The specialized relationship between an insect pollinator and its flower can be far more extreme than merely having correspondingly long tongues and nectaries. Some insects have coevolved with their floral hosts, the two becoming so specialized for one another that when one diverges into two new species, so too does the other. Textbook examples of coevolving pollinators are fig wasps and yucca moths, and the figs and yucca plants for which each is named. Figs are critical sources of food, and in some forests they can comprise up to 70 percent of the diet for the associated animals, be they birds, monkeys, or even humans. An immature fig fruit has a small opening that serves as the tunnel through which a mated female fig wasp, who is often scarcely larger than the head of pin, will crawl. The space is narrow and her body is slender, frequently with a rather peculiarly flattened and elongate head. Despite her adaptive size, she will usually have her wings torn from her sides as she attempts to make her way inside the fruit. Once inside, she lays her eggs and simultaneously deposits the pollen she had picked up from the original fig plant where she was born. The fig’s flowers are minute and line the inner chamber within the immature fruit. Trapped within the fruit, the fig wasp mother dies. Her offspring emerge as larvae and are provided safety and nourishment by the fig fruit, which, now pollinated, matures. After pupating, male fig wasps, many of which are neotenic (see page 90) and therefore flightless and more worm-like than recognizable as wasps—mate with the emerging females. As his final act, a male will bore a hole through the mature fig fruit, eventually reaching the outside, where he will die. The new females use the tunnel dug by the male as a means of escape, picking up pollen as they go and then departing to perpetuate the cycle.
Impressive pollinators such as the giant sphinx moth (Cocytius antaeus), with a wingspan up to 7 inches (17.8 centimeters), have elongate proboscides to enter the deep tubular flowers which they pollinate, including the rare ghost orchid (Epipogium aphyllum). From Maria Sibylla Merian, Over de voortteeling wonderbaerlyke veranderingen der Surinaemsche insecten (1719 Dutch edition of Metamorphosis Insectorum Surinamensium, 1705).
The intimate association of insects and flowers is a common theme in illuminated entomological texts, as exemplified here in the finely colored title page from the third volume of August Johann Rösel von Rosenhof’s Der Insecten-Belustigung (1746–1761).
Figs and fig wasps have been at this for about seventy million years, and we can find fossils of fig wasps, complete with the fig pollen they are carrying, preserved with lifelike fidelity in ancient amber. The fig–fig wasp mutualism is complex and varied across the hundreds of species involved, but what is perhaps truly remarkable is that we seem to have had a rudimentary appreciation of this method of pollination for thousands of years. The ancient Greek writer Herodotus (ca. 484 – 425 BCE) reported in his book, The Histories, how the Babylonians, who cultivated figs, understood that the fruits would not ripen unless they were first entered by a miniscule “fly” and that mature figs had the insects within them. The wasps were so small that in the absence of optics with which to magnify them, it is easy to see how they might be confused as gnats or other flies. While they could not have imagined the real complexity of the relationship or the underlying mechanism of what was going on, the growers of Babylon were keen observers and knew that ripened fruits were only achieved by the entry of the insect.
Yucca moths are the obligate pollinators of yuccas, as well as herbivores that partially devour them. Pollinating yucca moths are perhaps the only example of a pollinator in which the animal vector intentionally, rather than inadvertently, pollinates the plant. The adult moth has modifications of its proboscis that permits it to collect yucca pollen. The moth then bores a hole into the flower’s ovary and stuffs the packet of pollen into the stigma, thereby pollinating the plant. She will also deposit her eggs within. The moth larvae feed exclusively on yucca seeds; on cursory inspection this would appear to be disadvantageous to the plant, as the latter obviously needs the seeds in order to reproduce. The larvae, however, feed only on a sufficient number of seeds to complete their development, but never all of them, leaving behind a shared portion for the plant. In this way, the moth and yucca share in the harvest, and neither can survive without the other.
We have long held flowers in high esteem, and gardens, floral folios, and flower societies attest to this adoration. Many of the illuminated entomological texts of past centuries revolved around this fondness, showcasing the largest and most spectacular of insects in association with flowers, ranging from Maria Sibylla Merian’s engravings of metamorphosis (see pages 94-96) to Edward Donovan’s butterflies of India and China (see pages 188-189). Libraries and galleries are filled with such art, the peaceful joining of insects and their pollinators. If you will, pollination represents a form of détente in the otherwise ancient battle between plants and their dominant herbivores—insects. Through pollination, plants and insects become collaborators rather than combatants, and as this relationship blossomed, so too did our world.
There remains much to discover about the intimate entanglement of flowers and insects. While certain aspects of such discovery may appear esoteric—of interest to no one other than singularly obsessed horticulturists or bespectacled entomologists—they all have far-reaching ramifications. Our very lives may depend on discovering, learning about, and conserving insects as “invisible” as thrips or as conspicuous as gently humming bumble bees. The avenues of discovery are as innumerable as insects themselves. Even the seemingly common and well-trodden faunas of our own backyards are filled with novel and important discoveries to be made, from new species to new revelations into insect songs, dances, rituals, and rites. Insects are significant, and it is only appropriate that some of us devote our lives to more fully understanding theirs.
The number of biodiversity entomologists who undertake this work is few, and the task before us great. Many of the spellbinding discoveries and scientific artwork of the past were expertly executed by amateurs, the “citizen scientists” of their day—informed clergy, passionate medical doctors, artists, and explorers. The awe-inspiring achievements of insects, past and present, readily enliven a passion for engaged research, and such study is not the privileged reserve of only a few in lofty towers of ivory.
Insects are not merely innumerable, for their diversity is disproportionately large when compared to those other lineages of life around us. As J. B. S. Haldane (see page xiii) rightly intimated to his august companion the archbishop of Canterbury, insects are inordinate—inordinate in the legions of individuals that surround us, inordinate in the endless variety of species, and inordinate in the myriad manners by which they live out their lives and underpin, or undermine, those of other organisms, ourselves most of all. Inordinate, however, does not mean incomprehensible, intractable, or impossible. Inordinate instead should inspire, invigorate, enthrall, enliven, and call for eager engagement. We can all be entomologists, for in our capacities to learn and create we too are inordinate.
The resplendently ornate title page to Robert H. F. Rippon’s Icones Ornithopterorum (1898–[1907]), a three-volume work that covered everything known of the great birdwing butterflies and their relatives. Rippon, a student of the great polyhistor John O. Westwood, provided all of his own illustrations, including this title page.
“There is grandeur in this view of life, with its several powers, having been originally breathed by the Creator into a few forms or into one; and that, whilst this planet has gone cycling on according to the fixed law of gravity, from so simple a beginning endless forms most beautiful and most wonderful have been, and are being, evolved.”
—Charles R. Darwin On the Origin of Species, 1859
The gilt cover of E. F. Stavely’s British Insects (1871).
ACKNOWLEDGMENTS
For Mnemosyne: a Bee, a Bloom, a Breeze
A work such as this attempts to thread a fine needle, representing two seemingly different books interwoven into one. On the one hand it tries to provide a narr
ative of insect diversity and evolution, an evolutionary history of 400 million years on six legs. On the other, it chronicles, albeit with great lacunae included, humankind’s past exploration of insects, highlighting some of the now rare tomes that represent both artistic as well as scientific achievements. A wiser author would see these as disparate and unmixable tales, and the reader is left to judge the mongrel hybrid produced of this forced union.
Although it may be my words that appear on the pages bound herein, a work such as this is truly a collaborative endeavor, both directly and indirectly. At the American Museum of Natural History, Tom Baione, Harold Boeschenstein Director, Department of Library Services, was a cheerful and wealthy store of wisdom and assistance throughout. He and Mai Reitmeyer, senior research services librarian, were as patient as Job while I feverishly poured over the fine works under their care, and they tolerated my many large requests. I am further grateful to Tom for composing the beautiful foreword that graces the start of this volume. David Grimaldi, curator and professor, Division of Invertebrate Zoology, generously suggested I should make this undertaking, and for the joyful labor I cannot be too grateful. Dave, Tom, Mai, and Valerie Krishna, professor emerita of English at the City College of New York, offered many constructive suggestions, and any lapses that remain are my fault alone. They and my other dear friends at the Museum have represented a tremendous source of support, devoting time and energy beyond measure. Like Aeneas recalling the siege of Troy, they may each claim “quorum pars magna fui”! Although no longer with us, distinguished entomologists Kumar Krishna and Charles Michener seemed to have been with me throughout, and fond memories of either pouring over rare volumes with each or visiting antiquarian book stores in London with the former and Valerie dispelled frustrations whenever they arose.
Also of particular note at the American Museum of Natural History in the office of Global Business Development are Sharon Stulberg, senior director; Elizabeth Hormann, former assistant director; Joanna Hostert, marketing manager, and Courtney Edwards, business manager, with additional support from Jill Hamilton. Roderick Mickens, senior photographer in the Museum’s photography studio, dedicated countless hours to the photography for this book with assistance from Barbara Rhodes, conservation librarian. At Sterling Publishing, I am grateful to Barbara M. Berger, executive editor, for encouragement, listening to my entomological musings, and forgiving those unexpected challenges that stumble any writer. Also at Sterling I would like to thank Scott Russo, associate art director, for his stunning cover and his interior art direction; as well as Jo Obarowski, creative director; and Ellen Hudson, production manager. At Tandem Books, special thanks to Ashley Prine for the beautiful interior design and to Katherine Furman for her proficient copyediting. For their tolerance of my absences while I hid myself away to read and write, I am thankful to my students and colleagues at the University of Kansas, Zachary H. Falin, senior collection manager, Division of Entomology, Biodiversity Institute; Jennifer C. Thomas, associate collection manager, Division of Entomology, Biodiversity Institute; and Victor H. Gonzalez, director, Human Anatomy Laboratories, Undergraduate Biology Program, all of whom leapt to pick up matters when I was otherwise occupied.
I am grateful to the many writers and artists who preceded my own entomology, and left behind such grand and sumptuously illustrated works. They universally inspired and entertained, and left me awestruck over their genius, skill, passion, and bravery, all in the name of the mighty majority. If it were not for their many labors, there would have been no tales to tell.
Lastly, no prose is sufficient to thank my family for their forbearance, confidence, and love. Supporting my obsessions and frequent absenteeism have been my parents, A. Gayle and Donna Engel, without whom none of this would have been possible, quite literally. My siblings, Elisabeth and Jeffrey, have endured my prattling on over dusty books and all things “buggy” while my nieces and nephews—Grace, Kate, Leo, and Isaac—and extended family have been a constant source of joy, rejuvenating the soul on weary days. Foremost of all, I am thankful to my wife, Kellie. During late nights and long days she assisted in tracking down obscure historical sources, read and edited text, and kept spirits high when faculties and energy were stretched to their most thin. It is fair to say that without her constant faith and aid, my efforts would have been in vain. To her, and my whole family, I am most deeply appreciative, and before them I am always humbled.
SUGGESTED READING
Buchmann, Stephen L., and Gary P. Nabhan. The Forgotten Pollinators. Washington, DC: Island Press, 1996.
Dethier, Vincent G. Crickets and Katydids, Concerts and Solos. Cambridge, MA: Harvard University Press, 1992.
Eisner, Thomas. For Love of Insects. Cambridge, MA: Belknap Press, 2003.
Grimaldi, David, and Michael S. Engel. Evolution of the Insects. Cambridge, UK: Cambridge University Press, 2005.
Hoyt, Erich, and Ted Schultz. Insect Lives: Stories of Mystery and Romance from a Hidden World. New York: John Wiley & Sons, 1999.
Marshall, Stephen A. Insects: Their Natural History and Diversity—With a Photographic Guide to Insects of Eastern North America. Richmond Hill, ON: Firefly Books, 2006.
Seeley, Thomas D. Following the Wild Bees: The Craft and Science of Bee Hunting. Princeton, NJ: Princeton University Press, 2016.
Shaw, Scott R. Planet of the Bugs: Evolution and the Rise of Insects. Chicago: University of Chicago Press, 2014.
Wilson, Edward O. The Diversity of Life. Cambridge, MA: Belknap Press, 1994.
Zinsser, H. Rats, Lice and History: A Chronicle of Pestilence and Plagues. Boston: Little, Brown, 1935.
WORKS FEATURED
Aldrovandi, Ulisse. De Animalibus Insectis: Libri Septem cum Singulorum Iconibus ad Vivum Expressis. Bologna: Apud Clementem Ferronium, 1638 (1602).
Audouin, Jean Victor. Histoire naturelle des insectes, traitant de leur organisation et de leurs moeurs en general. Paris: F. D. Pillot, 1834.
Bates, Henry W. “Contributions to an Insect Fauna of the Amazon Valley. Lepidoptera: Heliconidae.” Transactions of the Linnean Society of London, vol. 23. London: Taylor and Francis, 1862 (1791–1875).
Biologia Centrali-Americana. Insecta. Coleoptera. London: Published for the editors by R. H. Porter, 1880–1911.
Biologia Centrali-Americana. Insecta. Diptera. London: Published for the editors by R. H. Porter, 1886–1903.
Biologia Centrali-Americana. Insecta. Lepidoptera-Heterocera […] London: Published for the editors by R. H. Porter, 1881–1900.
Biologia Centrali-Americana. Insecta. Neuroptera. Ephemeridae. London: Published for the editors by Dulau, 1892–1908.
Biologia Centrali-Americana. Insecta. Orthoptera. London: Published for the editors by R. H. Porter, 1893–1909.
Biologia Centrali-Americana. Insecta. Rhynchota. Hemiptera-Homoptera. London: Published for the editors by Dulau, 1881–1909.
Butler, Charles. The Feminine Monarchie, or the Historie of Bees. Shewing Their Admirable Nature, and Properties; Their Generation, and Colonies, Their Government, Loyaltie, Art, Industrie, Enimies, Warres, Magnanimitie, &c. Together with the Right Ordering of Them from Time to Time: and the Sweet Profit Arising Thereof. Oxford: Printed by William Turner, for the author, 1634 (1609).
Curtis, John. British Entomology; Being Illustrations and Descriptions of the Genera of Insects Found in Great Britain and Ireland: Containing Coloured Figures from Nature of the Most Rare and Beautiful Species, and in Many Instances of the Plants upon Which They Are Found. London: Printed for the author and sold by E. Ellis, 1823–1840.
Cuvier, Georges. Le règne animal distribué d’après son organisation: pour servir de base à l’histoire naturelle des animaux et d’introduction à l’anatomie comparée. Paris: Fortin, Masson et cie, 1836–1849.
Darwin, Charles. The Various Contrivances by Which Orchids Are Fertilised by Insects. New York: D. Appleton, 1895 (1862).
Denny, Henry. Monographia Anoplurorum Britanniae; or, An Essay on the British Species of
Parasitic Insects Belonging to the Order of Anoplura of Leach, with the Modern Divisions of the Genera According to the Views of Leach, Nitzsch, and Burmeister, with Highly Magnified Figures of Each Species. London: H. G. Bohn, 1842.
Donavan, Edward. Natural History of the Insects of China. London: R. Havell and H. G. Bohn, 1838.
———. Natural History of the Insects of India. London: R. Havell and H. G. Bohn, 1838.
Drury, Dru. Illustrations of Exotic Entomology, Containing Upwards of Six Hundred and Fifty Figures and Descriptions of Foreign Insects, Interspersed with Remarks and Reflections on Their Nature and Properties. London: H. G. Bohn, 1837.
Dumont d’Urville, Jules-Sébastien-César. Voyage au pôle Sud et et dans l’Océanie sur les corvettes l’Astrolabe et la Zélée, exécuté par ordre du roi pendant les années 1837–1838–1839–1840, sous le commandement de m. J. Dumont d’Urville, capitaine de vaisseau, publié par ordonnance de Sa Majesté sous la direction supérieure de m. Jacquinot, capitaine de vaisseau, commandant de la Zélée […] Paris: Gide, 1842–1854.
Ehrenberg, Christian Gottfried. Symbolae Physicae, seu, Icones et Descriptiones Corporum Naturalium Novorum aut Minus Cognitorum, Quae ex Itineribus per Libyam, Aegyptum, Nubiam, Dongalam, Syriam, Arabiam et Habessiniam […] Berlin: Mittlero, 1828–1845.
Forel, Auguste. Histoire physique, naturelle et politique de Madagascar, Hymenoptères. Les Formicides. Paris: Imprimerie nationale, 1891.
Forsskål, Peter. Descriptiones Animalium, Avium, Amphibiorum, Piscium, Insectorum, Vermium; Quae in Itinere Orientali Observavit Petrus Forskål. Copenhagen: Mölleri, 1775.
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