Book reviews.

Myrmica Ants of the Old World

von Teleutotje » Freitag 14. Oktober 2016, 17:17

“FAUNA MUNDI Volume 3: Myrmica ants (Hymenoptera: Formicidae) of the Old World.”
Alexander G. Radchenko and Graham W. Elmes, 2010.
Publisher: Natura Optima Dux Foundation, Wilcza 64., 00-679 Warszawa, Poland.
ISSN 2081-4615, ISBN 978-83-930773-1-1
789 pages, 333 figs, 163 maps, hardcover, 165 x 235 mm.
Price: 150 euro.

Reviewed by Marc Van der Stappen
Biologist (State University Genth, Belgium, 1987), working in the public transport sector. Interested in behavior and systematics of ants for more than 30 years.
teleutotje@live.be

This “little” book is in a lot of ways much better than I hoped for when I ordered it. It is also not as boring as I, and probably also a few others, might have thought. Someone who orders this book to find descriptions of all the species: well, he will be disappointed. This is not to say you won’t find any descriptions in it; you can find in this book the descriptions of:
M. arisana (first description of queen and male), M. bactriana stat. rev. (redescription of worker and male), M. bakurianica (redescription of worker, queen and male), M. kozakorum n. sp. (description of worker, queen and male), M. lobulicornis (first description of queen and male), M. pleiorhytida (first description of queen), M. pulchella stat.rev. stat. nov. (first description of queen), M. schoedli (first description of male), M. turcica (first description of male) and M. wesmaeli (first description of male).
Beside the already indicated stat. rev., stat. nov. and n. sp., two nomenclatural changes are given: M. ruzskyana nom. nov. (for M. exigua Ruzsky, 1915) and M. slovaca (priority over M. curvithorax). And last but not least, 20 new synonyms are indicated (one each for M. bactriana, M. deplanata, M. rubra, M. scabrinodis, M. schencki and M. transsibirica, two each for M. kurokii and M. sulcinodis, three each for M. ruginodis and M. specioides and four for M. kozlovi.).

But what makes this book so marvelous? The book contains, besides an abstract and a preface, five chapters, each with interesting and good scientific content. It starts with a preface that recalls the beginning of the research on “red ants” (old common name for ants of the genus Myrmica) and the origin of the collaboration of the authors, Radchenko and Elmes. In it, two names are worth remembering: J. Ray and W. Gould (p. 8). Both were important in the 18th Century as pioneers in the systematic and natural history of ants (including Myrmica!).

Chapter 1 describes a “General background and biology of Myrmica”. It gives the scope and the layout of the work, the general biology of the genus, a brief but thorough history of the taxonomy of the Old World species, a review and critique of morphometrics and, last but not least in the chapter, small biographies of all the authors who were important in describing species of Old World Myrmica’s.
The chapter starts with N. A. Weber and his first and the only prior attempt at a worldwide review of the genus. Here I find one of the rare acknowledgements of Jean Bondroit (I. H. H. Yarrow was another important myrmecologist who recognized Bondroit’s work!) “… who had a far better record in naming “good species” of Myrmica compared to his contemporaries…” Radchenko and Elmes originally started a world revision of Myrmica but later omitted the Nearctic species for a number of reasons (p. 13-14). The most important reason is that in the New World there is so much confusion about which names are connected with which species. This part they end with: “In our opinion, the time is ready for a complete revision of North American Myrmica that should start with no pre-conceptions and a fresh examination of the collections to erect a new taxonomy based on the modern concept of species variation in the genus Myrmica. Only then, should the types be examined and the existing names fitted to the modern taxonomy.” They also hope that their book “might stimulate a New World revision.”
After an overview of the book comes the review of Myrmica-biology/ecology/physiology. It’s a nice but quick review that, sadly, is a little bit too quick and jumpy in two paragraphs (p.17 on population dynamics and p. 19 on the frequency distribution of queens).
The part on taxonomic history is divided into a pre-1950 and a post-1950 account. On page 31 the authors indicate a few times that still more species await discovery in the Old World and that even now, after their revision, some problems still exist in the taxonomy of certain species. The review of morphometrics also discusses the critiques about this method.
Last, this chapter ends with the biographies of the most important Myrmica-taxonomists. While most get from ¼ to ½ page, Bondroit has a biography of more than 1 page. This befits their approval of Bondroit as a taxonomist as indicated throughout the book: “…we consider that his appreciation of the genus Myrmica was far superior to that of many of his better known contemporaries.” My impression is that Bondroit’s biggest problem, later used against him, was that he didn’t read the descriptions of species published in reviews from other regions than France or Belgium or that he didn’t understand about which species they really talked. So, except for his four really good species, most of the other names he used or described were known under other names. I also agree with Radchenko and Elmes that Bondroit should be appreciated more by modern taxonomists.

Chapter 2 describes the taxonomic position and definition of Myrmica (description of worker, queen and male) and how it is divided into species groups. Only one little omission in this chapter: although they say in the text that Nothomyrmica was synonymised with Myrmica, in the synonymic list they forget to include Nothomyrmica.

Chapter 3 is the bulk of the book (602 pages!). In it all the species are reviewed. First the 142 extant species, then the five fossil ones and the nine names that are incertae sedis. Each extant species has a full synonymic list, list of type localities and type specimens, material examined, distribution, etymology of the species name and all the synonyms, notes (why names are synonymized, what problems still exist in the species – morphological gradients, cryptic species, eco-types... –, problems with types, eventually descriptions – see the start of this review – and so on) and ecology. Also, all the known and described castes are depicted, drawn as much as possible from type material.
One of the biggest surprises – for me – about the ecology stands on page 103: while there are a few species known to live in very salty environments, workers of M. bergi, when their nest is flooded by a salty lake nearby, are known to “actively swim, sometimes for several tens of meters.” This is described in a Russian publication of 1998. The only other known actively swimming ant is Polyrhachis sokolova from northern Australia.
Other nice ecological points are 1) that colonies of M. pulchella were found in internodal cavities of bamboo (p. 222-223) and 2) on page 233 they describe why M. rubra is such a successful invader.
The most difficult species in the whole book is M. tulinae. The workers are almost identical to M. sabuleti-workers and the males are almost identical to M. scabrinodis-males. M. tulinae belongs to the scabrinodis-complex of the scabrinodis-group (the sabuleti-complex also belongs in this group!). This makes the separation of these species very difficult in regions like Middle Europe. All three species are found there and most collections of these ants are workers (series) or lone males... So, we are obliged to collect nest-series INCLUDING males to separate these species in the future!
Sadly, there are a series of errors, omissions and mistakes in the chapter (most of them are luckily minor ones!):
– Page 107– 108: Eidmann1941 and 1942 should all be 1941.
– Page 108: Although the male is known for M. cagnianti it isn’t depicted.
– Page 115: (see notes to M. rugulosa) should be on page 114 at the end of the notes for M. constricta.
– Page 118-119: The authors didn’t indicate why M. plana is synonymized under M. deplanata. Also, in the synonymic list, M. plana is indicated as described as a subspecies of M. lobicornis but on page 270 Radchenko and Elmes say it was described as a subspecies of M. schencki.
– Page 143: The queen of M. gigantea is known but not depicted.
– Page 146: The etymology of caucasica is easy but not given.
– Page 158: The queen of M. juglandeti is known but not depicted.
– Page 176: Etymology of M. ruzskyi isn’t given (but of course it is obvious!).
– Page 193: First description of the queen of M.lobulicornis but on page 191 it is indicated that the queen of M. alpine (a synonym of M.lobulicornis) was described by Stärcke in 1927.
– Page 239: Types were studied for M. silvestrii but they aren’t indicated in the list of material examined on page 237.
– Page 259: In the synonymic list, under M. rolandi, stands “see notes below and…” but in the notes I can’t find anything about M. rolandi.
– Page 313 under M. turcica: In the notes is indicated “… M. kozakorum (see Notes to that species).” But there you can’t find any comparison with M turcica. Look instead under M. georgica where the story is told (as indicated in the second line of p. 313)!
– Page 382: Queen syntype? No syntype queen indicated on page 113 under M. constricta.
– Page 458: A paratype queen is depicted for M. kirghisorum but the original description is of worker and male (page 167) and no queen.
– Page 489 depicts a paratype male of M. luteola but the male was described four years after the description of worker and queen (p. 197).
– Page 497: Only the head of the male of M. myrmecoxena is depicted. The authors indicate in the description of the species that they didn’t see any males and the figure of the head is after Kutter (1977).
– Page 591: M. specioides, a paralectotype male is depicted but on page 284 the male is excluded of the type-series and on page 288 males are included as paralectotypes. So, are they included in the types-series or not?
If you look at this list you may say “So many errors!” But no, if you consider that for the review of extant species 574 pages are needed, I found very few errors (17 to be exact). Of these only the reason M. plana is synonymized, the list of examined material of M. silvestrii and the notes for M.rolandi really matter (and are what I would also like to know!).
After reading the book I looked up one of the M. plana problems I noticed. It is originally described as Myrmica lobicornis var. plana by Karavaiev in 1927 but in 1929 Karavaiev placed it under M. schencki as M. schencki var. plana. Later, in 1934, Karavaiev synonymized it under M. deplanata (in the same year Arnoldi made plana a "natio" of M. deplanata). So, who can follow all this?
This review of extant species, together with the fossil species and the names incertae sedis, are the core of the book and, yes, they are very well reviewed!
The chapter ends with lists of the nomina nuda, unavailable names and the transferred/excluded species. In the last list Radchenko and Elmes did forget to mention for five species which castes were described (M. nylanderi, M. rugiceps, M. semipolita, M. sordidula and M. striatula.).

Chapter 4 describes the zoogeography and evolution of the genus. This is a chapter everybody needs to read! If you consider Myrmica to be a genus of the plains, forget it: Myrmica is originally a genus of mountainous regions! The boreal fauna is the derived group. Also, a lot of endemic species await discovery in the Central Asian Mountains (some are already being described), South and South-East Asia, the Tibetan Mountains and the Mediterranean region. For the rest of this chapter: it is so good you have to read it yourselves, I’m not going to reproduce it here!

Chapter 5 gives good keys to the species but separate for certain regions. Maybe not the best method for the Old World as a whole but easier for the regions! For the most part they are keys for workers only but for the Western part of the Old World you also need the males to separate a few species.

For the references that end the book, very nice! Only the citations of Weir, J. S. need to be corrected (‘58a and ‘59a are the same and ‘58c and ‘59c are also the same!).

My final and personal judgment: Very, very good. If Nothomyrmica, M. plana, M. silvestrii, M. rolandi and Weir, J. S. were corrected/included the book would be near-perfect. For all working in the Old World, it is surely a book you should have (notwithstanding the price!) or have read.

P.S.1: Both authors of the book have seen a draft of this review and appreciated the comments in it.
P.S.2: Since the publication of this book B. Seifert reviewed the M. salina species complex in 2011, H. Bharti, Y. P. Sharma and I. Gul described nine species from the Himalayas in 2011 and 2012, and A. Radchenko and Z. Yusupov described one species from the Caucasus in 2012.

17 March 2013.
Revised: 31 December 2013.

http://www.asian-myrmecology.org/public ... n-2014.pdf

Battle of the gi-ants: A fight between a fire ant specialist, an author and a reader of "Fire Ants"!
Or: A review of "Fire Ants" by S. W. Taber and it's review.

FIRE ANTS by Stephen Welton Taber Texas A & M University Press. 368 pp. ISBN: 0890969450

"In the early years of the twentieth century, South American fire ants crossed the Caribbean and invaded the shores of the south-eastern United States. These imported fire ants quickly found a niche in Gulf Coast fields and lawns, overpowered the native species, and began spreading. In the process they became a notorious pest to some, a beneficial ally to others, and a potential killer to allergy sufferers. As a result, they are among the most intensely studied insects in the world." Because the literature about Fire Ants has grown to such a massive amount, S. W. Taber decided to write a survey of what has been written about these ants.
The first person to review this book was Mark A. Deyrup, a Fire Ant specialist in Florida. Starting with this review, the debate about the book got sometimes very angry and I decided, reading the book, to review not only the book but also M. A. Deyrup's review. If you allow, I'll start with this first and then look at the rest of the book.
The first problem dealt with the Tropical Fire Ant (TFA) and the elaiosomes of a violet species (Viola odorata) . Both are native to the southern U.S.A. and Middle America . Taber wrote: "In Central Texas the plant "Wedelia hispida" uses fire ants to disperse its seeds. The seeds have oil-rich appendages called "elaiosomes" that the insects eat after they collect the seed and take it back to the nest. The discarded seeds germinate at a distance from the parent plant, thus perhaps avoiding competition with others of their own species. The plant's strategy is called myrmecochory. The arrowroot plant "Calathea microcephala" of tropical Mexico has similar success. But the ploy does not always work. The TFA takes the elaiosomes of the related arrowroot "C. ovandensis", leaving its seeds behind, still attached to the plant and undispersed. The ants build soil tunnels over these seeds while they do their work. At least one violet species is able to save its seeds from fire ant predation with an elaiosome. In the Dominican Republic the TFA disperses seeds of the pope's head cactus "Melocactus communis" in a similar fashion." (p. 73-74). Deyrup's remark in the review was: "The function of the elaiosome on seeds of a species of violet is not to "save its seeds from fire ant predation.""
The quoted passage of "Fire Ants" is all that you can find in the book about the function of elaiosomes. The elaiosome is developed as an organ to attract ants (including TFA) and stimulate them to take the seed to the nest. There the ants eat the elaiosome and the seed stays in the nest or is taken to the refuse heap. In short, the seed is transported to a different place (dispersed) that gives a form of protection (from animals that eat it) and is richer in nutrients (food-products) for the seed! Also, while eating the elaiosome, the ant does not eat the seed. Thus, the plant and the ant developed together and all functions are clear. So, as a way of saying, the seeds are saved from predators, including the Fire Ants themselves!
In the original article about the violet, Beattie and Lyons wrote: "One V. arvensis seed was found destroyed and empty, while five V. odorata seeds had been gnawed, the elaiosomes removed, and returned to the surface intact. …the results of this experiment suggested that predator avoidance plus dispersal could be achieved by providing large seeds with tough seed coats but very large food bodies as bait." and "When the elaiosome is small or perhaps nonexistent the ants are more likely to eat the seed itself. When the elaiosome is present the ants were at least sometimes content with this food and did not follow through on seed destruction." (p. 717).
The second remark of Deyrup was: "Polygyne fire ant nests that bud off daughter nests are not an example of "asexual reproduction."" This is related to two passages in Taber's book: "Queens of the polygyne social type disperse to begin nests of their own by simply leaving the colony in an overland march with a contingent of workers. This is dispersal by a kind of "budding" that calls to mind the asexual reproduction of yeasts or protozoa." (p.24) and "… ; (2) polygyne nests that bud off daughter colonies in a manner like that of a budding yeast cell, when queens simply leave with a contingent of workers; …" (p. 54).
In both cases Taber compares the budding of a colony with the asexual reproduction of yeast or protozoa (to make it visual!) but does not say that it is asexual reproduction! It is only a comparison that was misinterpreted!
Next, Deyrup writes: "The statement that one of the now-rejected chemical controls "might have done the job" strikes me as one of those retroactive fantasies of omnipotence typical of we middle-aged males. By the time such control was begun, the red imported fire ant had dispersed widely. Even if we modern experts had been in charge, we could not have stopped a widespread, highly mobile, generalist species that reproduces rapidly, is easily relocated by commerce, and is beautifully preadapted to the habitat disturbances diagnostic of the ecological niche of our own species."
Taber makes indeed that statement on page xv. But if you read chapter 11 ("Chemical Control and the Pesticide Issue.") you will see that there were a few good candidates to eradicate the Fire Ants! One was "Heptachlor" but it was to dangerous for humans and wildlife and stayed to long in the environment. The other was "Mirex" but this was rejected after a lot of years because they said e.g. that it caused cancer (never proven and related products were tumor-inhibitors.) and it killed and deformed rodent-embryo's (but it was originally released as a rodenticide!). So the two best candidates were rejected although they were very good against fire ants!
The forth dispute was about Taber saying: "An unidentified substance on the TFA exoskeleton decreases the viability of narcissus pollen. This may explain why ants are not pollinators, though they would appear to be ideal candidates because they are dominant arthropods and because pollination behavior is so widespread among their wasp and bee relations." (p. 77). Deyrup's remark goes: "Ants would not make "ideal pollinators" were it not for pollen-inhibiting chemicals, because ants travel everywhere on foot, automatically making them lousy pollinators, irrespective of their body chemistry."
Now , I would restrict Taber's remark to fire ants and not to ants in general because there are ant species that pollinate flowers (e.g. Myrmecia workers in Australia that pollinate, together with a few other species, a series of Orchids. There are also a few other plants that are pollinated by worker or male ants !). For the same reason, Deyrup's remark isn't also correct. It depends on cuticular chemistry and behavior, but it doesn't exclude that there are ants that pollinate flowers!
Now we reach critique number five. Deyrup states: "…Ernst Mayr did not actually state or imply that "the introduction of an exotic is good for all but the invader's closest competitors because the presence of an additional species increases biodiversity.""
In the attacked passage, Taber is wondering if the Fire Ants are pests or blessings and this as an introduction to the chapter "Fire Ants Pro and Con". Now here you have to read a little bit more: "Some evolutionists and even a few ecologists seem to believe that the introduction of an exotic is good for all but the invader's closest competitors because the presence of an additional species increases biodiversity (Mayr 1963; Elton 1958). This view is definitely opposed to the mainstream opinion. However, the issue has never been properly addressed in the case of the fire ants." ( p. 194 ). In a reply to Deyrup's criticism , Taber quoted from page 76 of E. Mayr's "Animal Species and Evolution" (1963): "It should be mentioned that every new arrival in an area tends to add to the total diversity and to enrich thereby the opportunities of other organisms except the most immediate competitors..."
So , Taber does not say that it is good or bad to introduce ants and he is not quoting (exactly) Mayr or Elton. He is just thinking: Is it good or bad and what are the opinions of others? He even says that most are against introductions. The problem is that nobody has looked at the case "The Fire Ants" and in the rest of the chapter he provides points pro and contra for Fire Ants. At the end of the chapter he makes the remark "…that fire ants have a beneficial side that has been largely overlooked." (p. 215) and that we don't have to see them only as big noxious pests!
The sixth remark of Deyrup is: "One can read the entire volume and still lack answers to reasonable questions about fire ants." Taber did not have the intention to answer unsolved questions (A synthesis doesn't give answers to things that aren't looked at!), but to review all that was written about Fire Ants ("My own purpose is a synthesis of the enormous literature on the fire ants of the world.", p. xv.) and, according to me, he succeeded in it! I'm amazed how much information can be included in such a small book! You also have to remember that this book was written in 1999 and published in 2000, so don't expect scientific findings that were published after 1999 (e.g. the review of "Pseudacteon" flies.)! Also, sometimes a synthesis brings answers in the open that stayed in the dark, but not always, and this book doesn't!

Related to this is the next thing Deyrup says: "The vision, complete with map, of future distribution shows the red imported fire ant sweeping up the West Coast to Canada. This seems highly unlikely: the extremes of temperature in Seattle might not be too cold for fire ants, but the northern west coast marine climate is unfavorable to a broad spectrum of ants for other reasons." Now this is something I don't like so much either, predictions about the distribution of an animal in the future. There are so much variables that influence these distributions that speculations and predictions are unreliable! Also, probably most of the interfering factors aren't known yet or aren't quantifiable at this moment! So I don't like these computer-simulations because they are limited in their predictive powers! Taber gives them for the RIFA (p. 219-221) and makes remarks about these simulations (e.g. cold tolerance and global warming problems, p. 223 + p.225.). All this between information on the distributional history of the RIFA. But indeed, these simulations are all unreliable but Taber included them, according to me, to make his literature review complete!
And finally, Deyrup's last remark is: "In the key to species one must check the "dorsolateral junction of the propodeum", a hitherto unknown feature, which is absent from the glossary." This will lead me to the part of the book I didn't like to much ("Appendix 2") and into my own review of "Fire Ants".
If you read the revision of fire ants by J. C. Trager (1991, A revision of the fire ants, Solenopsis geminata group [Hymenoptera: Formicidae: Myrmicinae], J. New York Entomol. Soc. 99: 141-198.), you notice that Fire Ants are not an easy group (Some specimens are, at the moment, not determinable to species!). So I would like to see the following points considered or included in the next edition of the book. First, if you work, for a part, with coloration differences between species in the determination keys and you give pictures to show these differences, don't give the pictures in black and white but in color. I would suggest that at least figs. 1.8, 1.9, 1.10, 1.11, 1.12, 1.13, 1.14 and all the photographs of "Appendix 2. How to Identify U.S. Fire Ants" should be in color . Second, the terminology in the determination keys is not so clear as in Trager's revision (e.g. the feature mentioned above by Deyrup takes, in Trager, 2 to 3 lines of explanation, starting with "…propodeal carinae originating…", in each lug of couplet 3!), but it is still detectable what is meant ! The third point is about the determination keys. Why dividing them in two parts, one for imported species and one for natives? You must get the chance to determine them with the keys even if you don't know if it's introduced or not! Yes, the division is explained on page 231 (If you need a quick determination, you take automatically the keys and not the text 3 pages earlier to start the determination of the specimen!), but I would have liked to see it in the keys! And lastly, the key I prefer is the more elaborated one of Trager! Why not asking Trager if it is possible to include his determination key (couplets 2-7/7c !) in the book. This key is more elaborate and, according to me, it is the best at this moment!

The rest of my review is about some small remarks I have about "Fire Ants".

The first remark is about the sex ratio's mentioned in the book. On page 32 Taber says "…total sex ratio is 1:1 sexual female to male,…" and on page 109 he says "…do produce approximately three times as many sexual females as males." Both quotes are about RIFA! These are contradicting results! But in the field of social evolution, these are important differences. In one case (a monogyne colony with a queen who mated only once!) it is known that females prefer a sex ratio of 1:1 and workers of 3:1 (This has to do with the asymmetric relatedness between mother and son/daughter at one side and sister and brother/sister at the other side respectively.). Later on, in the discussion about his book, Taber said "For example, I think we'll find that there is no contradiction regarding sex ratio. If one investigator finds and reports one thing, and another finds and reports a second thing. They don't have to agree at all. This isn't math, it's the variation of nature and the variation of investigators we're confronted with. I'll have to look. In any event, it probably won't be MY contradiction. For example, the sterile females are of the female sex, sterile or not, and I was merely drawing attention to what the numbers would be like IF the calculations were done differently. And remember, please don't curse the messenger..." This last remark of Taber is about the following phrase on page 32 were he says "The sex ratio would be enormously biased in favor of females if the hordes of sterile workers were included in the calculations." This is never done! Only the possible future reproductive individuals are counted.
Second, on page 43, when talking about polygyne colonies, Taber says "…, the recognition pheromone of one appears to be more attractive to workers than that of any other queen,…". In that case the other queens loses in the end and, after execution of these "losers", one queen remains and the colony becomes monogynous! This can't be the outcome in nature where a polygynous colony stays polygynous.
The third remark is about the location of the metapleural gland on the "…front part of the abdomen…" (p.44) and on "…the anterior abdomen." (p.77). On page 77 Taber also uses the terms head, thorax and gaster (The correct and accepted terms in Myrmecology!) so I would have preferred that he said "…at the lower hind corner of the thorax."
On page 48, for my remark number four, you can read "…, but because the vast majority of males in polygyne nests are sterile for some reason,…". These males are from inseminated eggs (diploid) but the information on the corresponding genes is identical (mostly as a result of inbreeding!). Because of that, these males have problems with the production of sperm cells and that is the reason why these males are sterile! These results are from research done by Ross and Fletcher .
The fifth remark is a very small one about "Chapter 10. Medical Importance of Fire Ants." (It starts on page128.). This is a very good review of fire ant venom and it's effect on humans! The only remark I can make about this chapter is that I would have liked some figures of the alkaloids and the allergenic proteins mentioned in this chapter.
Then, on page 148, I read about "the Mrak report of 1969." But I would appreciate it if I could find the reference for it in the "Bibliography."
Remark number seven goes about a passage you can find on page 151. Talking about virgin egg laying females in polygyne colonies, Taber writes that they "…will probably be limited to the production of male offspring, which are commonly sterile in polygyne colonies anyway." Now, this is the combination of two different things, first that virgin queens lay eggs that aren't inseminated and these are producing males (haploid and fertile!) and, second those males I referred to in my remark number four.

The next, number eight, is about the "Bibliography" that starts on page 243. On page xvi of the "Preface" Taber says a few times that he used "Biological Abstracts" for compiling a big part of the bibliography . Why didn't he also used the "Fire Ant Literature Database" (FALD) or "FORMIS" (In this the FALD is included as one of the foundations.). I know that "FORMIS" doesn't include all ant related articles and books (That's why they keep expanding the old references in it while they make updates!) but these are already vast resources to find scientific Fire Ant literature!
And lastly, there are some typographic errors, not to many and not disturbing ones, but these are not mistakes made by the author and they aren't included here.
Now, in this concluding section, I want, before my closing remark, to mention two paragraphs that I found confusing. Their contents is correct but, to be clear and bright, they needed more space in the book to explain them!
The first one is this: "Tropical fire ant queens do not seem to produce the diploid variety of the male sex, though the RIFA can. Diploid males have twice as many gene-bearing chromosomes as the more common haploid variety of male ant. Nor are the winged TFA virgin queens likely to shed their wings and begin laying eggs when the mated mother queen is removed from her nest. The tendency to do so seems to be unique to one or a very few fire ant species and perhaps evolved when recently mated queens attempted to usurp the reproductive role after joining established colonies." (p. 81). Just don't let you be distracted when you read this passage and everything is o.k.!
For confusion number two, we go to page 114: "It is often found that hybrids are superior in some ways to both parental species, a condition known variously as hybrid superiority or hybrid vigor. An increase in genetic variation is considered an indication of the phenomenon, and although this increase is apparent in the case of the RIFA x BIFA hybrid, the supposed superiority over the parental types does not seem to hold because hybrid fire ants which resemble a given parent more than the other are inferior in competition with that same parent. Yet the hybrid is encroaching upon the BIFA's range and may eliminate the original import with the help of the red imported fire ant. In fact, the range of the hybrid now exceeds that of the BIFA. This trend would seem to conflict with the generalization that hybrids are inferior to the parental types that surround them, and with the generalization that hybrids more like one parent than the other suffer in competition with that parent." It is correct, this passage, but a little bit mind-buzzing like it is written here! I had to read it a few times on a quit place before it sank in and got clear to me! This is the darkest passage in this book!

And now my closing remark to end this long and enduring review: I didn't "find cause to cringe and twitch on almost every page.", no, I enjoyed reading it and did find things I didn't know (e.g. "It is now possible to culture fire ant ovaries in the laboratory for a period of at least eight months. This…" you can find on page 54!). For me this was a very good review of Fire Ant literature (except the determination-key.)!

Van der Stappen Marc

FIRE ANTS by Stephen Welton
Taber Texas A & M University Press. 368 pp.
ISBN: 0890969450

Stephen Taber has gone out fishing on the flood tide of fire ant literature, and has hauled in a ton of facts. Tossed in gleaming heaps on the deck of his 368-page book, these facts fill the entomologist with admiration, as well as with gratitude that any personal obligation to troll the murky depths of a hundred journals is thus obviated. Here, laid out in a row, are the ant-conquered southern states, extending east and west from the Alabama entry point. Here, still feebly and reflexively gasping and flipping their tails, are the bloated and ignominious fire ant eradication programs. Here are the problems of fire ant taxonomy, still tangled in a snarl of fishing line, but out in the open. The entomologist is not the only beneficiary of this bounty; the journalist should be equally pleased. News items, such as, "Fire Ants Kill Easter Chicks at Local Pet Store," a headline that the journalist belatedly realizes hardly requires actual text, can be nicely padded out with educational bits on the potency of fire ant venom and the murder of hatchling quail out in the countryside.
While delighted to see this huge harvest of information all in one place, I would not want my understanding of fire ants to depend on this book. One can read the entire volume and still lack answers to reasonable questions about fire ants. Why are fire ants, especially the imported species, so much more aggressive around the nest than most other ants? How, exactly, do they so successfully displace other species? Why do they spread so rapidly in some regions and much slower in others? Why are some areas within their range wall-to-wall fire ants, while other areas have few nests? What forces brought about the decline of the devastating outbreaks of tropical fire ants in the Caribbean, and does this decline offer hope elsewhere? Symptomatic of this weakness in synthesis, the overviews of past and future fire ant scenarios seem questionable. The statement that one of the now-rejected chemical controls "might have done the job" strikes me as one of those retroactive fantasies of omnipotence typical of us middle-aged males. By the time such control was begun, the red imported fire ant had dispersed widely. Even if we modern experts had been in charge, we could not have stopped a widespread, highly mobile, generalist species that reproduces rapidly, is easily relocated by commerce, and is beautifully preadapted to the habitat disturbances diagnostic of the ecological niche of our own species. The vision, complete with map, of future distribution shows the red imported fire ant sweeping up the West Coast to Canada. This seems highly unlikely: the extremes of temperature in Seattle might not be too cold for fire ants, but the northern west coast marine climate is unfavorable to a broad spectrum of ants for other reasons.
It is easy for inaccuracies to creep in when one is dealing in an uncritical and non-selective way with a large literature (the list of references takes up 56 pages of tiny print). For this reason, one needs to exercise judgement before quoting from the book. For example, Ernst Mayr did not actually state or imply that "the introduction of an exotic is good for all but the invader's closest competitors because the presence of an additional species increases biodiversity." Ants would not make "ideal pollinators" were it not for pollen-inhibiting chemicals, because ants travel everywhere on foot, automatically making them lousy pollinators, irrespective of their body chemistry. The function of the elaiosome on seeds of a species of violet is not to "save its seeds from fire ant predation." Polygyne fire ant nests that bud off daughter nests are not an example of "asexual reproduction." In the key to species one must check the "dorsolateral junction of the propodeum," a hitherto unknown feature, which is absent from the glossary. There are many more of these little mistakes, and also many places where it would be easy to get the wrong impression about some aspect of fire ant biology.
I do not recommend this book for the more sensitive myrmecologist, who would find cause to cringe and twitch on almost every page. For tougher specialists, this book is a great labor-saving compendium.

Mark Deyrup

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FIRE ANTS by Stephen Welton Taber
Texas A & M University Press. 368 pp.
ISBN: 0890969450

The motive behind Deyrup's review of my book FIRE ANTS is transparent, as witnessed by the tenor of the first sentences, the choice of words therein, and by the same weaknesses exhibited in the review's conclusion. I have dismantled the reviewer's thinly-veiled jealousy at the website known as the Ant Farm's Message Board. I would encourage those interested in fair treatment of scholarly work to see Vogt's review in Quarterly Review of Biology, vol. 77, March 2002, page. 73

Stephen W. Taber swtaber@aol.com

A little excursion through the world of books:
Edward Osborne Wilson, Jr., and Berthold K. Hölldobler, Natural History of Social Insects and "The Superorganism.".

([(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.

The 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:

Hölldobler, B. & Wilson, E. O., 1990, The Ants.

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.).
The 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 greatest concerns:

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.

In 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.

What 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.

My 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.

What 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.).
Also 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.

How 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.
The 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.
The 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!).
When the three preceding chapters gave the genetic basis and how decisions are made, now we start with the most important external characteristics of a superorganism.

This 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.).
For 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.).
The 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.
The way how cast are determined are also reviewed (genetic and nongenetic or environmental causes.).
The chapter ends with adaptive demography (and the observation that specialized workers, although good in doing a specific job, don't do everything well) and teamwork.

One "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.

How 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.
The 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.
The chapter ends with the communication of resource-holding potential (with the tournaments of Myrmecocystus as the most important example.).

It 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.

One 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!!!

On 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.

This 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.

One 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.

Here 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.).
- On 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.).

The 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!

If 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.

Two 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 main text!).
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.
It is a fairly complete review, even the relative nutritional values of different fungus-parts are mentioned.

At 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.

The 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!

One 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!
A 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)".

One "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.
Here 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.
For 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)".

Epilogue.

"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.).
The 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.
They 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.).

Acknowledgments.

As always in Wilson's books, a complete list of all who helped with the book in any possible way.

Glossary.

A 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.

Index.

For me, a little bit too small. I always like a big, detailed index to find everything directly when looking for something.

In 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.
More specific:
- How does it come that up to nine queens can coexist in Pachycondyla tarsata colonies and what are its genetic consequences?
- 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 "receiver"?
- 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?

For 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.

My 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…

When 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,…

A reprint of William Morton Wheeler's "founding" article (for ants!) would have been very nice in the book. It is: Wheeler, W. M., 1911, "The ant-colony as an organism." Journal of Morphology, 22 (2): 307-325.

With 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.

I 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:
- On 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?
- On 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 …

Supplement: Complete references and abstracts of mentioned articles:

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Strohm, E., Herzner, G. and Goettler, W., 2007, "A "social" gland in a solitary wasp? The postpharyngeal gland of female European beewolves (Hymenoptera, Crabronidae)." Arthropod Structure & Development 36: 113-122.

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Abstract:

Exocrine glands play an important role in maintaining the integrity of colonies of social Hymenoptera. The postpharyngeal gland (PPG) of ants is crucial for the generation of a nest odour that enables nestmate recognition. The evolutionary history of this gland is unknown and it was thought to be restricted to ants. Here we describe an exocrine head gland in females of a solitary crabronid wasp, the European beewolf, Philanthus triangulum, that resembles the PPG of ants in many respects. The newly described gland has the same location and the same glove like shape as in ants, and it also has a monolayered epithelium with similar ultrastructure. Unlike in ants, the epithelium bears hairs that reach into the lumen of the gland. Although the PPG of beewolves serves a completely different function it is also associated to an allogrooming behaviour as in ants. Based on these morphological and behavioural similarities as well as similarities in the chemical composition of the content of the PPG of both taxa, we hypothesise 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.

Keywords: Philanthus triangulum; Crabronidae; Sphecidae; Formicidae; Postpharyngeal gland; Exocrine gland.

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Herzner, G., Goettler, W., Kroiss, J., Purea, A., Webb, A. G., Jacob, P. M., Rössler, W., and Strohm, E., 2007, "Males of a solitary wasp possess a postpharyngeal gland." Arthropod Structure & Development 36: 123-133.

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Abstract:

The postpharyngeal gland has long been thought to occur only in ants. Here we characterize, by use of light and electron microscopy as well as 3D reconstruction based on nuclear magnetic resonance (NMR) imaging data, a large cephalic gland reservoir of males of a solitary digger wasp, the European beewolf, Philanthus triangulum. Several lines of evidence suggest that this reservoir is a postpharyngeal gland. The gland reservoir originates from the posterior part of the pharynx and consists of two pairs of unbranched tubular structures that occupy a large portion of the head capsule. Its wall is composed of a unicellular epithelium that is lined by a cuticle. The gland contains a blend of hydrocarbons and compounds with functional groups, and we show that the hydrocarbon fraction of the pheromone is congruent with the hydrocarbons on the cuticle. We discuss the implications of our findings for the evolution of the postpharyngeal gland in ants.

Keywords: Postpharyngeal gland; Crabronidae; Sphecidae; Philanthus; Sex pheromone; NMR imaging.

The latest work of two of the greatest myrmecologists in the world:

Hölldobler, B. & Wilson, E. O., 2011, The Leafcutter Ants. Civilisation by Instinct. W. W. Norton, New York and London. XIV + 160 pp.

It includes a prologue, 13 chapters, a glossary and references.

Why did they write the book? In their own words: “If a congress of naturalists were to gather to choose the seven wonders of the animal world, they would be compelled to include the bizarre and mighty civilizations of the attine leafcutters.” (p.1), “If visitors of another star system had visited Earth a million years ago, before the rise of humanity, they might have concluded that leafcutter colonies were the most advanced societies this planet would ever be able to produce. Yet there was one step to take, the invention of culture, making it possible to write this book about them.” (p. 4), “ …, 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 on Earth discovered to the present time.” (p. 127) and “This book is based on one chapter of our recent book The Superorganism (2009), substantial expanded. We have written this book because we want to highlight the remarkable societies of the leafcutter ants, the “ultimate superorganisms,” and to extend and update the fascinating discoveries made by an international assembly of scientist.” (p.129).

Prologue

A little introduction (4 pages) of leafcutter ants, where you can find them, what most people see of them and a general picture of how they live.

Chapter 1 The ultimate superorganisms

This chapter zooms in on leafcutters. It starts with predictions about how many living species there are on earth, how many species of insects and how many eusocial insects. Here follow the characteristics of eusocial insects and who they are. Going on: a general view of ants and a list of some of the “pinnacles in ant evolution”. The chapter ends with a description of a superorganism and what the benefits are from this concept.

One little hiccup: On page 7 H. & W. say that “The ants are divided at the present time into nineteen taxonomic subfamilies.” For the moment, there are at least twenty-one extant subfamilies.

Chapter 2 The attine breakthrough

Two big groups of social insects employ agriculture, the Macrotermitinae and the Attini. After a short introduction of the Macrotermitinae, this chapter gives a review of the attine genera and their symbiotic fungi/yeasts. This chapter includes a time-calibrated phylogeny of the fungus-growing ants and a discussion of the vertical - and horizontal transfers of the fungi. Photographs of workers of the most important genera are also included.

Chapter 3 The ascent of the leafcutters

This little chapter (2 pages) gives some general info of the genera Acromyrmex and Atta (the leafcutters) and their importance in their ecosystems.

Chapter 4 Life cycle of the leafcutter ants

Production of queens and males, nuptial flights, matings (attine queens mate more than once, three hypotheses about the benefits of this are discussed.), nest/colony foundation and the first steps of a colony are reviewed. The chapter ends with a review of the food of the ants.

Chapter 5 The Atta cast system

This is a review of the morphological worker-caste system, of the division of labor based on it, of age polyethism and the adaptiveness of these systems during colony-growth.

In this chapter you notice that this little book is based on one chapter in The Superorganism. The authors make the same hiccup as in that chapter: On page 53 they say “… and strongly skewed to the larger-size classes.” It must be “… and strongly skewed to the smaller-size classes.” In The Superorganism they made the same mistake!

On the same page on line 28 a small typographic error: “(see Plates 20 and 22)” must be “(see Plates 21 and 22)”.

Chapter 6 Harvesting vegetation

A review of how plant material is harvested, from cutting and transport to substrate-choice.

Chapter 7 Communication in Atta

A very nice review of the chemical – and vibrational communication between the ants. It even includes description of the olfactory pathways in the ant-brain. It ends with the question how queens communicate with their workers.

Here I did also notice a small mistake. In the explanation of Plate 34 we read (in the section “Lower right”) “AN (yellow): Antennal nerve” but in the picture the nerve is green!

Chapter 8 The ant-fungus mutualism

It there a kind of communication between the ants and the fungus about the collected plant-material? How are the ants informed about the quality of the material (the feedback) and how do the ants react to that info? Also the recognition of fungal strains is discussed, how these are protected and how fungal strains can be changed.

Chapter 9 Hygiene in the symbiosis

This chapter starts with a discussion of the function of the substances produced by the metapleural glands. The rest of it is about “the ”agricultural pathology” of the ant fungus gardens.” In The Superorganism we read about the interactions between the ants, their fungus, the microfungus Escovopsis and the bacterium Pseudonocardia. In some instances, instead of Pseudonocardia we find a Streptomyces. All is reviewed here also but two more players are introduced in the story, a black yeast that eats Pseudonocardia and a nitrogen-fixing bacteria (Klebsiella?).

Chapter 10 Waste management

All about waste removal (and task partitioning during that.) and how to get a new fungus when the old one is death.

Chapter 11 Agropredators and agroparasites

A little chapter about Gnamptogenys, Megalomyrmex, Pseudoatta and social parasitic Acromyrmex species.

Chapter 12 Leafcutter nests

A review of nest architecture, carbon dioxide – and temperature – regulation and how the ants adapt their nest architecture to humidity.

Chapter 13 Trails and trunk routes

All about foraging tunnels, trails and routes, their construction and maintenance and how they translate to foraging area. This chapter also ends the text with a few last remarks about superorganisms.

Glossary

It gives definitions of all the encountered scientific jargon. I’m only dissatisfied by the definitions of colony odor and nest odor.

References

16 pages of references in the form of notes to the text.

The little book is lavishly illustrated with a few figures and a lot of photographs. Those photos are made by H. Heilmann, H. Herz, B. Höllbobler, M. Kaib, M. Poulsen, F. Roces, W. Thaler, A. Wild and C. Ziegler. A conservative guess is that more than half the photos are made by our Myrmecos!!!

My general impression of the book is that it is good, very good. My favorite chapters are numbers 7, 8, 9, 11 and 12! I will recommend it to all who are interested in leafcutter ants, to all interested in ants and to all interested in superorganisms.

Worth the 23.95 euro’s I payed for it!

Joshua Blu Buhs: About Ants…

This review is about one book and four articles, written by a historian and sociologist of science, J. B. Buhs. His thesis at the University was about The Fire Ant Wars (adapted for the book) but he got therefore access to some archives that nobody saw before and gave him some material incorporated in the book and the articles. They are:

Joshua Blu Buhs.
"Building on Bedrock: William Steel Creighton and the Reformation of Ant Systematics, 1925-1970."
Journal of the History of Biology, Vol. 33, Issue 1: 27-70, 2000.
Kluwer Academic Publishers. – 44 pages

Joshua Blu Buhs.
"The Fire Ant Wars. Nature and Science in the Pesticide Controversies of the Late Twentieth Century."
Isis, Vol. 93, Issue 3, 377-400, 2002.
The History of Science Society. – 24 pages

Joshua Blu Buhs.
"Dead Cows on a Georgia Field. Mapping the Cultural Landscape of the Post-World War II American Pesticide Controversies."
Environmental History, Vol. 7, Issue 1, 99-121, 2002.
Forest History Society and American Society for Environmental History. – 23 pages

Joshua Blu Buhs.
"The Fire Ant Wars. Nature, Science, and Public Policy in Twentieth-Century America."
x + 216 pages, 5 halftones, 13 line drawings, 2004.
The University of Chicago Press. – 226 pages

Joshua Blu Buhs.
"Ground Zero in the Fire Ant Wars."
Alabama Heritage, Issue 82, 24-31, Fall 2006.
The University of Alabama and The Alabama Department of Archives and History. – 8 pages

You should read them in the order they are published!

First you have "Building on Bedrock", the history of the writing of the 1950 classic book on the ants of North America, the reactions on the book and life of W. S. Creighton, the author. In it the "Happy Harvard Team" is introduced, E. O. Wilson and W. L. Brown, Jr., who first agreed with the book but later on got very critical about it and wanted also to control Myrmecology. One of the disputes (with publications on both sides!) was about the subgenus Raptiformica. This subgenus was, for North America, finally revised by W. F. Buren. Creighton and Buren, together with other Myrmecologists, fougth against the "Happy Harvard Team" and, in the end, survived the dispute.

The article "The Fire Ant Wars" describes the technical way Buhs studied these Wars and gives a general overview of the history of the ways the USDA wanted to eradicate the fire ant in North America and the opposition against this chemical war.

The article "Dead Cows on a Georgia Field" describes the history of the first death cows in Georgia after the first sprayings against fire ants and how they had to prove that the chemicals of The Fire Ant Wars were responsible! At a certain moment the USDA said that you can't compare domesticated cows and wild animals!

The book "The Fire Ant Wars" gives a complete history about those wars. First a natural history of the fire ants and the invasion of North America (and why this was possible!). Second the view of the USDA and their allies - chemical eradication. Third the view of R. Carson and allies - control and integration. Fourth the history of the wars, politics and justice included. Fifth the lessons learned and what now - biological control. In this book the discussions between Creighton/Buren and Wilson go on, Buren revises the fire ants and proves Wilson wrong, Creighton blocks support for the USDA because Wilson supported the USDA...

And finally "Ground Zero" reviews all about the fire ants in Alabama/North America (with e.g. some more quotes from Creighton about the fire ants and the wars against them).

This series is a very good review of Creighton and The Fire Ant Wars. There are only a few typographic errors (In the first article and the book a few small words too much) and only two remarks on the book. At the time of publication there was some uncertainety about the name S. invicta (but this is now fixed) and how many times the ant was introduced in North America (now known to be twice). That's all!

For those interested in the history of Myrmecology: Read them all if you can!!!

Buhs did win two prizes with his articles.
"Building on Bedrock": Winner, Forum for the History of Science in America Young Scholars' Publication Prize, 2003.
"The Fire Ant Wars" (Isis article): Winner, Henry and Ida Schuman Prize, 2001. This article was also reprinted in "Environmental History and the American South: An Anthology of Recent Work", Edited by Christopher Manganiello and Paul S. Sutter (Athens, University of Georgia Press, 2009), 345-71.

"An Account Of English Ants" by William Gould, 1747. Together with de Réaumur, Mutis and de Geer the four ground-layers of myrmecology. Huber made myrmecology a big science in 1810 but those four were forgotten or not known for a long time but are as important as Huber. Now I have Gould, de Réaumur and de Geer as copies or books and a review of Mutis. I will tell you already one thing: Gould recognizes only five different ants. That's less than 10 percent of the known English ants...

Gould was a very good observer and, contrary to what Wheeler once wrote, did some experiments (simple and primitive, but real experiments!).

Chapter one describes the difference between workers of the five species recognised by Gould in 'England" and a general description of the worker ant. It includes a description of the petiole and it differentiates the Myrmicinae that have a petiole AND a postpetiole (Gould doesn't use this terminology but it is clear from the descriptions that he means this.). He even includes a description of the four palpi around the mouth of the ant.
One error: Gould says the antennae are use to feel what is in front of or just above the ant. Now we know that its most important function is the detection of chemical compounds (smell and communication.).

Chapter two gives a description of the nests with chambers and corridors, how the workers build them with the help of their mandibles and how they keep them clean. Gould says also that the workers remove the death out of the nest.

Chapter three concerns the queens and workers. The queens lay eggs and start colonies, workers feed the larvae and protect the nests. Gould gives a general description of the queen (stressing the differences with the workers: ocelli, bigger thorax with "hollow or indented Place, which shows as if she had been originally adorned with Wings", gaster full with eggs.) and describes the differences between the queens of the five species. The workers show their respect towards the egg-layer(s) and form a protective group (now: retinue.) around the queen(s). The queens move lower and higher in the nest depending on the season. In the winter no eggs are laid, most of them in July and August. When the queen dies, the workers look after the remaining eggs and larvae. Gould even mentions that some species have only one queen (monogyny.), others a few (oligogyny.). A few errors: Gould thinks that alates are the males (now: males and virgin queens.), workers are sex-less (now: sterile females.), workers walk on their hindlegs and dance for the queens. One funny thing: Gould says that workers are neutral (sex-less.) individuals because if they were males and females, there would have been to much interruptions in their duties for breeding the young.

Gould, de Réaumur and Mutis are, taken together, very good for describing ants, how they live in Europe and, for South America, how leafcutters and army ants live. Yes, all of them have some errors but keep in mind that they are written some 230-270 years ago.

Chapter four explain that queens lay the eggs and workers care for and feed the eggs/larvae. When you cut open a queen you can see it is full of eggs. When the queen lays the eggs, the workers collect them to look after them. The workers don't lay eggs and, when you look inside them, no eggs, no sperm. Gould then describes the eggs of the three castes (worker, queen, male.) and when the queen lays them.
The only problem here is that I'm not sure from the description of queen and male eggs that these are really eggs. With the worker eggs, no problem, these are real eggs.

Chapter five gives the story about the eggs and what becomes of them . It describes how the workers care for the eggs and relocate them (temperature, moisture,...), how eggs "transform" to become larvae, how these larvae grow, how these larvae are fed and protected by the workers,.... Gould describes the worker, queen and male larvae and how long this stage lasts (including their winter-pause.). He mentions the difference between the larvae of different species and explains why the workers relocate the larvae in the nest.

Chapter six is about the pupal stage (the quite phase between larva and imago.). Gould describes how the larva, when full-grown and in an ideal spot, spins a cocoon and how it transform in a pupa. Again, Gould describes the pupae for the three castes and how long the ants stay in this phase and he also gives the differences between the different species. The pupae are relocated in the nest depending on moisture and temperature. Gould even disects pupae and describes how the contents of them becomes a fluid and then reassembles. At this point he also describes the meconium. Gould isn't sure what it is but one of the possibilities he mentions (King's) is almost totally correct. The workers attend the pupae, protect them and recognizes the pupae of their own species. Gould's Red Ant pupae have no cocoon.

Chapter seven starts with the external development of the pupae, how the body parts of the later imago's become visible, how the coloration of the different parts changes,... When the pupae are fully developed but not yet eclosed, they can already move certain parts of the body. The workers open the cocoons and take the pupa out. Here comes a very nice part of the book: Gould tries, and succeeds very well, to explain why he thinks the big pupa with wings and the big alates are females (future queens.) and the small pupae with wings and the small alates are males. He bases it all on morphological characters (with descriptions and comparisons of the pupae-virgin females [both with wings] with the queens.). En passant Gould describes how an alate keeps her/his wings in rest. He describes the alate females (and the differences between the species.) and the males (and the differences between the species.). The males have hooks at the end of the body and Gould says they are, when you compare with other insects, external genitalia. There are clear differences between the females and the males and their fate is different. Females start new colonies and are morphological (external and internal) as queens of established colonies but with wings. Gould describes also how you can, with a little pressure, pop out the male genitals. Males keep their wings because the need them to fly, females lose them because they don't need them. The male and alate females are at the same moment in the nest and together they leave it. They are almost equal in numbers because that is needed (you know...) and there are many of them (because a lot are eaten by birds,...). Females also lose their wings so they can live longer and so can become queens in colonies. At the end Gould also describes the eclosion of worker-pupae and mentions that workers are unique among insects because they don't have wings. In July you can find the most different stadia in an ant nest: worker, female and male pupa, worker, female and male larvae, eggs, alate males, alate females, workers and the queen.
In this chapter are two things that need to be said/mentioned. First, at a certain moment Gould describes that he found a few females together without workers and that, when he kept them at his home, after a while a few of them started laying eggs. The females didn't care for the eggs. Gould doesn't recognize this as queens starting a new colony (contrary to Wheeler's claim.). Second, Gould describes a white worm, lying in a spiral, that you can find in the gaster of females (three worms together.) or males (lonely worm.). This is a Mermithine I think.

Chapter eight is about the work the workers do. Gould starts with saying that he only takes in account the English ants because a lot has been said about the work of different species of ants but he knows only the English ants and doesn't know how other ants live in other places/seasons/climates. Workers are constantly busy from Mach to October and have three main tasks.
1) Building and care of the nest. The workers collect small sticks,... and put them on the nest. They are used as hiding places (to escape for predators.) and to regulate the temperature in the nest (for the young.). Earth is brought to the surface of the nest. In these earthen hills are chambers and tunnels and larvae and pupae are moved up and down in them so they can profit of the best conditions to grow and develop.
2) Care of the young: feeding, relocating (to best places qua temperature and moisture.), protection against predators and heavy rains, opening of cocoons,... The biggest task of these is feeding the larvae. To feed the larvae (and all the other inhabitant.) a lot of food is needed. The workers store the food in the alvus (now: crop.), adapt and improve it there and than regurgitate it for the larvae.
3) Collection of food. The first question Gould tries to answer is if ants store grain for the wintermonths/ make granaries. After a small review of ancient texts, Gould says that England has a different climate and ants don't have granaries there because in the cold winter English ants don't need food (they are in a sort of winter-numbness.). Maybe, for small species, the grains are to heavy, but even big ants don't have granaries. Gould does a few experiments but non of the English ants make granaries or collect grain. He even discovers with his experiments what ants like (sugar, fruit-juice, insects,...) or not (e.g. grain.). Jet ant have a trail system, these street they keep clean and on them cut grass,... This trail system changes with the distribution of the good food-sources. When a colony is relocated, a trail system is reformed within one week. He also discovers that some ants eat other ants, prey items are placed in galleries with larvae, in the winter English ants only need water and peace.

Chapter nine is about the use of ants: Why are there ants on this earth? First, as food for birds. Their yearly and daily cycle and their behavior is adapted at the development of the birds that feed on them. Second, some products and phases are used (in that time.) in medicine. Third, they are food for certain mammals and insects. Some arthropods/insects also live in or on ant-nests.
This chapter and book is ended with a list of what is still needed to be known:
1) Why are there species with bigger and smaller workers,
2) Sometime workers carry others with their jaws (without hurting them.). Why do they do this? A kind of sport?
3) Why do some ants feed on their own species? Mostly weaker or injured individuals are attacked.
4) How is a new colony started? And how do they disperse?
5) Why do they like fruit? But at the same time they destroy a few fruits they also attack other animals that are harmful to fruit and trees.
6) How best to destroy a colony? At that moment nests were destroyed in winter but the queens are at that moment deep underground and the colonies mostly survive. In the summer the queens are higher up in the nest and so the colony is more vulnerable to destruction.
7) When a queen dies, workers go on with their jobs. This is because the larvae are so important. For what other things are ants so important?

To end:
A marvellous little book, well written, that can make you fall in love with ants (if you aren't it at the moment!)... They should bring out a new edition of it and, in the same style as Gould wrote, add a chapter about colony-founding and dispersion of ants. The title should then be: "An Account Of Common English Ants"!

"William Morton Wheeler, Biologist" by Mary Alice Evans and Howard Ensign Evans, 1970, Harvard University Press.

What a book!!! Let me start with the mistakes in it (quick so that I don’t forget them!): Two typographic errors , both related to dates (p. 69, line 15, 1897 must be 1891; and page 203, line 5, 1923 must be 1922) and one unclear passages (on page 233, when talking about De Vries in 1901, they say “… a decade before the rediscovery of the work of Gregor Mendel…” and a few lines further “… a year earlier had been one of the three workers who independently rediscovered Mendel’s paper…”). But for the rest...

What a marvelous book! Shame for the current Publisher (not H.U.P.) that they don't want to sell this book in Europe. Most biographies I've read I'm not impressed by but there are five very good ones that are worth reading as far as I know, two about Einstein ("Subtle is the Lord. The life and work of Albert Einstein." and "Einstein was here.", a companion for the first book, both by A. Pais), one about Tolkien ("J. R. R. Tolkien, a biography.", by H. Carpenter), one about Columbus (sadly, can't remember the author) and then this book about W. M. Wheeler.

Evans and Evans wrote a book about a scientist (not only the myrmecologist but the complete biologist), his life and his work (like insect-embryology, myrmecology, study of insect sociality and sociology, ethology, …) that is unbelievable good (and as far as I know discusses every facet of his work and his evolving thoughts!) and should be a must for every student of (the history of) myrmecology. It also includes introductions to others Wheeler met or was influenced by and fragments of the letters between Wheeler and his closest friends. Sadly Charlotte Sleigh (in “Six Legs Better; A Cultural History of Myrmecology.”, 2007) noted that the family-archives of the Wheeler's (used by Evans and Evans) are mostly destroyed including a lot of letters, so a lot of interesting stuff has disappeared...

This leads me to this: Shame for Sleigh for two big mistakes in her book. First, on page 291 in her book, she says that the Evans’s wrote this book “… in order to bolster the role of whole animal biology (and myrmecology) when it was going through a rough patch in Harvard’s departmental structure. Its polemic purposes are not too intrusive,…” Of this I didn’t notice anything and I’m inclined to think this is not true. About all his professional jobs, Evans and Evans report in the same way (or, at the moment the book was written, all biology-departments on the included universities should have been in danger of disappearing!). Second, she includes Wheeler in the morphological/sociological myrmecologists. This is the second phase between the psychological (e.g. A. Forel) and the communicational myrmecologists (e.g. E. O. Wilson). But when I read Evans and Evans, I get the impression that Wheeler evolved from a morphological (Loeb et al)/sociological (the superorganism) myrmecologist to a psychological myrmecologist (a reasonable evolution according to Wheeler and Evans and Evans!)…

But, back to Evans and Evans. One of the things I find marvelous is their prophetic views. Writing in 1970, they said that Wheeler was the last one to write a review of everything that was known about social insects (1923 and 1928) but it needed to be replaced (one year later Wilson’s “The Insect Societies” appeared) and, on pages 270, “… contemporary students are… providing the material for a fuller synthesis at some future date.” That synthesis was published by Harvard U. P. in 1975 “Sociobiology. The New Synthesis.”, also by Wilson.

I knew already a lot about Wheeler but I must say I’ve also learned a lot about him that I didn’t know just by reading this book. One of those things: I knew Wheeler compared the ant colony with an organism (1911) but his “Emergent Evolution” (connected with this concept of the superorganism!; 1926, 1927 and 1928) was new for me!!! Wilson aborted in 1967 the superorganism to revive it around 1990 (with other scientists!) and, together with Hölldobler, in 2009 wrote the big reintroduction of it. I also read things about Wheeler that, knowing them before, I didn’t know Wheeler was the author of (like one of the jokes below!) or said it...

So, for everybody, if you have the chance, READ THIS BOOK!

Let me end with one questions and two Wheelerisms (like they are called, jokes about a serious subject with a serious contents, like Wheeler only could make them!).

The question:
It is about the American Museum Congo Expedition. Wheeler described ants from that expedition and wrote about the distribution of African ants, determination keys for the genera and subgenera and synonymic lists. Probably he didn’t go to Africa but is this correct?

The Wheelerisms:
Page 195-196, about human reproduction and the family: “Evidently a very considerable proportion of the population is over-sexed, under-sexed, intersexed or no-sexed, and, therefore, not well-suited to family life of the old-fashioned, rural, or garden variety. It is, of course, very easy to tell all these people to go to hell, but many of them are so devilishly attractive and, apart from their sexual behavior, so very efficient socially, that they are constantly being married by those who are blessed with normal sexual proclivities and ideals.”
Page 228, about psychoanalysis: “… Insects undoubtedly sleep. Do they dream? If they do, what a pity that we shall never be able to apply the Freudian analysis to the dreams of that symbol of sexual repression and sublimation, the worker ant!”

For those that can read it: Enjoy this marvelous journey through history!

In this review I will discuss some quotes from :

Wheeler , W. M. , 1926 , " The Natural History of Ants . From an Unpublished Manuscript in the Archives of the Academy of Sciences of Paris by René Antoine Ferchault de Réaumur . Translated and Annotated by William Morton Wheeler " .

Sibling species .

In the chapter about " The Life and Work of Réaumur " , W. M. Wheeler writes the following about sibling species ( pp. 31-32 . ) :

" Furthermore , in entomology as in other domains of zoölogy we are now confronted with the problem of physiological and ethological species , that is , with forms which show little or no appreciable difference in structure but which differ greatly in behaviour . Giard in his paper on poecilogony called attention to a long list of forms which are almost or quite indistinguishable as adults but differ in their development , and others have from time to time detected differences of habit and habitat , seasonal distribution , and immunity to disease , etc. , in forms otherwise indistinguishable . No doubt many more such cases will be brought to light and will have to be provided for in our classifications . Hence the taxonomy of the future may be expected to place increasing stress on physiological and ethological characters . "

This is indeed a problem in a lot of animal groups , including ants . Examples are e.g. the genera Lasius , Formica , Myrmecia , .....

Classification of animals .

In the same chapter W. M. Wheeler says about classification ( p. 32 . ) :

" The future task of taxonomy , it will be admitted , is a formidable one , but must be undertaken if classification is to be a system embodying in condensed form all the ascertainable data , functional as well as structural , relating to the genetic affinities of organisms , past and present . "

We are already good under way but still must do the biggest part of it !

The Manuscript .

The original manuscript was written between October 1742 and January 1743 . The " Notes " are from the years 1720 , 1734 , 1735 , 1742 , 1743 and 1744 . So , for the following quotes , they are that old !

The original " Histoire des Fourmis " is 66 pages long , followed by 20 pages of " Notes " . They are written in very old French ! The translation " The Natural History of Ants " is 67 pages long , also followed by 20 pages of " Notes on Ants " . At last , the 162 " Annotations " comprise 42 pages .

In the following chapters , de Réaumur's passages are followed ( in brackets ) by W. M. Wheeler's translation !

Huns and Tartars .

We start with a little quote from de Réaumur ( p. 47 . ) :

" Des fourmis de plusieurs autres especes n'ont que des etablisements passagers , elles peuvent etre comparees aux tartares . Elles campent pour ainsi dire , elles sont toujours prestes a quitter le lieu ou elles s'etoient arrestees , des qu'il cesse d'avoir les commodites qui les avoient determiné a le choisir . " ( Wheeler , p. 135 : " Ants of several other species have only temporary abodes and may be compared with the Tartars . They bivouac , so to speak , and are always ready to quit a spot where they have tarried as soon as it ceases to afford the conveniences that led them to select it . " ) .

It sounds a little bit like the phrase of W. M. Wheeler ( in his 1910 book about ants . ) " .. the Huns and Tartars of the insect world ! " and de Réaumur also talks about bivouacs BUT it is not about Eciton ( or any other army ant genus / species ! ) ! It is about the flooding of ant nests ! Don't forget : This was written in the 18th century !

Earthen Anthills .

Then , about earthen anthills in Maine , France ( p. 52 . ) :

" Elles gatent les prés a un point qui engage ceux qui les afferment de metre pour une des conditions du bail qu'on sera oblige chaque annee d'abbatre les fourmilleres , d'applanir les endroits ou elles se trouvent . " ( Wheeler , p. 140 : " They injure the meadows to such a degree as to induce those who let them to require among the conditions of rental the obligation to destroy the formicaries every year and to level the ground where they occur . " ) .

I wouldn't like to be the owner of a lot of land in those times !

Lubbock Nests .

De Réaumur kept bees in transparent glass hives . He covered the glass panes with shutters ( leaving between both a distance of an inch or more ! ) so that , when not observing the bees , the bees where in the dark . When he wanted to do observations , he opened the shutters . A few species of ants regularly made their nest between the glass panes and the shutters to " profiter de la chaleur que les abeilles entretienne dans leur habitation " , " de profiter d'un vuide ou elles peuvent jouir constamment d'une chaleur douce " ( p. 57 ; " to profit by the warmth which the bees maintain in their habitation " , " profit by a cavity in which they can constantly enjoy a pleasant warmth " , p. 145 . ) .

Now , he wanted not to disturb the ants any more and do also observations on them . So he adapted his hives so that it became possible to do this . Here follows the rest of his story ( p. 58 . ) :

" J'ai arreste un carreau de verre en dehors de la ruche , a fleur de l'endroit ou se trouvoit la surface d'un volet fermé . Ainsi le nid a ete renfermé entre deux carreaux , et n'etant plus appuie contre le volet on pouvoit ouvrir celui ci sans derranger aucunement les travaux des fourmis . Autraver d'un verre bien transparent on observoit aisement a quoy elles etoient occupees . Rien n'etoit cache de l'interieur d'un nid si mince et plein de cavites disposées par etages , on le voioit enfin dans toute son epaisseur , et du haut en bas . " ( Wheeler , p. 146 : " I attached a pane of glass outside the hive at a level with the inner surface of the closed shutter . Thus the nest was enclosed between two panes and , being no longer supported against the shutter , it was possible to open the latter without in any way destroying the work of the ants . Through a very transparent pane it was easy to observe what they were doing . Nothing could be concealed in the interior of a nest so thin and full of cavities arranged in storeys : its whole thickness was visible from top to bottom . "

Wheeler , on page 229 , adds here note # 25 : " The description shows that Réaumur actually constructed for his ants the same type of artificial nest as that employed by Lubbock and since regarded as his invention ! "

So , de Réaumur had multi-purpose hives : On one side you could see bees and on the other side ants !

The Ant and the Cicada .

When de Réaumur mentions " La charmante fable de la fourmi et de la cigale " ( The charming fable of the ant and the cicada ; pp.59 and 148 . ) , Wheeler gives a similar fable from the Maoris of New Zealand ( Note # 28 on pp. 229-230 . ) :

" The cicada is treated in Maori fable as the personification of slothful carelessness and the ant as the emblem of industry and forethought . "

And then the fable :

" The pokorua ( ant ) said to the kihikihi ( cicada ) , ' Let us be diligent and collect food during the summer , that we may retain life when the winter arrives . ' ' Not so , ' remarked the cicada ; ' rather let us ascend the trees and bask in the sun on the warm bark . ' Even so , the ant laboured at collecting and storing food for the winter . The cicada said , ' This is true pleasure , to bask in the warm sun and enjoy life . How foolish is the ant who toils below ! ' But when winter came and the warmth went out of the sun , behold , the cicada perished of cold and hunger , while the ant , how snug is he in his warm home underground , with abundance of food ! "

It seems that there are Maori songs sung by the ant and the cicada , but Wheeler doesn't give them .

This Maori fable is almost the same as that from De la Fontaine ! They only make one mistake : For the ant they say " he " in stead of " she " ! Most people know the version with the grasshopper but it appears that in the " primitive " human tribes ( Greeks , Maori , ... ) it is a cicada , not a grasshopper !

Sexuals and nuptial flights .

Now , this is a very interesting part :

Swammerdam , in his publication of 1737-1738 , talked about the castes of ants but thought that the big winged individuals where males and the big wingless individuals were queens . He didn't observe the small winged individuals .

In 1741 Carl von Linné ( Linnaeus . ) described the winged castes as females and males ( and compares them with the queens and drones of bees ! ) and describes the deälation of the queens AND males .

De Réaumur was the first ( in this manuscript , 1742-1743 ! ) to get it OK . Big alate individuals were the virgin females , big wingless individuals the inseminated queens and small winged individuals the males . Queens lose their wings after copulation , males not ( but later in his text he says that males lose their wings also ! ) . He reports in this book also other observations made by him in 1731 : He was the first to see nuptial flights and sees also as the first the copulation in ants ( a Myrmica species , later also with other species . ) ! He only thinks that females and males of the same nest copulate . In 1747 William Gould published the same observations and in 1777 De Geer .

So , following the dates of observation and publication , de Réaumur was the first to get it all ( almost . ) correct and the first to make the observations about nuptial flights and copulation .

I don't include here the original version ( pp. 72-81 . ) and Wheelers translation of it and his notes ( pp. 161-170 ; notes # 44-57 , pp. 241-244 . ) because they are much to long . Wheeler only makes one big mistake : Linnaeus is called by him Carl von Linnaeus !

Ponera .

De Réaumur mentions a few times that after the nuptial flight the males shed their wings and start to do the same work as the workers . He also says that the males and workers look alike but can be distinguished by the genitalia . Wheeler , in his note # 62 on page 245 , says that de Réaumur was not clear about the male caste and that this was in part due to de Réaumur's crude way to determine the castes by their genitalia . The worker's may look like those of the male . My remark about this is the next : De Réaumur is indeed mistaken by the shedding of the wings in European ants BUT he can be confused by the presence of wingless males in some European ants ( e.g. some Ponera species ! ) .

Colony - and larval development .

Now we get to the most remarkable part of the book , that about the development of colonies and larvae ! It is a very big part ( original text from page 84 to page 96 , translation on pp. 173-185 and notes # 65-84 on pp. 245-249 ! ) so forgive me that I don't reproduce them here !

First , de Ráumur starts with the foundation of ant colonies . Gould , in 1747 , gave an account of the founding of ant colonies but only the very beginning ( untill the queen laid her first eggs ! ) and we had to wait until 1876 to get a complete description ( untill the first workers are there ! ) by Lubbock . BUT in this book , de Ráumur gives the result of research done by him a few years before . He did find three queens running around and did put them together in a formicary . The queens made a common primary chamber and started to lay eggs . The queens didn't leave this chamber to forage ( so a claustral colony foundation ! ) . He also saw the first larvae coming out of the eggs . That was the end of his experiment ( he never saw here the complete development to workers ! ) . So he was further in this research than Gould .
But de Ráumur didn't know if the queens did need workers to start a colony or not . But because queens were hard to find alone in nature , he thought that they needed the help of conspecific workers to start their colonies and to help them to produce their first workers !