Beiträge von Teleutotje

...and a key:

A new species of Boloponera from Sekhukhuneland, South Africa (Hymenoptera, Formicidae, Ponerinae)
Peter G. Hawkes
https://zookeys.pensoft.net/article/28606/download/pdf/

Here follows an example of how your impressions of a book can change when you read it....

The book:

Karl Gößwald
Die Waldameise. Biologie, Ökologie und forstliche Nutzung
Prof. Karl Gößwald hat sich Zeit seines Lebens dem natürlichen Biotop- und Artenschutz verschrieben. So hat er stets die These vertreten, dass die „Rote Waldameise“, vorwiegend aus der Gattung Formica bestehend, von größter Bedeutung für das ökologische Gleichgewicht des Lebensraumes „Wald“ ist. Gerade in einer Zeit, wo dieses Ökosystem durch zahlreiche äußere Einflüsse erheblich gestört wird, ist eine Rückbesinnung auf biologische Vielfalt dringend erforderlich. Hier nimmt die Waldameise eine wichtige Rolle ein: Neben der Bekämpfung von Schadinsekten fällt die Verbesserung der Bodenstruktur, die Verbreitung von Sämlingen, der Schutz von Lachniden u.v.a.m. in ihr „Ressort“. Grund genug, das ursprünglich zweibändige Lebenswerk dieses Waldschützers in Form einer einbändigen, auf das Praxisfeld zugeschnittenen Sonderausgabe neu herauszugeben.
1. Auflage, gekürzte Sonderausgabe in einem Band 2012. 652 S., 143 s/w-Abb., 126 Strichabb., 22 Farbtaf., 22 Tab., geb., Format 17 x 23 cm.
Preis: 49,95 € ISBN: 978-3-89104-755-2 Best.-Nr.: 315-01129 Aula Verlag, Wiebelsheim.

The original books are:
Gösswald, K., 1989: Die Waldameise, Band 1: Biologische Grundlagen, Ökologie und Verhalten. (xi + 660 pp.)
Gösswald, K., 1990: Die Waldameise, Band 2: Die Waldameise im Ökosystem Wald, ihr Nutzen und ihre Hege. (x + 510 pp.)

The new book:
Gösswald, K., 2012: Die Waldameise. Biologie, Ökologie und forstliche Nutzung. (xii + 630 pp.)

First impressions:

Gösswald died in 1996, so he didn't help with the new edition of his book(s). And I think it shows! As a reduced new edition of the two original volumes, I thought it would be easier for everybody to read and to follow. And it is also adapted for use by nature-conservationists, so, hopefully practical.

When I looked through the new one for the first time, my impression was "We're back in the 80's!". And when I got at the references "Yes, they improved that!". You must know, the biggest negative point about the original books was the literature lists. But now, a modern-looking list (with the titles of the articles!), Yes, Good.

Now, except those two literature lists in the original books, both books were very good. They were exhaustive in everything (only when you got to the details of the details of the details of the... you were referred to another publication!). But those 1170 pages did let you know everything about Formica s.s. in Europe that you had to know when you were interested in how they looked, their anatomy, their systematics, their determination, their behavior, their benefit for the forests, their protection, handling and keeping them,...

And now I have the new book... I'm going to read it, yes (I did start with it already, got to page 8!) and will tell you what I think of it... And I hope that I can tell you very good things about it but at this moment I can only start with negative points (except for the literature list, see above!). First a few minor things: typographic errors. I met already a few of them: wrong letters (e.g. Prothorokalganglions, must be Prothorakalganglions) and missing spaces (e.g. Formicapolyctena, must be Formica polyctena).

But more, in the explication of Farbtafel 3 (Color plate 3) I find a reference to fig. 196 that gives the respiratory system of a Formica-worker. In this book, fig. 196 is about woodantprotection through the protection of nests and the division of nests. In this new book there is no picture of the respiratory system of an ant! But in Band 1 from 1989.....

The systematics are still the same as in Gösswald's time. So, NO mention of F. paralugubris! A conservationist in the field should also be in the possibility to determine it I think!

The general description of a woodant (and the difference between male, queen and worker!) and it's anatomy occupies only 4 pages with, e.g., almost nothing about the exocrine glands that are so important for the ants. In the original books you were over flooded with info about all those glands!!!

So, I'm sad about a bad start for a new book!

And it gets not better.....:

Yes, me again. I won't do a full review of this book... Why? It's clear for me now already. I'm not going to recommend it to anybody anymore. Why? Reached page 12 now and I did find a second reference to a figure that isn't in this edition any more. More? Yes, even the reference list isn't good (contrary to what I thought at first glance!). After 4 of the 69 pages of references I got really annoyed with it: One reference without title, one repeated twice, one twice under different dates and one twice under different authors.

I'm still going to read it completely but only for myself but no review of it! This because my judgment is too much influenced by the very sloppy work done by the editor/publisher of the book. I even think Gösswald, if he was still alive, wouldn't like it with all the errors/mistakes in it.

I only hope I can find something I didn't know or that I did forget but is worth to rediscover but, no, don't bother about it. For everybody that wants to read Gösswald's work about woodants, try to find the original books from 1989 and 1990.

I will read this one just for me and, when needed, will go to the originals when I need a good review of European woodants. Luckily, I have them here at home. The publisher hasn't got the originals any more on offer. Shame, should have kept those on sale and not this "reworked" version!

And so it ended, without finishing the book:

Merkur (at that moment in time) hopes I'll continue my review of this book but why? For making a long list of all that can go wrong in a book?

I completed the literature list and the types of possible mistakes has got longer:
- There are a few references where you find author, year and part of the title and then... finished, nothing anymore.
- One starts, gives another reference and then concludes the first.
- One is repeated twice, first time with pages 127-198 indicated, then with pages 170-173.
- For Gösswald, his 1985 book is twice in it and two of his publications of 1951 are each four times in it.

For the text, I've reached page 26 and misery goes on.
Again, a few times genus name and species name united.
What is much worse: A determination key with words missing.
Then, when reading about Raptiformica, in the middle of the paragraph, starting about something completely different.

I think that this says enough about the editors of this book. But they are safe, their name is nowhere mentioned so they go unharmed away from this book. Worse is of course the damage done to Gösswald's name.

So, sorry Merkur, but I won't go on with this. It's not a bad book written by Gösswald BUT a very bad edition made by one or a few editors. So I don't say anything anymore about this book because it's sloppy appearance is NOT Gösswald's fault.

For everybody: Read the original 1989 and 1990 edition and forget this one!!!

End of my review. Won't say anything anymore about it!!!

And that was it. I never looked in it again! And when asked about Formica s.s. and working with them, I keep referring to the two original books.

Second , the development of workers / sexuals . He begins with saying that an ant develops in this sequence : eggs to larvae to nymphs / pupae ( with or without cocoon ! ) to adult workers / sexuals and that the place of the larvae and pupae in the nest are regulated by temperature and humidity ( Swammerdam , 1752 , also observed that ants relocate their young in the nest , depending on the temperature ! ) . And he also says that by disturbances the workers bring the larvae and pupae in security !
But this is not yet finished . He gives descriptions of ( followed by some other observations by him . ) :
- The eggs : They swell in six days six times in diameter . He doesn't know how this happens but gives a few hypothesizes how it could be .
- The larvae in the eggs before they come out of them : There is also liquid in those eggs that serve to feed the larvae .
- The hatched larvae : De Ráumur was the first to describe the feeding of the larvae by liquid and not solid food ( earlier than Gould , 1747 ! ) . He follows by saying that the workers also licked the larvae . The duration of the larval stadium he didn't know but he knew the larvae moulted a few times ( he found their skins ! ) and supposed that it was about fourteen days ( but this was to short ! ) .
- The pupae of workers , females and males and the changes in coloration they go through : In some of the species , larvae spin cocoons , others not . Those larvae that spin cocoons stay for two days in it before they change into pupae . De Ráumur gives a detailed account of the spinning of the cocoon and that the duration for making it is around 29 hours . But then , he gives , as the first entomologist ever , the next observations : For starting the cocoon , the ant-larvae need solid bodies , like particles of wood , to glue the first treads on . When the cocoon is finished , the workers remove the solid particles ! De Ráumur doesn't know how long the pupal stage lasts , but the callow ants escape from the cocoons on their own . Since a long time , it is believed that De Geer , 1771 , was the first to see that the workers assist the callows to escape from the cocoons by opening them , but the first one to witness this was Lyonet in December 1743 , followed by Gould in 1747 .

Carrying of workers .

When studying the disturbance of Formica pratensis-nests ( pp. 101-105 in the original manuscript , pp. 190-194 in the translation and note # 96 on page 251 . ) , de Réaumur observed that some Formica-workers transported some rounded granules . When looking closer , he saw that they carried another worker ant . Bonnet was the first to observe this kind of behavior in Leptothorax ( and later on in Formica . ) in 1739 and published it in 1745 but he interpreted this behavior wrongly ( as part of hostility or irritation ! ) . De Réaumur however got it OK ! He describes that both ants had grasped one of the mandibles of the other , the carried ant brings its gaster under its thorax and fold up his legs . He also describes how an ant asks to be carried and how the other can refuse this to do . The carried ants are mostly workers but possibly sometimes a male !

Notes on Ants .

Now follow a few observations in the " Notes on Ants " !

- De Réaumur describes in short the difference between larvae of workers on one side and larvae of queens / males on the other side . He reports that the workers lick the larvae and that they relocate the larvae depending on the humidity of the chambers in which they are placed ( pp. 113-114 , pp. 202-203 and note # 116 on p. 256 . ) .
- De Réaumur describes an experiment in which he relocates larvae from one colony to another and that the workers of the last colony accept these larvae ( pp. 115-116 , pp. 204-205 and notes # 117-119 on p. 256 . ) .
- De Réaumur reports about an experiment he conducted about colony foundation in ants . He placed two queens together with four workers . The deälated queen carried workers like described before ( pp. 118-119 and p. 208 . ) .
- He describes how ants pass liquid food from one ant to the other : " Les fourmis qui se rencontrent se donnent a manger . L'une presente sa langue qui est lechee par celle de l'autre " ( p. 122 . ) , " Ants , when they meet , feed one another ; one of them presents her tongue , which is licked by that of the other " ( p. 211 . ) . Wheeler , in his note # 138 on p. 259 , says " This seems to be the first description of the method of mutual feeding among ants " .

Some other things et al.

- In this work and in the third and fourth volumes of the " Mémoires pour servir à l'Histoire des Insectes " , de Réaumur describes the relation between plant-lice / scale-insects and ants ( he was the first to discover this ! ) . He says that the ants lap up the sugary liquid produced by these insects . He makes one mistake , he says that this liquid in plant-lice is excluded by the anus ( correct ! ) and the two cornicles ( wrong , they produce a defensive substance ! ) . In scale-insects he was not sure about the source ( the same as in the plant-lice ! ) . See pp. 64-66 , pp. 152-154 and notes # 32 and 34 on pp. 231-232 .
- In the discussion about the formation of polydomous colonies ( pp. 96-103 , pp. 186-192 . ) , de Réaumur gives some observations made by three others , Jean Baptiste du Tertre , Jean Baptiste Labat and Maria Sibylla Merian . They published some info about ants in Martinique ( the first two . ) and Surinam ( the last one ! ) . Now , these are a mix of observations , attributed to one ant , that are now allocated to different species and subfamilies ( on one side Eciton burchelli , E. hamatum or Labidus praedator belonging in the Ecitoninae and on the other side Atta sp. , a Myrmicine ) . The ants have fixed nests . The forays of these ants are explained as swarms that need a few days to find their new , fixed lodging . They cut leaves with their long mandibles . These parts of the leaves are used as larval food . The ants form living bridges between objects to far apart . But de Réaumur is very doubtful about all this ( see pp. 97-99 , pp. 186-188 and notes # 85-91 on pp. 250-251 . ) !!!
- In note # 37 , Wheeler describes the invasion of different islands by " Formica omnivora Linn. 1767 ( Myrmica omnivora Latr. ) " and " Formica saccharivora Linn. 1767 " . The first one ( 1 ) is originally described as " Formica domestica omnivora Linn. 1767 " and is now determined as Solenopsis geminata Fabr. The second is based on " Formica minima saccharivora Brown " ( 2 ) and " magnitudo Formica caespitum " ( 3 ) . ( 2 ) is now determined as Tapinoma melanocephalum and ( 3 ) as Paratrechina longicornis Latr. In some publications ( 1 ) and ( 3 ) are inter-changed with the other explanation causing some big confusions ! This note is to be found on pp. 232-239 ( almost 7 pages ! ) .

The End : A Myrmecological Time Machine !

It is clear the Reaumur was far ahead of his time in his understanding of ants and in the methods to study them . No doubt , this must be the reason why W. M. Wheeler felt compelled to translate this manuscript . It is wonderful to have these glimmerings from the dawn of modern myrmecology .

René Antoine Ferchault de Réaumur was the first modern entomologist that studied ants ( among other insects . ) . This history about ants was never published because he started a discussion with an other big zoölogist of that time , Buffon . Buffon had so much power that he decided or something would be published or not . Unlucky for us , Buffon decided against the publication . The manuscript wasn't know for so long and even now , most myrmecologists don't know it . I'm very happy W. M. Wheeler published it !

Hope you all liked this little trip back in time ! I did , so .....

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 !

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 ! "

"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!

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"!

"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.

Seifert, B. (2018): The ants of Central and North Europe. Lutra Verlag- und Vertriebsgesellschaft, Tauer.

An illustrated determination key to 190 species recorded for the area and directly adjacent regions.
ISBN: 978-3-936412-07-9
407 Seiten, 581 Fotos und Zeichnungen von Ameisen, 1047 Titel im Literaturverzeichnis.

Radchenko, A. G.; Dlussky, G. M. 2018. A new species of the extinct ant genus Electromyrmex (Hymenoptera, Formicidae). Vestnik Zoologii 52:361-366.

http://www.antcat.org/document…lectromyrmex_wheeleri.pdf

Taxonomy of coccids (Hemiptera: Coccidae: Coccus L.) associated with Crematogaster ants (Hymenoptera: Formicidae) in the stems of Macaranga plants (Euphorbiaceae) in Southeast Asia.
PENNY J. GULLAN, TAKUMASA KONDO, BRIGITTE FIALA, SWEE-PECK QUEK.

http://mapress.com/j/zt/articl…ad/zootaxa.4521.1.1/16573

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.

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!!!

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!

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.

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 …

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.

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.

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!

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.