquin
Moreira, Douglas Morse, Paul Ode, Paulo Oliveira, Steve Passoa, Moria Robinson,
Jay Rosenheim, Carlo Seifert, Tim Schowalter, Mike Singer, Angela Smilanich,
John Stireman, Doug Tallamy, Heiko Vogel, Martin Volf, David Webb, Martha
Weiss, James Whitfield, and Meron Zalucki.
In addition, we would like to express our appreciation to the Entomological
Society of America for the initial opportunity to bring many of the authors to St.
Louis in November 2019 to contribute to the symposium that led to the initial pro-
posal for this book. Finally, we appreciate the enthusiasm and support from our
colleagues, employees, and students at Florida International University and the
University of Missouri-St. Louis during the completion of this volume.
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