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Book reviews

Termites: Biology and Pest Management.M. J.Pearce. 1997. XII+ 172 pp., 6 tables, 59 figures, 32plates. CAB International, Wallingford, United King-dom. ISBN 0–85199–130–0 (HB) price £35.00, US$65.00.

A definite niche for an introductory book to the worldof termites exists. Currently, newcomers to the dis-cipline have to consult the two volumes (over 1200pages) of Krishna & Weesner (1969/1970)Biology ofTermitesor the three volumes (over 2000 pages) ofGrassé (1982–1986)Termitologia.For more recent ad-vances on Isoptera, they have to search the scatteredliterature.

In Termites: Biology and Pest Management,Pearce divides the topic into seven chapters: (1) Ter-mites as Insects, including evolution, termite castesand classification; (2) Distribution; (3) Termite Biol-ogy and Behaviour with sections covering communi-cation, feeding, water requirements, defence, foragingand nest building; (4) Nest Systems; (5) Termite Ecol-ogy, including sections such as the effects of soiland vegetation type on termite biology, benefits tothe environment, and predators and parasites; (6) Ter-mites as Pests, giving information on food preferences,damage recognition and detection and damage assess-ment; and (7) Control Methods, detailing chemical,physical, cultural and biological control, safety andfuture trends. Four appendices detail methods andprocedures for collecting, identifying and culturingtermites; for monitoring termite activity and assessingtheir damage to field crops; and for conducting a rangeof laboratory tests (effects of chemicals and biolog-ical control , food preferences, termite resistance ofmaterials). The References are divided into general,taxonomic and others covering biology and control.

The book has a number of valuable features whichbring together information which is not readily acces-sible or would take considerable effort to compile. Forexample, features are a table of all termite genera andgeographical regions in which they occur, world dis-tribution maps for several pest genera of termites anddetailed instructions and descriptions of methods forlaboratory and field experiments with termites (in theappendices).

Unfortunately, there is a definite disjunctive stylein the book. Chapters (1) to (5) dealing with termitebiology appear to have been put together somewhathastily and phrasing is often sloppy and somewhatconfused. Paragraphs frequently have facts strungtogether without apparent linkage, and individualsentences are loaded with significant information insupplementary sentences. Two examples may suffice:

p. 55, under ‘Defence’ the following paragraphoccurs which illustrates the stringing and unusualarrangement of facts: ‘In mounds in the Macrotermiti-nae the royal cell has solid walls with a line of smallexit holes which are easy to defend. Mechanical de-fence (mandibles) and chemical secretions are foundin soldiers of many termites (ref.).’ - Specific situationsare given first, with general facts attached rather thandescribing general features first and adding interestingspecifics.

p. 66 under Nest Systems – Wood-dwelling Ter-mites, confused, sloppy writing and unrelated factsabound: ‘Small colonies living inside a piece ofwood...may never grow beyond a few hundred individ-uals, but some can reach a few thousand. Mastotermescan have nests containing several million individu-als....Coptotermes nests have been recorded as havingseveral million individuals. Wood-dwelling termiteshave fewer competitors than the mound or subter-ranean termites, but as the wood becomes eaten outwith increasing colony size, more predators can en-ter. As with wood-dwelling cockroaches, galleries areparallel with the grain of the wood. Termites canlive entirely within the food source and move to othersources of wood. Drywood termite faeces are dry seed-like pellets that are thrown out of holes made in thewood, which are resealed afterwards. Wood can betotally isolated from the ground in a tree, building oreven a boat or in contact with the soil, where the lowerhalf may be damp. In hot climates termites are foundmainly in wood close to the ground. Wood can be tim-ber or living, which then provides a water source.’– Inthis quote, facts from groups of termites with differentlife histories which are active inside wood are some-what combined, although the main emphasis appearsto be on drywood and dampwood termites, with infor-mation on population sizes of different termite groups

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added in between. Further, this is the only section inthe book where the author deals with the populationsize of termite colonies (and the index fails to lead thereader to this paragraph). Separate paragraphs dealingwith the relevant facts for drywood, dampwood andsubterranean termites and one on colony size couldhave turned this section into very informative text.

In contrast, the chapters (6), (7) and the Ap-pendices covering termite management etc., areas inwhich the author can draw on a wealth of his ownexperience and research, read well and the informationtherein appears reliable.

The arrangement of topics is often unexpected.For example, developmental pathways are covered ina section dealing with ‘Replacement Reproductives’(p. 18). The section on ‘Feeding’ (Chapter 3, Ter-mite Biology and Behaviour) deals, amongst othertopics, with ‘Wood Feeding’ (p. 47/48) and ‘OtherFood Sources’ (p. 48). The latter mentions oddities(fodder, old sacks, discarded shoes etc.), yet the greatdiversity of cellulosic materials different groups oftermites utilise as their staple is only covered in thesection ‘Food Preferences’ (p. 91) in Chapter 6 (Ter-mites as Pests). See also the above example on termitepopulations.

The book contains a number of errors which couldhave been easily avoided, and a number of statementswithout any reference to support them. Here are exam-ples from just a cursory scanning of the text:p. 3, Table 1, has names of some genera misspelt orarranged in the wrong families/subfamilies;p. 20, maximum number of cerci segments in termites:Archotermopsiswith 6–7 – not 5 as inZootermopsisandMastotermes; ‘In Mastotermesthe first soldier hasno teeth’ (presumably mandibular teeth?) – it doeshave them; ‘Compound eyes are present inMastoter-mes, Hodotermes and Anacanthotermes[soldiers]’ –yet for Mastotermesthey are omitted from the illus-tration in Figure 3, further, soldiers in this genus donot have the type of compound eye found in those ofHodotermes.p. 37, the distribution ofCoptotermesin the US isshown incorrectly. – The southern states which havenotorious problems withC. formosanusand fromwhere the invasion started [Louisiana (New Orleans’famous French Quarters is heavily damaged byC.formosanus!), Florida etc.] are shown asCoptotermes-free areas while on the East coast the species extendson the map much further to the North than in reality.p. 38 has an example of the many notable statementswithout giving adequate background information or

references: ‘For termites to survive in cold regionsthey must compensate for loss of population in win-ter by increasing numbers during the warm season.’– Species would require many attributes to survive incold regions. Is Pearce implying that such species havehigher reproduction rates than termites from warmerregions?p. 47, The statement: ‘In the lower termites that haveprotozoa in their gut, salivary glands are lost just be-fore moult but are regained later by trophallaxis.’ isincorrect. The protozoa are lost and regained, not theglands.p. 52 says: ‘Animal dung from wildlife is often eaten bytermites, but dung from domestic animals is not eaten.’– This may apply to Africa, but certainly does nothold true for Australia (and elsewhere?) (for Australia:Hill 1942, Gay 1968, Ferrar & Watson 1972, the latterrecorded well over 40 species from 3 families in cattledung);p. 53 has another incorrect statement: ‘Cannibalism iscommon in termites where the food source is low, andoften appendages are bitten off. In the Kalotermitidae,one may often find wing-pad or leg scars resultingfrom attack by other workers.’ – This behaviour hasa caste regulatory function (Myles 1986) and is notlinked to cannibalism as a result of ‘low food sources’[meaning low in nitrogen?];p. 54 has an odd statement, meaning?: ‘In dry areas itis especially useful for termites to have access to thesoil surrounding soil particles, as this is where watercan be found.’p. 55 says: ‘In Nasutitermes exitiosus large soldiersare present with the foragers while small soldiers arenear the nest.’ – It is the other way around. Smallsoldiers accompany foragers and are also numerousthroughout the nest (mound). Large soldiers are foundonly inside the nest, but not all colonies have them.p. 78 has: ‘However, by opening up gaps one losesgroups such as the soil feeders Coptotermes and Na-sutitermes,...’– These genera are not soil feeders.p. 87 states: ‘Echidnas from western Australia re-ject Drepanotermes and Nasutitermes,..., but eatTrinervitermes.’ – Trinervitermesdoes not occur inAustralia [according to Table 1, p. 6 it is a genus fromthe ethiopian and indomalayan regions.]p. 112 informs that: ‘Waller (1966) has looked at thepossibility of using urea solutions for controlling ter-mites in Australia.’– The work withReticulitermeswas done in the US; this genus does not occur in Aus-tralia.p. 119, Contrary to what is stated, the fungal pathogen

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Metarhiziumhas not yet been registered for termitecontrol in Australia (only limited experimental permitshave been issued).p. 127, Contrary to what is stated,Heterotermescul-tures will produce supplementary reproductives (e.g.Roonwal, 1979).In the References, the mode of citing Chinese authorsis erratic: those cited more than once have their namespelt differently on each occasion.

One certainly can argue how well or otherwise theauthor has managed to condense the knowledge on ter-mites and their management into a small volume: it isa difficult task by any standard. Several of the specialfeatures, notably in the appendices and the last twochapters, as mentioned earlier, clearly present valueand make the book interesting to those with involve-ment in termite management. However, the apparentlack of care in checking numerous facts and the some-what confused writing in many sections on the biologyof termites, render the book into a flawed introduc-tion to the world of termites. It will be necessary forthe reader to seek confirmation on statements in othersources and to turn to older and more voluminous ref-erences and various review papers on specific topics tofully understand the material.

MICHAEL LENZ

CSIRO Entomology,GPO Box 1700,

Canberra ACT 2601, Australia

Soft Scale Insects: Their Biology, Natural Enemiesand Control. Edited by Y. Ben-Dov & C. J. Hodgson1997. Volume 7A (452 pp) and 7B (442 pp.), Elsevier,Amsterdam. ISBN O–444–89303–2 (7A, HB), priceUS $270.00 (NLG 470.00) and ISBN 0–444–82843–5(7B, HB) US $264.50 (NLG 460.00).

This is Volume 7 in the series World Crop Pests(editor-in-chief M. W. Sabelis) devoted to the familyCoccidae. It follows the same well tested pattern andhas the same high standard as the previous volumes. Involume A, morphology, systematics, phylogeny, biol-ogy, physiology, and ecology of coccids are reviewed,while volume B treats first natural enemies and sec-ond damage and control. Together with two otherrecently published monographs by the same two edi-tors, this volume is meant to cover the entire spectrumof knowledge on soft scales.

Volume 7A was written by 25 authors, seven withmultiple contributions, from 12 countries; volume 7Bby 23 authors, six with multiple contributions (onlyfour from 7A), from 10 countries. Overall, authorsfrom 15 countries, from all continents except SouthAmerica, have contributed. The coverage of Coccidae(with about 1100 described species), one out of 21families of the scale insects Coccoidea, is thus trulycomprehensive.

Here follows a short description of the family, asknown from most entomology text books: Coccidsare bisexual but have wide ranging parthenogeneticspecies with diploid arrhenotoky. Females have ahemimetabolous life cycle, while males can be con-sidered holometabolous, having resting pupae andwinged adults. Most are protected by wax. All arephytophagous, mostly on perennial plants, where theyfeed on phloem and produce honeydew. Many arepests, directly through feeding and indirectly throughsooty mould that grows on the honeydew. The hon-eydew is often removed by ants with whom coccidshave a successful symbiosis. Ants protect them againstnatural enemies. Coccids often succeed in encapsulat-ing eggs or larvae of their internal parasitoids. Thesewell known facts are described, sometimes in severalchapters, from different angles and in great detail.

In view of the fact that this is now the third,relatively slim, volume treating similar insects withsimilar natural enemies and pest status (after aphids involume 2 and armored scales in volume 4) and againonly with black and white photographs, does not thismake for boring reading ? The answer is definitivelyno. The further I read, the more I became enamoredby the treatise, which succeeds in giving insights intomorphology, ecology, and integrated pest management(IPM) through the prism of this one family.

The first 250 pages are devoted to morphology ofthe different stages, which may look old-fashioned,but is extremely useful. They cover the adult fe-male (D. Matile-Ferrero), adult male (J. H. Giliomee),and immature stages (M. L. Williams). Taxonomy(C. J. Hodgson) is largely based on young females,which can often be found only during a short timeof the year. Adult males (J. H. Giliomee), thoughfascinating creatures, are less useful for taxonomy be-cause they are highly variable and few characters areavailable to distinguish between different taxa. Thetaxonomy of immature stages (M. L. Williams andG. S. Hodges), mostly first instars, is – as the au-thors say – still in its infancy. ‘The male test’ (G. L.Miller and M. L. Williams) turns out to be a mor-

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phological structure, a protective covering useful fortaxonomy. The female test (Y. Tamaki), which can at-tain 98% of the total weight of the insect, is composedof waxy material (aliphatic fatty acids and alcohols,with unique cyclic compounds inCeroplastesspp.)but also aqueous components, which are also called‘interior honeydew’.

The resulting classification of the Coccidae and re-lated coccoid families (C. J. Hodgson) shows a variedand tortuous history of groupings of families related tococcids. Coccidae are then separated into 10 subfami-lies. Taxonomy is difficult partly because intraspecificvariation of taxonomic characters (E. M. Danzig) ishigh, which is attributed to the wide geographic distri-bution and polyphagy of these insects. Moreover, mor-phology is influenced by parthenogenesis, seasons,and the presence of parasitoids.

Zoogeographical considerations (F. Kozar, Y. Ben-Dov) deal with the question as to how many speciesthe coccids comprise. The answer depends on howyou count what you do not know. One intriguingapproach: Among the conspicuous and well studieddanaid and papilionid butterflies, 1% of all specieswere described for the first time between 1960 and1980. For the Coccidae the same statistics is about13%. In fact, in the last 100 years, a linear steepincrease in number of recognized genera has been reg-istered, with no sign of flattening of the curve. InEurope, where this family has been relatively wellstudied, it is particularly rich in species living at thebase of Cyperaceae and Poaceae, and the roots of otherplants. In other regions, species from this niche aremuch less common, presumably because they have notbeen collected sufficiently. On the basis of these con-siderations and despite the fact that more species andgenera are known from the Palaearctic region, Cocci-dae are thought to have their centre of diversity andtheir origin in the tropics. A small group of generaoccurs in more than three zoogeographical regions,but even in this cosmopolitan group, only 73 speciesare distributed on more than one continent. Cladis-tic analysis leads to a phylogeny (D. R. Miller, C. J.Hodgson) with 10 recognized subfamilies.

In the section on biology, the general life historycharacteristics (S. Marotta), including the embryonicdevelopment (E. Tremblay), are presented. We learnthat the ‘Krassilstschick cell’ of diaspidids has nowbeen discovered in coccids. At least, this gives us newmaterial for crossword puzzles.

Endosymbionts (E. Tremblay) have been knownsince 1854, their hereditary transmission through the

egg has been described in 1880, and they were recog-nized as symbionts in 1907. These organisms, whichoccur in the haemolymph and normal or polyploidizedfat cells have, however, a weak association with theirhost, to whom they impart as yet unknown benefits.

For the tinkerer and lover of mechanical gadets,the chapter on morphology and anatomy of honeydeweliminating organs (C. P. Malumphy) describes an as-tonishing mechanism. Unlike other families, coccidshave a complex anal apparatus to achieve the coatingand disposal of droplets. We also find a useful def-inition of honeydew involving the notions of sugarydeposits as a liquid elimination of a waste product byphloem feeders. Sooty moulds (R. K. Mibey), whichgrow on this honeydew, are taxonomically separatefrom the black mildews and phytopathogenic fungi.Sooty moulds are often mixtures of species, whichmay be pleiomorphic. Five distinct families, all withinthe order of Dothideales, are recognized as beingspecific to this niche.

In a few instances, soft scales are known as ben-eficial insects. Honeydew offers forage for honeyproduction (H. Kunkel), particularly in regions withlarge forests with dry summers and/or where weedshave been decimated. As a sales pitch for honey, fla-grant flowers are better than the reality, i.e., faecesof unknown insects. In China, the Pela wax scale isused for commercial wax production (T-K. Qin). Forover a 1000 years, wax produced by male nymphsof Ericerus pelagrown mainly on ash and privet hasbeen collected. The chapter describes in loving de-tail the involved, labour intensive rearing of the malesand females in separate fields. We get a glimpse intothe candle production in old China and learn of themodern use of this wax in ‘high tech’ industries.

In at least one case, coccids have an indirect effecton their host plant (J.A. Vranjic) through a pathogen.The beech scaleCryptococcus fagisugatransmits thepathogenic fungusNectria coccinea. Since 1890, thedate of the accidental introduction of this scale intoNorth America, 85% of all beech trees on the continentdied from the resulting disease. Gall formation (J. W.Beardsley) is known from only one coccid, the AfricanCissococcus fulleri, which produces galls on the vineCissus.

Knowledge of crawler behaviour and dispersal (D.J. Greathead) and of the seasonal history and diapause(S. Marotta & A. Tranfaglia) are essential to under-standing the population dynamics of these insects. Ifound the description of the symbiotic relationshipswith ants (P. J. Gullan) particularly fascinating. It in-

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volves the coccid, parasitoids and predators, otherphytophagous competitors including large mammals(including you, if you get in the way of an ant nest),and the host plant, which sometimes offers extra shel-ter in so-called domatia. The outcome of these inter-actions depends on the ant species and has often beendemonstrated in beautiful experiments. Encapsulationof parasitoids (D. Blumberg) is the scale insect’s wayto overcome these natural enemies. Most informationis, however, from only four species of coccids andinvolves mainlyMetaphycusspp. (Encyrtidae). Inter-estingly, the effectiveness of encapsulation sometimesdepends on host plant.

Lest we get carried away with these fascinatingglimpses in the life of coccids, the chapters on tech-niques for collecting and mounting (Y. Ben-Dov & C.J. Hodgson) and laboratory and mass rearing (M. Rose& S. Stauffer) remind us of the technical basis neededto investigate these creatures.

In the second volume, part two deals with thenatural enemies of scale insects in almost the samedetail as was given to the host. The taxonomy of ento-mopathogenic fungi (H. C. Evans & L. Hywel-Jones)has mostly been worked out by plant pathologists andmost literature is>70 years old. The entomopatho-genic nature of such fungi was only discovered in 1894and, most often, the insect host was not identified byan entomologist. Moreover, entomopathogenic fungiare sometimes overgrown by mycoparasitic fungi. Itis therefore not astonishing that the field needs muchmore work and only a few attempts at biologicalcontrol have been made.

By contrast, Coccinellidae (D. J. Ponsonby & M.J. W. Copland), with about 4200 species, are wellknown predators of coccids, as described in this ex-haustive review. Interestingly, male-killingRickettsia,which are not contagious, were discovered a long timeago, but their benefit to ladybirds is still an enigma.The authors conclude that none of the coccid-feedingcoccinellids so far investigated appears to have all theattributes deemed desirable for successful biologicalcontrol. Some coccinellids do, however, assist biolog-ical control of coccids when given the opportunity tofeed also on other hosts and/or when assisted by otherbiological control agents. Although Nitidulidae (Cy-bocephalinae) are host specific and good dispersers,they are considered as less effective than coccinellids.

Among the Cecidomyiidae (K. M. Harris) thereare specialized as well as opportunistic feeders. Lit-tle is known about those on coccids, most commonlyDiadiplosis coccidivora. While interest in coccid-

feeding cecidomyids seems to be declining,Aphido-letes aphidimyzais now used commercially againstaphids.

About 830 spp. in 45 genera of Encyrtidae (G. L.Prinsloo) are internal parasitoids of soft scales, halfof them primary, half secondary (with the exceptionof someMicroterys spp., which are egg predators).The morphology of the larvae and pupae and theirspecialisations to attach to the tracheal system of thehost are described. A key to genera and useful tax-onomic notes are given. Encyrtids attack some 250soft scale species. The most serious pests also havethe largest recorded spectrum of encyrtid parasitoids,but only a small proportion of encyrtid species areused in biological control programmes. Aphelinidae(M. Hayat) are the most successful family in bio-logical control, but mostly on diaspidid scales. Oncoccids, only nine genera are recorded, including fourprimary parasitoids and five hyperparasites. Again, auseful key to the genera and notes, including a tablewith doubtful and unusual records, are given. OtherChalcidoidea are less important parasitoids of coccids.Eulophidae, Pteromalidae, Eupelmidae, Signiphoridae(G. Viggiani) are briefly mentioned. It is sobering tolearn that the biology of most species of the ubiquitousgenusTetrastichus, which includes both primary andsecondary parasitoids, is unknown.

In Part 3, covering damage and control, the eco-nomic importance (R. J. Gill & M. Kosztarab) of coc-cids is described superficially and qualitatively only,which might well reflect the status of knowledge.

After a description of insect development and re-production disrupters (B. Darvas), biological controlof soft scale insects in interior plantscapes in the USA(S. Stauffer & M. Rose) gives an interesting, holis-tic picture. The mentioned landscapes are plant col-lections, botanical gardens, zoological parks, modelecosystems, and plantings that beautify office build-ings and shopping malls, i.e., the daily ecosystemsof urban populations. Perennial plants, polycultures,planted for comfort of humans, with humidity, light,and temperature levels all adjusted downward, withoutmechanization, and high pest tolerance characterizethese systems and distinguish them from the shortlived glasshouse crops. Natural defense mechanismsof plant species are maintained and public concernabout pesticides should eventually lead to reduced in-secticide use, a status which does not yet seem to havebeen attained. A wide number of insect species inhabita great variety of microhabitats. Sedentary species arefavoured and it is therefore not astonishing that soft

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scales constitute an inordinate proportion (about 1/4to 1/3) of all pest problems. Until recently, relianceon the use of pesticides has excluded biological con-trol, but the authors are enthusiastic about prospects,citing the need for better education of the managers ofthese plantscapes. This unorthodox chapter gives a lotof practical and well founded advice.

Volume B closes with a series of rather conven-tional chapters on coccid pests of important crops,which review information given in previous chaptersin a new light. This section covers citrus (R. J. Gill),olive (G. Pellizzari), avocado, mango, guava, persim-mon, and other tropical fruits (E. Swirski, M. Wysoki,Y. Ben-Dov in various combinations), deciduous fruittrees (D. G. Pfeiffer), grapevine (G. Pellizzari), sugar-cane and bamboo (A. J. M. Carnegie), coniferous anddeciduous forest trees, ornamentals and house plants(each by M. Kosztarab), coffee (S. T. Murphy), cocoa(C. A.M. Campbell), tea (D. Greathead), and coconutand rubber (each by T. H. Chua).

In summary, this volume demonstrates the stateof the art in coccid research. The chapters on con-trol reflect very much the status of IPM in general,whose implementation is often hampered by the lackof basic knowledge. This highlights the amount of re-search still needed to arrive at a full understanding anda satisfactory management of these fascinating scaleinsects.

The authors are to be congratulated for this work.Like its predecessors, this volume will form an in-dispensable part of all entomological libraries and,despite its hefty price, many a private entomologistwill decide to buy it for a leisurely reading at home.

PETER NEUENSCHWANDER

Plant Health Management Division,International Institute of Tropical Agriculture,

B.P. 08 0932, Cotonou, Benin (West Africa)

Pheromone Communication in Social Insects.Edited by R. K. Vander Meer, M.D. Breed, M.L. Win-ston & K.E. Espelie, 1998. 368 pp. Westview Press,Boulder. ISBN 0–8133–8976–3 (HB). US $75.00,£57.95, Can. $107.00.

This book is the result of two symposia held at theannual meeting of the Entomological Society of Amer-ica in 1994. Its chapters have been written by thespeakers at these symposia, and some chapters byother authors have been added to obtain a wider cov-

erage of the field. The book is divided into four parts:Sources and Secretions, Nestmate recognition, Re-leaser pheromones, and Primer pheromones of thesocial insects.

Although communication is part of the title, it isonly treated in its proximate sense. There is no reflec-tion in this book concerning the changes that have oc-curred in the last 10–15 years in our understanding ofcommunication in general, such as that signals need tobe honest (Zahavi, 1986), and, connected to this, thatpheromones are not means to suppress behaviouraland physiological processes in nestmates but insteadserve for mutual adjustments (Keller & Nonacs, 1993).For proximate analyses, such distinctions are indeedless useful; however, for understanding evolutionaryprocesses related to the emergence of sociality, andits rather complex forms we witness nowadays, thesedistinctions are fundamental.

Related to this is the distinction made in this bookbetween releaser and primer pheromones. The dis-tinction goes back to Wilson & Bossert (1963). Theydefined releaser pheromones as those substances thatevoke immediate behavioural responses, while primerpheromones were those substances that lead to phys-iological effects in the receiver. This distinction wasbased on the idea that primer pheromones would actdirectly on, for instance, endocrinological regulatoryprocesses, while releaser pheromones should be per-ceived through sense organs. This distinction provedto be incorrect, and later this separation was changedinto releaser and primer effects (see Hölldobler &Wilson, 1990), making it possible that one substancecould lead to both types of response. These differenteffects could be the consequence of different routes forprocessing the chemical stimulus perceived throughsense organs. This mode of action is discussed in thevery readable chapter by Vargo (Primer pheromonesin ants).

Primer pheromones in the original sense probablyoccur in the termites, where caste determination de-pends on substances, such as juvenile hormone, whichare passed around in the colony. The idea stems fromLüscher (see his 1974 paper) and is now supportedby experimental evidence. It is described in anotherstimulating chapter, this time by Henderson (Primerpheromones and possible soldier caste influence on theevolution of sociality in lower termites).

Also the introductory chapter (by Billen & Mor-gan: Pheromone communication in social insects:sources and secretions) has in part a general valuein that it discusses principles that go beyond the ex-

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amples treated. Several other chapters, however, havethe characteristics of encyclopedia: a number of sub-stances and specific effects have been found and arelisted. Such chapters are probably quite useful for thestudent who wants to have full details in a narrow,specific area, but are less suitable for a larger reader-ship. It is the consequence and probably unavoidableeffect, of having many authors contributing to such abook, some of them successful in giving an overview,others in reporting on the achievements of their ownresearch group. Together, these chapters give an ex-tensive overview of the field of pheromone research inthe social insects. Undoubtedly, there is something toadmire in the book for every sort of reader, dependingon their interest in this aspect of sociality.

References

Hölldobler, B. & E. O. Wilson, 1990. The Ants. Springer-Verlag,Berlin, 732 pp.

Keller, L. & P. Nonacs, 1993. The role of queen pheromones insocial insects: queen control or queen signal? Animal Behaviour45: 787–794.

Lüscher, M., 1974. Kasten und Kastendifferenzierung bei niederenTermiten. In: G. H. Schmidt (ed.), Sozialpolymorphismus beiInsekten. Wissenschaftliche Verlagsanst., Stuttgart, pp. 694–739.

Wilson, E. O. & W. H. Bossert, 1963. Chemical communicationamong animals. Recent Progress in Hormone Research 19: 673–716.

Zahavi, A., 1986. Reliability in signalling motivation. Behavioraland Brain Sciences 9: 741–742.

HAYO H. W. VELTHUIS

Comparative Physiology,Ethology & Socio-ecology Group,

University of Utrecht, Padualaan 14,3508 TB Utrecht,The Netherlands