A fascinating examination of socially parasitic invaders, from butterflies to bacteria, that survive and thrive by exploiting the communication systems of ant colonies.
Down below, on sidewalks, in fallen leaves, and across the forest floor, a covert invasion is taking place. Ant colonies, revered and studied for their complex collective behaviours, are being infiltrated by tiny organisms called myrmecophiles. Using incredibly sophisticated tactics, various species of butterflies, beetles, crickets, spiders, fungi, and bacteria insert themselves into ant colonies and decode the colonies' communication system. Once able to "speak the language," these outsiders can masquerade as ants. Suddenly colony members can no longer distinguish friend from foe.
Pulitzer Prize-winning author and biologist Bert Hölldobler and behavioural ecologist Christina L. Kwapich explore this remarkable phenomenon, showing how myrmecophiles manage their feat of code-breaking and go on to exploit colony resources. Some myrmecophiles slip themselves into their hosts' food sharing system, stealing liquid nutrition normally exchanged between ant nestmates. Other intruders use specialized organs and glandular secretions to entice ants or calm their aggression. Guiding readers through key experiments and observations, Hölldobler and Kwapich reveal a universe of behavioural mechanisms by which myrmecophiles turn ants into unwilling servants.
As The Guests of Ants makes clear, symbiosis in ant societies can sometimes be mutualistic, but, in most cases, these foreign intruders exhibit amazingly diverse modes of parasitism. Like other unwelcome guests, many of these myrmecophiles both disrupt and depend on their host, making for an uneasy coexistence that nonetheless plays an important role in the balance of nature.
Preface [Bert Hölldobler]
1. Superorganisms: A Primer
Mating and Colony Founding
Division of Labor among Worker Ants
Division of Labor in Reproduction
Parasites inside the Superorganism
2. Inside and on the Bodies of the Ants
Mutualistic Symbionts: The Case of Blochmannia
Internal Parasites and Parasitoids that Affect the Ants’ Behavior
Phoretic Myrmecophiles and Parasites on the Bodies of Host Ants
3. Recognition, Identity Theft, and Camouflage
Identity Theft and Other Means of Intrusion
Cuticular Hydrocarbons Revisited
4. The Lycaenidae: Mutualists, Predators, and Parasites
Predators and Socially Parasitic Myrmecophiles in the Genus Phengaris
Miletine Predators, Tripartite Symbiosis, and Indirect Parasitism
Parasitoids Attacking Lycaenid Myrmecophiles
5. Foraging Paths and Refuse Sites
Ant-Mugging Flies on the Ants’ Trails
Bengalia: Brood and Booty Snatcher Flies
The Beetle Amphotis marginata: Highwayman of Lasius fuliginosus
Predators and Scavengers: The Story of Some Pella Species
On the Ants’ Trails
6. Spiders and Other Mimics, Pretenders, and Predators
Transformational and Compound Mimics
Backpacks, Shields, and False Heads
Phoretic Spiders Take Flight
Spider Predators of Veromessor pergandei
7. The Mysteries of Myrmecophilous Crickets
Strigilators and Thieves
Host-Specialists and Host-Generalists
Odor Mimicry and Trail Following
The Mystery of Cricket Body Size
Phenotypic Plasticity and Cryptic Speciation
Gaster Mimicry and Egg Mimicry
Island Endemics and Island Hoppers
8. Grades of Myrmecophilous Adaptations
Pella humeralis: The Predator and Scavenger
Dinarda Beetles: The Sneaky Thieves
Breaking the Code: Lomechusa and Lomechusoides
Claviger: Pretending to Be a Piece of Booty?
Paussinae: The Myrmecophilous Dracula Beetles
9. Myrmecophiles in the Ecosystem of Ant Nests
The Nest Ecosystem of the Harvester Ant Pogonomyrmex badius
The Role of Myrmecophiles in the Ecosystem of Mound-Building Wood Ants
The Case of Army Ants, Especially the Leptogenys “Army Ants”
Networks and Colony-Level Censuses
Infestation by Myrmecophilous Parasites and Colony Traits
10. Vertebrates and Ants
Myrmecophiles of a Feather
Fish and Amphibious Associates
Reptile Eggs in Ant Nests
Horned Lizards, Blind Snakes, and Legless Lizards
Bert Hölldobler is the Robert A. Johnson Professor in Social Insect Research at Arizona State University. He was previously Professor of Biology and Alexander Agassiz Professor of Zoology at Harvard University and subsequently held the chair for Behavioral Physiology and Sociobiology at the University of Würzburg, Germany. He is an elected member of many academies, including the American Academy of Arts and Sciences, the U.S. National Academy of Sciences, and the German National Academy of Sciences, Leopoldina. He has received many awards, among them the Pulitzer Prize for The Ants, co-authored with E. O. Wilson.
Christina L. Kwapich is Assistant Professor of Biology at the University of Massachusetts, Lowell, and President-elect of the North American Section of the International Union for the Study of Social Insects.
"This beautiful book is a true classic of biology and destined to be a standard work on the subject of symbiosis for many years to come."
– Edward O. Wilson, author of Sociobiology
"A marvelous tour of the wonderland of ants and the non-ant 'guests' that live with them. Bert Hölldobler and Christina Kwapich have combined fascinating science and first-rate scholarship to share what myrmecologists have learned about a rarely seen part of the natural world."
– Thomas D. Seeley, author of The Lives of Bees
"Prepare to be astounded, surprised, and charmed time and time again. This book is destined to become the authoritative work on ants and their guests for decades to come."
– Walter R. Tschinkel, author of Ant Architecture
"A deeply inspiring and masterful account of how myrmecophiles interact with their hosts, trick them by breaking their communication codes, and make their living inside the ant colonies. Lavishly illustrated, this book is a thorough and fascinating study."
– Rüdiger Wehner, Professor and Director Emeritus, Institute of Zoology, University of Zürich
"Hölldobler and Kwapich have masterfully marshalled a vast literature on the evolution and behavioral ecology of ants and their many associates. This book will have wide appeal to ecologists and evolutionary biologists, as well as anyone interested in the natural history of social insects and their friends and foes."
– Naomi E. Pierce, Hessel Professor of Biology and Curator of Lepidoptera, Museum of Comparative Zoology, Harvard University
"A spellbinding journey to the exotic world of ants and their 'guests.' By far the most comprehensive treatment of this fascinating subject, this book is a delight."
– Daniel Kronauer, author of Army Ants
"Superbly illustrated and written with enthusiasm and delight, The Guests of Ants is unparalleled. This book provides first-class, fascinating coverage on the behavioral biology and mechanisms enabling myrmecophiles to coexist with, and frequently exploit, the ant superorganism."
– Paulo S. Oliveira, coauthor of The Ecology and Evolution of Ant–Plant Interactions
"This magnificent book takes the reader into a striking world largely unknown, even to most biologists. Comprised of lively, informative writing spiced with fascinating illustrations, The Guests of Ants highlights some of the most bizarre and unbelievable organisms on earth. I would have loved to have had such a book when I started my own research years ago."
– Konrad Fiedler, University of Vienna, Austria
"An outstanding celebration of natural history as modern science. World-renowned scientist Bert Hölldobler and entomologist Christina Kwapich present a treasure trove of information about the ant nest microcosm, complete with spectacular photos of members on the colony's guest list, including butterflies, worms, fungi, and more."
– Raghavendra Gadagkar, author of The Social Biology of Ropalidia marginata
"This book provides a captivating, excellently illustrated overview of the complex interactions among ants and the many organisms who live inside their nests. A must-read for everyone who enjoys nature and the fascinating world of social insects."
– Jürgen Heinze, University of Regensburg