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Academic & Professional Books  Insects & other Invertebrates  Insects  Bees, Ants & Wasps (Hymenoptera)

The Guests of Ants How Myrmecophiles Interact with Their Hosts

By: Bert Hölldobler(Author), Christina L Kwapich(Author)
559 pages, 205 colour & b/w photos, 16 colour & b/w illustrations
Publisher: Belknap Press
This unique and beautifully illustrated book takes a look at one of the most delightful aspects of ant biology: the many organisms that live amongst them.
The Guests of Ants
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  • The Guests of Ants ISBN: 9780674265516 Hardback Jul 2022 In stock
Price: £59.95
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About this book

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
    Nestmate Recognition
    Identity Theft and Other Means of Intrusion
    Cuticular Hydrocarbons Revisited
4. The Lycaenidae: Mutualists, Predators, and Parasites
    The Mutualists
    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
    Collective Mimicry
    Backpacks, Shields, and False Heads
    Locomotory Mimicry
    Odor Cloaks
    Phoretic Spiders Take Flight
    Spider Predators of Veromessor pergandei
7. The Mysteries of Myrmecophilous Crickets
    Strigilators and Thieves
    Specialized Mouthparts
    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

Customer Reviews (1)

  • A critical review that is exquisitely presented and thoroughly engrossing
    By Leon (NHBS Catalogue Editor) 19 Dec 2022 Written for Hardback

    It has to be one of the more delightful details of the natural world: the ecosystem of an ant's nest is home to its own constellation of creatures that specialise in living within or nearby it. Daniel Kronauer's book Army Ants first drew my attention to these so-called myrmecophiles and their sometimes bizarre adaptations. I was stoked when Harvard University Press announced it would publish a monograph focusing on just this aspect of ant biology, authored by entomology professors Bert Hölldobler (a frequent co-author to E.O. Wilson) and Christina L. Kwapich. The Guests of Ants gives a beautifully illustrated, wide-ranging, and critical literature review of this delightful corner of myrmecology. Will ants make it to my personal top 5 for a third-year running? This book is a very strong contender.

    Before delving in, a quick word on what is not in the book. Excluded are the large topic of interactions between ants and plants, the subject of trophobiosis (i.e. the bugs and some butterflies that provide honeydew to ants and in return are protected by them), and the hundreds of socially parasitic ant species that exploit other ants. This book is also not an exhaustive listing of all known myrmecophiles, as done in review papers and books such as Parasites in Social Insects and The Ants. Instead, Hölldobler & Kwapich use numerous examples to review the behavioural mechanisms by which myrmecophiles coexist and often exploit their hosts. How do they break the internal communication codes that otherwise make ants such a successful superorganism? This self-imposed limitation nevertheless leaves plenty of material and the first thing that will strike you is the book's sheer size. Measuring some 24 × 24.5 cm it is as big as Army Ants, though it is a good deal thicker. What it shares is that it is chock-a-block with glorious macro-photography. The acknowledgements thank no fewer than 84 people for providing (mostly) photos and illustrations and the book is a feast for the eyes.

    So who are these guests? Beyond two chapters on the really small (bacteria, internal parasites, and fungi such as crowd-pleaser Ophiocordyceps) and the really large (vertebrates including birds, lizards, and amphibians), most myrmecophiles are arthropods. Some flies and beetles rely on chemical mimicry to blend in and e.g. hitch a ride on ant bodies, lay their eggs inside the protected environment of ant nests, or forage here for detritus and ant corpses. Caterpillars of lycaenid butterflies possess glands that secrete so-called appeasement substances, allowing them to live amidst ants as mutualists, parasites, or even predators. Refuse sites inside the nest, and foraging trails outside it, host numerous beetles and flies that steal booty and prey the ants bring home, or attack the ants directly. Crickets behave as thieves, technically as kleptoparasites. Using a mixture of chemical stimuli, such as odours, and tactile stimuli, such as antennae and mouthparts with which they tickle ants, they either get ants to regurgitate liquid food for them (as seen on the book's cover) or get in between two ants practicising what biologists call trophallaxis, the transfer of food between each other. But the most mind-blowing chapter showcases the unbelievable morphological and behavioural mimicry of certain spiders. Some spiders, as they grow larger, resemble different ant species in turn. Other spiders use parts of their body such as chelicerae (mouth parts) and fold these in front of their heads to resemble an ant carrying a nestmate.

    The above is just a small selection of a mind-bogglingly diverse cast of myrmecophiles. Two further chapters focus more on underlying explanations. One looks at a possible evolutionary pathway that explains how free-living insects became fully integrated into ant nests, illustrating it through studies on different rove beetles. The other chapter looks at various ant ecosystems and their guests, including harvest ants, wood ants, and army ants. It also discusses studies that have tried to apply network analyses to document foodweb interactions (i.e. myrmecophiles eating ants and each other), and studies that have looked at links between colony traits, such as genetic diversity, and rates of myrmecophile infestation.

    A few things struck me in particular about this book. First is the amount of detail. The authors thoroughly show just how diverse are myrmecophiles and the ways in which they trick ants. If I have to sound a critical note, it is that, though the individual chapters are organised around themes, I was missing a final chapter to tie it together and provide some sort of overview or tabulation. The bewildering diversity, fascinating as it is, now remains just that: bewildering. But perhaps that is the nature of the beast. Second was a fascinating lesson on cuticular hydrocarbons. These are carbon-and-hydrogen-containing chemicals that insects sport on their outer layer, their cuticle, as a kind of chemical signature. You would think that mimicking or acquiring cuticular hydrocarbons is key for myrmecophiles to be accepted or ignored by ants, rather than attacked and killed. But numerous studies have thrown up conflicting results: "we have to guard against a hasty conclusion that the hydrocarbon match represents the effective mechanism for the acceptance of myrmecophiles by their hosts" (p. 147). Other chemicals as well as behaviour seem to play an overriding role in some species. Third and final was a welcome critical attitude from the authors in their review. They frequently point out shortcomings in experimental protocols, comment on the appropriateness of certain methods, and tone down sweeping conclusions. None of this is mean-spirited but is done to encourage rigorous experimental protocols. The ecosystem of ants and their guests is incredibly complex and studying them is hard; drawing firm conclusions requires excluding alternative explanations.

    The Guests of Ants is a must for myrmecologists and entomologists, but also more generally for biologists with an interest in social insects and insect ecology. The book provides a welcome entry point into a remarkably rich literature (45 pages of references are provided). The work is detailed and fairly, but not overtly, technical; effectively this is at the level of a series of literature reviews as one might find in e.g. Trends in Ecology & Evolution. There is nothing here that amateur entomologists or an interested lay audience could not handle, with a 14-page glossary taking care of technical terms. For my part, I have to congratulate the publisher for producing yet another exquisitely presented, in-depth monograph on ants that highlights a thoroughly engrossing aspect of these fascinating insects.
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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.

By: Bert Hölldobler(Author), Christina L Kwapich(Author)
559 pages, 205 colour & b/w photos, 16 colour & b/w illustrations
Publisher: Belknap Press
This unique and beautifully illustrated book takes a look at one of the most delightful aspects of ant biology: the many organisms that live amongst them.
Media reviews

"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

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