REVIEWS EDITED BY M. ROSS LEIN Thefollowing reviews express the opinions of theindividual reviewers regarding the strengths, weaknesses, and value of thebooks they review. As such,they are subjective evaluations and do not necessarily reflect the opinions of theeditors or any officialpolicy of theA.O.U.--Eds. The Florida Scrub Jay: demographyof a cooper- geneticparents in the care of young that are not off- ative-breeding bird.--Glen E. Woolfenden and John spring of the helpers.To establishthat an individual W. Fitzpatrick. 1984. Princeton, New Jersey,Prince- is a helper, one must observeit caring for young that ton University Press. xiv + 406 pp., 1 color plate, are known not to be its own. There is no evidence many figures.ISBN 0-691-08366-5(cloth), 0-691-08367-3 in the chapter on procedures that Woolfenden and (paper).Cloth, $45.00;paper, $14.50.--For nearly two Fitzpatrick employed this criterion, although they decades three long-term studies of communally were aware of it (p. 4). Instead, throughout the book breeding birds, well known to readersof this review, jays are divided into breedersand helpers, implying have been in progress.The behavioral ecology of that if a bird is not a breeder it must be a helper these speciesis so complex that conclusionsreached ("Helpers are nonbreeders," p. 80). It is well known after only a few years of study can be quite mislead- for other speciesthat not all nonbreeders help, and ing. Eachnew year of studynot only enlargessample that individual nonbreedersvary significantlyin the sizesbut also provides insights that require impor- amount or intensity of their helping efforts. These tant reinterpretationsof earlier findings. Commend- are also possibilitiesfor the Scrub Jay. To remain ably, Woolfenden and Fitzpatrickbrought together strictly objectiveI recommendthat readerssubstitute in one convenient place some of the findings of the in their minds the term "nonbreeder"every time they first 10 yearsof their study of ScrubJays, demonstrat- seethe word "helper" in this monographbecause the ing many benefitsof a long-termstudy. Although it operationalcriterion used in identifying presumed presentsa wealth of information, however, this book helpersseems to have been that they were not breed- fails short of accuratelyrepresenting the prevalent ers. views of 1984 (and especially 1987) with respectto The failure to distinguish clearly and consistently the main issues.To understandwhy, I proposeto between the question of helping and that of non- examine the main issuescritically. breeding can have far-reachingconsequences. It may One of the major contributionsof evolutionarybi- be one reason the authors and others have confused ology in the secondhalf of this century is, in the the main issues.The question, "Why help?" is dis- opinion of many, the theory of inclusive fitness tinctly different from the question, "Why delay (Hamilton 1964). By generating testable predictions, breedingand dispersal?"Yet in this monographthese this theory hasstimulated many empiricalstudies of two questionshave been confounded.Of course,the natural populations. The focus of interest is the rel- two questionsare not independent. For example, the ative importanceof indirect fitness (Brown 1980) in hypothesisthat the authorsreject may be stated as the evolution of socialsystems. follows: Helpers give up chancesto breed to gain One approachto this issueis to ask whether indi- indirect benefitsfrom helping their parents.No oth- rect fitnessis really necessaryas a componentof an er benefit to the helper is considered except the in- explanation of social behavior, such as helping be- direct benefit. A secondhypothesis involving indi- havior. This view is commonlyassociated with Wool- rect selection is that some jays are prevented from fenden and Fitzpatrick (p. 6). Another approach, breeding by territoriality and the resultant habitat which I prefer, is to use the methodof alternativework- saturationand that only then, having no opportunity inghypotheses, traditionally viewed as the heart of the to breed, do the birds becomehelpers for the indirect scientific method. Using this procedure we set up benefit. A rule of thumb for suchbirds might be stat- alternativehypotheses, some employing indirect fit- ed as:"Breed if you can;but if you cannot,then stay nessand some not. Then the data are comparedwith home as a strategy for getting a territory and help the predictionsof eachhypothesis. Some hypotheses your parentsin the meantime."This is the basisfor will be rejected;others will remain. The "is it nec- the theory I proposedin 1969 and 1974. Needlessto essary?"approach, if it is used at all, should be ap- say, I was dismayedthat although I proposedthe plied to all hypotheses,not selectivelyto thosedis- theory specificallyfor jays and had the Scrub Jay in approvedby the authors. mind, the authors did not even consider it. Beforethe processcan even begin, we must know Koenig, Mumme, and Pitelka (1983) extended the exactlywhat we are talking about. What preciselyis tendency to think of helpers as nonbreederseven a helper? A helper is an individual that assiststhe further: they mistakenly defined helpers as non- 35O April 1987] Reviews 351 breeders.Quite possiblythe rather widespreadoc- authors,however, imply that their observationsare currenceof suchmistakes has contributed to the per- not"consistent with strictly kin-selectedhelping be- sistentconfusion of issuesby these five authors. havior" (p. 88). It would have been interestingto see After nine chaptersof competentpresentation and the data on actual helping at the nest by the unre- analysis of data, Woolfenden and Fitzpatrick distill lated 6%,compared with the correspondingdata for the main resultsin a final chapter that evaluatesthe the related 94%. These data were not presented. In data with respectto the question,"Why help?" In my opinion, the data in this monograph are com- their casethis means mainly, "Why stay home and pletely consistentwith the theory I proposedin 1974. delay breeding?" They conclude that the factor of Further discussionof this point may be found in greatestimportance for a young bird is the difficulty Brown (1987). of obtaininga breedingterritory. In other words,the Are the data also consistent with othertheories, not jaysdelay breeding("help") simply becauseit is dif- involving indirect selection?There is some discus- ficult to find a breedingopening. This is exactlythe sion of the possibilitythat somemales that help can conclusionI reached(1978a) after a review of quan- improve their chancesof gaining breeding status titative data on various species,including the Scrub through territorial budding. Personally, I find the Jay. Indeed, the authors postulatea variable, reflect- data unconvincing (see Brown 1987); but I am will- ing this difficulty,D, which correspondsto my vari- ing to allow, lacking the critical data, that interpre- able F, which correspondsto Emlen's(1982) variable tationscould go either way. The experimentsneces- psi. It appearsthat on this point Woolfenden, Fitz- sary to resolve the issuehave not been done. patrick, Emlen, and I are in perfect agreement.This For females,on the other hand, no plausibletheory monograph,however, gives the impressionof dis- is presented by the authors. Furthermore, the data agreement.The authorsrefer to their "decouplingof are not consistentwith any publishedtheory for the kinship benefitsand dispersal"as "our major depar- evolution of helping that does not invoke indirect ture from typical kin-selectionmodels," and cite me selection. For further discussion,see Brown (1987: incorrectly(p. 324). This uncouplingwas, of course, chapter 14). the majorpoint of the K-selectionphase of my 3-phase In my opinion the confusionin this monographis theory, which "uncoupled"the explanationfor de- the inevitableresult of failure to distinguishcareful- layed breedingand dispersal(habitat saturation) from ly and consistentlybetween two questions.First (Q1), kinship benefits. Why do young remain with their parentsin a non- Despite their hidden agreement,I am displeased breeding role? Second(Q2), Given this role, why do with their conclusion, not for what it is but for how they help? they arrived at it. Ideally, the method of multiple The loudest claims that indirect kin selection is working hypothesesshould have been used formally unimportant (i.e. this monograph,Koenig and Pi- and explicitly. All hypothesesfor delayed breeding telka 1981) are based on Q1. But for Q1, where is the shouldhave been laid on the table.An effort to reject theoretical and empirical literature that invokes in- each of them should have been made. One of the direct kin selectionformally and explicitly?If this is oldest known hypotheses for delayed breeding, an important scientific issue,such literature should namely age-dependentlack of skill in foraging,was not be hard to find. I have searched in vain for such not even considered.Perhaps in a future publication papers before 1984. My own papers have never in- they can present data with which the remaining hy- voked indirect kin selection for Q1, and I have ex- pothesescan be evaluated. plicitly rejectedthis hypothesison the basisof a re- Now what aboutthe main question,the evolution view of empirical data (Brown 1978a). Woolfenden of helping in the true sense?The authorshave little and Fitzpatrick failed to find such references;and to say on this subjectexcept for a few commentson their earlier (1978) reference to Brown (1974) was an the possibilitythat helpingbehavior is a neutraltrait.