Sweating Cicadas
CICADAS are famous for their “music”-or perhaps one should say they are notorious for it. Their singing has been compared to a chain saw driven at high speed against naked iron, to the frantic thumping of a tin can, and to shrilly, trilling flutes. A sing
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The Hot-Blooded Insects STRATEGIES AND ~ MECHANISMS OF -· THERMOREGULATION ~~~;~
BERND HEINRICH
SPRINGER-VERLAG BERLIN HEIDELBERG GMBH
Copyright © 1993 by Bernd Heinrich Originally published by Springer-Verlag Berlin Heidelberg in 1993 Softcover reprint of the hardcover 1st edition 1993 All rights reserved 10 9 8 7 6 5 4 3 2
ISBN 978-3-662-10342-5 ISBN 978-3-662-10340-1 (eBook) DOI 10.1007/978-3-662-10340-1
This book is printed on acid-free paper and its binding materials have been chosen for strength and durability. Designed by Gwen Frankfeldt
Contents
Prologue 1 Night-Flying Moths
17
2 Butterflies and Wings
76
3 Dragonflies Now and Then 4
117
Grasshoppers and Other Orthoptera
5 Beetles Large and Small
191
6 Bumblebees Out in the Cold 7 Tropical Bees
227
277
8 Hot -Headed Honeybees 9 The Tolerance of Ants
292 323
10 Wasps and the Heat of Battle 11
Flies of All Kinds
343
12
Sweating Cicadas
369
334
13 Warm Caterpillars and Hot Maggots 14 Fever 411 15
Cold Jumpers
422
16 Social Thermoregulation Summary References
143
510 525
Acknowledgments 585 Index of Authors Cited General Index 598
590
447
382
Prologue
N
o aspect of the physical environment is more important to insects than temperature. In most environments temperature fluctuates through time, but insects also experience extreme temperature variations in space. A large mass, such as a human body weighing 65 kg, would register no measurable temperature increase by stepping from shade to sunshine for several minutes; a 10 mg fly, however, heats up some 10 0 C in only 10 seconds when it lands in a sunfleck. Needless to say, the thermal environment faced by insects is potentially much more severe than it is to us or to any other vertebrate animal. And it is probably not an exaggeration to claim that insects have evolved some of the most amazing feats of thermal adaptation and thermoregulation in the entire animal kingdom. Yet, little over 20 years ago, that statement would have seemed eccentric. As we sat one night in 1969 in front of a white sheet illuminated by a lantern on Mount Kainde in New Guinea among dark forest trees festooned with orchids, moss, and tangles of lianas, one after another beautiful moth, of dozens of different species, flew out of the darkness. I captured the arriving insects by net and thrust a thermocouple probe into each one to measure its body temperature. My host, Peter Shanahan, owner of the coffee plantation and of the white sheet, kindly tolerated my unusual routine. I was grasping for any and all clues that might ultimately allow me to figure out what to me seemed a deeply puzzling question: How could moths, who have no sweat glands, no lungs, no capillary system for peripheral blood circulation, and no major muscles besides those used for flight-how could such creatures possibly regulate their body temperature and fly at the same time?
After two years of grappling with the problem for my Ph.D. thesis under George A. Bartholomew and Franz Engelmann at UCLA, I had only recently become convi
