A little excursion through the world of books:
Edward Osborne Wilson, Jr., and Berthold K. Hölldobler,
Natural History of Social Insects and "The
([(This is not a real book-review but a summing up of a few books, an elaborated index of one of them and a few comments on that last book.)])
E. O. Wilson (born June 10, 1929) and B. Hölldobler (born June 25, 1936) belong to a very small group of leading man in myrmecology. They are all-rounders, studying mostly the behavior of ants but also systematics, morphology, physiology, … Other great man in the second half of the 21st century were Barry Bolton and William L. Brown, Jr., both systematicist, and, according to me, Heinrich Kutter and Alfred Buschinger, studying mostly the behavior of parasitic ants.
Wilsons highest award is the Crafoord Prize, 1990, a prize awarded by the Royal Swedish Academy of Sciences in certain sciences not covered by the Nobel Prize, and therefore considered the highest award given in the field of ecology. Also in 1990, Hölldobler received the Gottfried Wilhelm Leibniz Prize of the Deutsche Forschungsgemeinschaft, which is the highest honor awarded in German research.
Before we start with a little review of the latest book by Hölldobler and Wilson, lets have a little look at their most important books of the past. I will do this in chronological order because the information in them is supported by ongoing research.
first book in line is:
Wilson, E. O., 1971, The Insect Societies.
This was the first big review of social insects since William Morton Wheeler's book in 1928. It reviewed, in depth, the ants, eusocial bees and wasps and termites. It included systematics, ecology, behavior and physiology, all as far as was known at that time. The books last chapter expressed the hope that someday a new field of science would explain the social behavior of all animals. That field of science was born with:
Wilson, E. O., 1975, Sociobiology: The New Synthesis.
This monumental work became the cornerstone of the science that studies the social behavior of all animals. The first halve reviewed all of the theoretical background, the second halve all the social and subsocial animals (humans included!). This sociobiology is now on the foreground in all studies of social behavior. This field was worked out more to humans by Wilson and collaborators but, in the beginning, it met with great resistance from the Human Social Sciences departments around the world. Now, everybody finds that this coalescence of biology and the "Humanities" was good for the study of human nature.
Wilson, E. O., 1990, Excellence in Ecology, Vol. 2:
Success and Dominance in Ecosystems: The Case of the Social Insects.
This is an important book that explains in depth why social insects were so successful in geological history and so dominant in modern times. Wilson gave here, as far as I know, his first comparison of social insects with superorganisms (an idea first proposed by W. M. Wheeler in 1911.). It occupied a complete chapter in a marvelous book. The "little" book appeared, as an ECI Prize winner's book, in the same year as:
This Pulitzer Prize winning book
was the first big encyclopedic work about ants since 1910. It included reviews of every topic in myrmecology like systematic, life history, mating, … and about
army ants, leaf-cutting ants, parasitic ants, … Also in this massive tome were determination keys for the workers of recent ant genera (by Barry Bolton, who
elaborated them in two books later on.).
Hölldobler, B. & Wilson, E. O., 1990, The Ants.
book also included unpublished material. One of the observations by B. Hölldobler mentioned in this book, and kept in my memory (and I still hope to see a
publication describing it in every detail!), is about the resting/sleeping behavior of Camponotus, one of the big
ant genera in the world.
After completing this book, Wilson decided to write a book about one of his
Wilson, E. O., 1992, The Diversity of Life.
After his cry for help for the natural world in 1985, Wilson wanted to bring the diversity of life in the spotlight. Being one of the organizers of the two first conferences about that topic (and one of the editors of the proceedings of both.) he decided to write a book showing the general reader some of the greatest wonders of animals and plants and what we, humans, are doing to destroy them. This topic is pursued by him later on in a few other books and originated, according to me, in his love for nature, biophilia.
Hölldobler, B. & Wilson, E. O., 1994, Journey to the Ants:
A Story of Scientific Exploration.
This second book from Hölldobler and Wilson was intended to be a general introduction to the ants for the general public. It included also a few recent advances in myrmecology since the publication of their first book "The Ants". It was translated in 26 languages (if I remember correctly.) and is still considered as the best introduction to the world of the ants.
Hölldobler, B. & Wilson, E. O., 2008, The Superorganism.
The Beauty, Elegance, and Strangeness of Insect Societies.
This third and, for the moment, latest book of both scientists together is a good review of the concept of the superorganism. It is a worked out version of the chapter in Wilson's 1990 "little" book.
Although many scientists hoped to see a book that reviewed everything about social insects in general or about ants in particular this was not the case. Throughout the book it is mentioned a few times that this was not intended to be such an encyclopedic work but more a lavishly exemplified book about the key features of a superorganism (e.g. page XX "This book is not intended to be as comprehensive a monograph as The Ants (1990)" or page 9 "…, we will draw examples from …".).
Let us have a quick guided tour of the book. It has 10 chapters, preceded by a note and followed by an epilogue. Dr Ant noted "The chapters do not seem to flow into one another, but rather to be an assortment of things they have wanted to write about but haven't had a good chance to expound upon before.". This is not the case however. The first four chapters are more about the theoretical foundation (with examples.) while the rest covers the more important characteristics of superorganisms. In the chapters are a lot of examples that are mentioned in their previous books.
So, here comes the introduction:
Note to the General Reader.
Wouldn't it be nice to be an alien? The idea of an Encyclopedia Galactica
goes back (for me!) to Carl Sagan and looks great, at least on the scale of the world we live in. For an alien some forms of life will attract more attention
than others and yes, for me ants, wasps and bees deserve a big lot of that attention! The authors go back in the past before humans appeared and introduce the
aliens, looking at eusocial insects. And by looking at them, all the characteristics of a superorganism are summed up, to be reviewed in depth in the rest of
the book (with a lot of examples!). Also is explained why it is important to study social insects for sociology and biology in general!
Chapter 1:The Construction of a Superorganism.
this chapter Wilson and Hölldobler explain 1) why insect colonies are superior (very short version of the 1990 "little" book!), 2) how superorganism
are constructed and 3) the levels of organization in the biological world. It is concluded with a brief history of insect sociobiology.
Although it was intended as a general introduction for the rest of the book, it
also gives a detail that can't I find outside the original article: How do we get the estimates of biomass?
Chapter 2: Genetic Social Evolution.
is the genetical background of eusociality and how did that theory evolve through history. Haplodiploidy, Hamilton, Wilson, inclusive fitness, kin selection,
altruism, multilevel natural selection, … all pass the spotlight. The possible steps in the natural history of insects and their behavior, on the way form
solitary life styles to societies, are also mentioned and the "eusociality gene" and the "eusociality threshold" are introduced.
only problem with this chapter is on page 36, in the mathematical argument of Kern Reeve. In the quote a negative C is needed by unrelated individuals and
because my math is not so good, I can't follow everything in this quote. But for the rest, a very good historical review.
Chapter 3: Sociogenesis.
is the general life cycle of a colony and how is it generated? Here we meet algorithms, decision or epigenetic rules, decision points, behavioral programs and
how they work together to make a colony functions like it does! The insect as a cellular automaton! All by all, a colony is self-organized (major examples are
the swarm raids of army ants and the regulation of colony temperature in termites.).
reviewed are phylogenitic inertia and dynamic selection (e.g. chemical trails and mass communication in ants and, for honeybees, the organization of foraging
with waggle dance, shaking signal and tremble dance.).
Chapter 4: The Genetic Evolution of Decision Rules.
many decision rules are there? Of what kind are they? And what are the genetic changes behind them? Here the authors elaborate on these questions and bring on
sociogenetics and sociogenomics. A part of the chapter deals with honeybee sociogenomics and this includes their waggle dance, round dances, shaking dance,
tremble dance, pheromones during the dances, memory, … All the information for this complex system of foraging communication is stored in a part of the 10,157
genes identified in the honeybee.
next point in this chapter is sociogenomic conservation and "the large amount of change that can occur by the modification of a very few genes within the
genome." (p. 77.). The big example here is the social structure of the fire ants in North America.
little chapter concludes with a small piece about variation in genetic information and the resulting phenotypic plasticity.
Chapter 5: The Division of Labor.
Wilson and Hölldobler start this chapter with the definition of a superorganism
and a comparison between it and a "normal" organism (p. 84-85.). This is important because it determines all the contents of the following chapters
in the book (this one included!).
the three preceding chapters gave the genetic basis and how decisions are made, now we start with the most important external characteristics of a
chapter reviews the division of labor and castes. It includes reproductive division of labor (between queens and workers.), dominance orders, polyethism (or
temporal cast system, changing of work with increasing age, and it's physiological control/expression.) and polymorphism (different work for different
worker-morphologies, physical castes.).
polyethism counts that how older the worker, how farther away from the nest-centre it works (nursing/brood- and queen care, storing food and building the nest,
foraging and defense.).
authors also discuss the flexibility of these "systems", how workers (especially ant workers!) "find" their "labor-tasks" and the
genetic division/variability of work allocation (different matri-/patrilines.). Task switching and behavioral plasticity are important facets in evolution.
Child/larval work is also looked at.
way how cast are determined are also reviewed (genetic and nongenetic or environmental causes.).
chapter ends with adaptive demography (and the observation that specialized workers, although good in doing a specific job, don't do everything well) and
"small" mistake hit my eyes. On page 139 in note 121 they speak of 16 tribes but it must be 16 subfamilies!
Chapter 6: Communication.
do social insects communicate? Not only trail and alarm pheromones, recruitment signals, multicomponent signals, ritualizations, modulatory communication,
necrophoric behavior, nestmate recognition, … and how the chemical signals work are reviewed but also motor displays and visual, tactile and vibrational
communication. Some forms of communication can be classified in more than one type (e.g. tandem running: chemical and tactile signals/modulotory
communication.). A part of the chapter is also devoted to trophallaxis.
best studied examples of communication in social insects are foraging- and territorial communication in weaver ants, mass communication in fire ants and,
indeed, the dances of honeybees.
chapter ends with the communication of resource-holding potential (with the tournaments of Myrmecocystus as the
most important example.).
is the biggest chapter in the book but it gives all the variations very clearly. However, the problem with social insects is that almost all is
"said" by them with the help of chemical signals. Social insects are batteries of pheromones producing glands! If the authors wanted to write a
detailed review the book would be three or four times as thick as it is now.
of the more intriguing observations in the whole book is the claim by Z. Reznikova and B. Ryabko that wood ants can count and can pass that information on to
nestmates (p.256.). Strange indeed!!!
page 277 is stated "The postpharyngeal gland, a large organ located in the head and comprising two bilateral glove-shaped halves, is unique to
ants.". But a few days ago I found references to two articles, one by E. Strohm, G. Herzner and W. Goettler
"A "social" gland in a solitary wasp? The postpharyngeal gland of female European beewolves (Hymenoptera, Crabronidae)." (Arthropod
Structure & Development 36: 113-122.), the other by G. Herzner, W. Goettler, J. Kroiss, A. Purea, A. G. Webb, P. M. Jacob, W. Rössler and E.
Strohm "Males of a solitary wasp possess a postpharyngeal gland" (Arthropod Structure & Development 36,
123-133, both published in 2007.). In the abstracts I read "…and it was thought to be restricted to ants.", "…, we hypothesize that the PPGs of
ants and beewolves have a common evolutionary origin. Thus, our results suggest that the PPG in ants might not have evolved in response to social requirements
but might have already existed in solitary predecessors." and "We discuss the implications of our findings for the evolution of the postpharyngeal
gland in ants.". (The European beewolf is Philanthus triangulum.).
Chapter 7: The Rise of the Ants.
chapter reviews the history of the ants as seen in the fossil record. The main focus is a behavioral/ecological one and not a strict phylogenetic one (a tree
based on genetic evidence is only given as a frame to see the relations between the different ant groups!). The different radiations/extinctions of the ants
are reviewed. It also includes discussions of the ponerine paradox (the Ponerinae are a "primitive" but very successful ant group in the tropical and
warm temperate regions of the world), tropical ground faunas, arboreal groups and the dynastic-succession hypothesis.
small "omission" is Martialis but this ant was only discovered after Wilson and Hölldobler wrote the
book. But in this chapter it would not have been introduced at full length because nothing is known about its lifestyle.
I find two little hiccups:
The genera Archimyrmex, Polanskiella and Ameghinoia are synonyms (established in 2003.) but Wilson and Hölldobler use their names as separate groups (p. 319.).
page 322 it seems that they prefer the use of Ponerinae in the pre-2003 sense and only give Bolton's division of 2003 with some hesitation. Also, in this
chapter and the next, they use the "poneromorphs" in a lose sense, but with one big fault, namely "… that the assemblage as a whole represents a
diversification from a single Mesozoic ancestor." (p.322.). Had they only looked at the phylogenetic tree on page 316 …
Chapter 8: Ponerine Ants:
The Great Radiation.
Although the Ponerinae are morphologically "primitive", they show a big
variation in their social organizations, reproductive cycles, regulation of reproduction and division of labor. Here we get a review of the most important
discoveries of the last decades about facets of their life histories.
The review starts with the incredible sophisticated social organizations and reproductive strategies of Harpegnathos saltator (queens and/or gamergates, elaborate nests.) and Dinoponera quadriceps (only gamergates!). It continues with Gnamptogenys (queens and gamergates in different colonies.), Pachycondyla (whit all the diversity it exhibits, its sociobiological hyperdiversity!), Diacamma (with its mutilation-practice.), Streblognathus (dominance and fertility are uncoupled.), Platythyrea (ergatoid queens.) and Odontomachus (dominance hierarchies in polygynous colonies.).
most incredible part of the book is "Harpegnathos: Resilience in Reproductive Behavior". How infertile
workers, without queens or gamergates, lay haploid eggs that become males that inseminate their mothers that produce diploid eggs that become workers!!! Did
you follow it? Yes, amazing!
you really want to know, chapters 7 and 8 are not really necessary in this book. But they give so much information about variety of ecology/evolutionary
replacements and of sociobiological diversity in "primitive" ants that I find they are "alright" in this book.
little typo's or so in this chapter:
Figure 8-14, on page 374, line 3, word 10: at must be or (see explanation in the
Page 399, line 3: Plate 26 must be Plate 33.
Chapter 9: The Attine Leafcutters:
The Ultimate Superorganisms.
"Because they possess one of the most complex communication systems known in
animals, the most elaborate caste system, air-conditioned nest architecture, and populations into the millions, leafcutter ants deserve recognition as
Earth's ultimate superorganisms." (p. 408.).
Then, an explanation is given of why the attines deserve their status
as "ultimate" evolutionary endpoint in ants and why Atta, with
their big societies that inhabit huge nests with many fungus gardens, are the subject of the rest of the chapter.
Hölldobler and Wilson continue with a review of the life cycle, caste system,
foraging behavior, communication, nest structure, trails and trunk routes of Atta species, their symbiosis with
the fungus (including hygiene and waste management.) and the agropredators and agroparasites.
is a fairly complete review, even the relative nutritional values of different fungus-parts are mentioned.
the end of this chapter, Hölldobler and Wilson explain on which point they differ in opinion. Wilson wants to call all eusocial insect colonies superorganisms
(including e.g. the poneromorph and myrmeciine societies!) while Hölldobler want to restrict that "title" to eusocial insects with greatly reduced or
no reproductive competition. For convenience, all are included in the book.
final paragraph of the chapter is: "But whatever criteria may be adopted, there can be little doubt that the gigantic colonies of the Atta leafcutters, with their interlocking symbiont communities and extreme complexity and mechanisms of cohesiveness,
deserve special attention as the greatest superorganisms discovered to the present time." (p. 467.). Yes, indeed!
not so clear point is made on page 411 about "a single widespread and sexual fungal symbiont species". A better explanation was surely needed here!
little "mistake" also on page 411 about the lower attines: "… only minor polymorphism in minor worker size" must be "…only minor
polymorphism in worker size." And more: page 431, line 8, "(Figure 9-3)" must be "(Figure 9-4)"; page 436, reference 77, first
"Atta" must be deleted; page 439, line 5, "(see Plates 49 and 50)" must be "(see Plates
59 and 60)".
"big" mistake is present in the part "The Atta Caste System" and don't ask me how it got
there. To start, let us go back to chapter 5 "The Division of Labor", part "Adaptive Demography". The second paragraph (p. 153-154.) in
this part is about the adaption of worker size classes in developing Atta colonies. In it we read (p. 154, line
3.) "… becomes more sharply peaked in the smallest size classes." This is illustrated in figure 5-13 on page 153. In "The Atta Caste System" we get in paragraph 3 (p.426-427.) an almost identical review of this adaptive demography (same
writer who wanted to bring in a little variation?) and here we read (p. 427, lines 10-11.) "… becomes more sharply peaked and strongly skewed to the
larger-size classes." This is clearly wrong! The first one is the correct one! It must be the smaller workers that are more represented, like fig. 5-13
shows. This is overseen in proof reading and I don't understand how that was possible!
Chapter 10: Nest Architecture and House Hunting.The last chapter deals with the 'houses" of social insects: Their architecture, how they are build (including a review of Stigmergy!), house hunting and emigration.
we meet again the waggle dance of the honeybees. Now the "house"-bees must get information not about food but about the quality of the new nest site.
And, in the end, all bees much reach an unanimous decision of the new nest site they prefer, but how is this done? After explaining the process of "quorum
sensing" by the bees, Hölldobler and Wilson take us back to the ants. Here, tandem running, "quorum sensing" and social carrying behavior are
the most important mechanisms to lead nestmates to the new "house" to live in. Also, the mechanisms by which bees and ants determine the quality of
the new nestsites are explained.
ants, the emigration of the queen is described and, for big colonies, the "pre-emigration" digging of a new nest (e.g. Pogonomyrmex and Atta.).
Another incredible piece in the natural history of animals is the formation of
hexagonal cells in honeybee combs. It is the combination of the thermoplastic properties of wax and the production of heath (more than 40°C!) by the bees
inside the cells that make the pure form of the cells appear.
Sadly, again a few typographic errors: page 480, line 16, "(see Plates 17
and 19)" must be "(see Plates 17-19)"; page 494, line 7, "(see Figures 6-25 and 6-26)" must be "(see Figures 6-26 and
6-27)", and line 20, "(see Figure 6-27)" must be "(see Figures 6-27 and 6-28)".
"Our knowledge of the social insects, and the phenomenon of the
superorganism they so beautifully display, has grown immensely during the past century. Yet we have only begun to explore this alien world." (p. 501.).
authors ask now what can be studied next (genetics, ecological pressures, …. and much more!) and, with comparison to history, conclude we can't foresee how
the scientific field under review will evolve the next fifty years.
end with a little comparison between social insects (with their "rigid" instincts.) and us humans (with our "intelligence and swiftly evolving
cultures".). Their hope is that we, humans, will live in harmony "with one another but also with the rest of life." (p. 502.).
always in Wilson's books, a complete list of all who helped with the book in any possible way.
very good glossary including most, if not all, words that need an explanation. Only with one I had a problem: Group Selection. It could be explained better or
my brain was a little bit clouded at that moment.
me, a little bit too small. I always like a big, detailed index to find everything directly when looking for something.
the book the authors mention the following topics that must be studied more:
Kin recognition in colonies.
Recognition of male brood (The mechanism is totally unknown!).
Recognition of individuals with different fathers.
How does it come that up to nine queens can coexist in Pachycondyla tarsata colonies and what are its genetic
The mandibular gland secretions in Atta species: There is a difference between subcastes but does the composition
also change with age? And the responses to them: Do they change according to the place where the substance is emitted and with the behavior of the
How do the Atta queens inform their workers if they are in the nest and how it is with their fertility?
Why do some mature Atta colonies emigrate and what is the communication
system by which these emigrations are organized?
me the book has only two great 'faults". The first one is the lack of a review of the "diseases" of a superorganism. All normal organisms
has to cope with bacteria, viruses, parasitic worms,… during his life. But superorganisms have comparable problems: commensalistic beetles, flies, … that
inhabit their nests and refuse belts, predatory and parasitic wasps and flies, inquilines (social parasites.), … All these can weaken a colony or even
eradicate it! Only the agropredators and agroparasites are covered but they are specialists in the fungus growing societies of the attines and not general
"attackers" of insect colonies.
second biggest problem is that no reference list is included at the end of the book. Normally, the first thing I do with an article or book is going through
the references to see if there is some article that seems interesting but that I missed one way or the other. Not here. By putting the references in notes at
the bottom of a page you make the book longer, all the full references coming back every time. Just a name and year referring to a reference list at the end of
the book would be better. When real explaining or additional information is needed you can put that in notes at the bottom of the page or in a separate
"chapter". But now, reading the notes I meet a lot of full references so many times…
I read the comment "… and they keep repeating the same fact about honey bees over and over again." I was a little bit surprised because when you read
a book about social insects you can expect the dances of honeybees to "surface" one or more times because this is one of the most studied behaviors
of social insects. I didn't hear the same comment for weaver ants, fire ants, gardening ants,… And some chapters indeed overlap. So the dances of honeybees
are covered a few (4) times (with complementary material, needed on that place, included.), just like trail pheromones in Solenopsis, tandem running by ants, castes in Atta gardening ants,…
reprint of William Morton Wheeler's "founding" article (for ants!) would have been very nice in the book. It is: Wheeler, W. M., 1911,
as an organism." Journal of Morphology, 22 (2): 307-325.
this book, Wilson finished a "grand tour of books" though the lives of social insects. Starting with "The Insect Societies" and
"Success and Dominance in Ecosystems: The Case of the Social Insects", about social insects in general, then on with "The Ants" and
"Journey to the Ants: A Story of Scientific Exploration" for his main field of interest and ending with "The Superorganism. The Beauty,
Elegance, and Strangeness of Insect Societies" as a general conclusion of what social insects are and why. For the last three he got the help from
Hölldobler, his co-worker in Harvard for so many years. This book is a must for everybody and a worthy ending of this tour of social insects.
will conclude this little "review" with a question: Why and how do some people read books? Why I ask this question? Let me give a few examples:
a Dutch forum a reader complained that chapter 1 of "The Superorganism" was too heavy and too technical for him, so I ask myself what he should do
with the rest of the book? Won't he read chapters 2, 3, 4 and 7 and parts of 5 and 6? Why did he buy the book then?
an American blog, one reader didn't find a definition of a superorganism in the book. How about the start of the "Note to the general reader"?
Yes, they don't say it are the characteristics of a superorganism, that's true but … Or in the "Glossary"? Or on page 84-85 where you can
find a definition of it and a comparison with a "normal" organism? Or on page 466-467 where a definition is given with a discussion of which eusocial
insects must be included (depending of the degree of reproductive competition.)? Didn't he read it? I only hope that a few readers of this post learn to
read their books better than they have done till now because … Why did they buy the book then?
Many readers had a totally different idea about "The Superorganism" in mind (qua contents and how it was worked out.) but the authors had their own
vision of what they wanted and, according to me, succeeded perfectly in it. It is always difficult to find a book good if you start with a fixed idea about
what it should be. So why buy the book and read it? Maybe some of them should write their own book …
(For our German speaking friends: In October 2009 a German translation will be published by Springer Verlag: Der Superorganismus.).