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Of the Books They Review. As Such, They Are Subjective Evaluations and Do REVIEWS EDITED BY WILLIAM E. SOUTHERN 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 officalpolicy of the A.O.U. Eds. The Shelduck: A study in behavioural ecology.-- for explaining many of the variablesin behavior and, I. J. Patterson. 1982. Cambridge, Cambridge Univer- indeed, is the main theme related to population lim- sity Press.x + 276 pp., 10 black-and-white drawings, itation that recursin discussionsthroughout the book. 101 figures, 68 tables. $49.50.--Traditionally, ethol- While this certainly seemsa proper emphasis,the ogistshave emphasizedstudies of causation,ontog- lack of precisedata on body weight in terms of pro- eny, evolution, and physiologicalbases of behavior, tein and lipids and the confounding variablesrelated often without concern for ecologicalvariables (e.g. to whole body weights (e.g. trap bias, gut contents, food supply) or relationships to population density. and prolonged breeding season so that individuals Similarly, population ecologists have emphasized are at different stagesof the reproductivecycle) re- numerical values in relation to ecological variables suit in a large amount of logical speculationand con- without data on variability of individuals or social clusionswithout critical data. Strengthsof the treat- behavior. Recent attempts to integrate ethology, ment are data on birds of known age and data on ecology,and population biology with the theory of food supply and use. Chapter 2 concludeswith a natural selectionhave produced a new field that is rather lengthy treatment of world distribution of now commonly termed Behavioral Ecology (e.g.J.R. Shelducksand the molt migration (pp. 17-39). Much Krebs and N. D. Davies (Eds.) 1978. Behavioral ecol- of this struck me as distracting detail, given the main ogy. Oxford, Blackwell Scientific Publications).This theme and strength of the book. integration providesa more full understandingof an Thereafter, the book proceeds in chapter form animal'sbehavior based upon knowledgeof costsand through the annual cycle. Chapter 3 treats gregari- benefits in terms of fitness of alternative modes of ousness in the winter flock. This section contains some action and organization in relation to population excellent data on arrival of Shelducks on breeding density and ecologicalvariables. Testing of ideas in estuariesin relation to age and sex (oldest and males BehavioralEcology will be assistedby detailed field first, 4+ yr olds preceding 3-yr olds, which precede studies using individually identifiable animals. 2-yr olds, which precedeyearlings), the common re- The goal of Patterson'sbook is to usedata collected pairing of previouslymated birds, spatial organiza- in a seriesof individual studiesby himself and sev- tion, the preponderanceof feeding activity in winter eral colleagues,dating back to 1960, on the ecology (February--63% of time), and the replacementof this and behavior of the Common Shelduck (Tadorna ta- activity with agonistic interactions and other behav- dorna)to addressseveral issues of behavioralecology, iors as the breeding seasonapproaches. Brief descrip- especiallythe effectsof socialbehavior on the limi- tions of displays are followed by much detail on tation of population size. Shelduckswere considered dominance(most of which waspreviously published excellentsubjects for studybecause of their relatively in journal format,but is neededhere for the integrity large size, conspicuousappearance, varied socialbe- of the book). Clear dominance hierarchies were evi- havior, and existence in small and accessible local dent that were related especially to pair-bonding, populationsin the estuariesof Great Britain. suggesting that females might stimulate aggres- Following a brief (Chapter 1:4 pp.) introduction sion-or prefer the more dominant males. Interest- of the subjectof behavioralecology, the author pro- ingly, dominance did not relate consistently to ad- ceedsin Chapter 2 with a treatment of the taxonomic vantage in obtaining territories, nor to successin positionand descriptionof Shelducks,which exhibit rearing broods,nor to survival. Facedwith these re- many goose-likecharacteristics (e.g. lack of conspic- suits, Pattersonsuggests that maintenanceof pair- uous sexual dimorphism, prolonged parental care, bonds may have been the main selection force for persistentpair bonds)combined with morphological dominant, aggressivebehavior, while leaving open and molt featuresthat are duck-like. The compari- the possibility that breeding population size might sonsend there, however, as there is virtually no fur- be regulatedby dominanceeven though this was only ther referenceto other waterfowl (or pertinent lit- weakly, if at all, supportedby the evidence. erature) thereafter. Included in the description is a Chapter 4 deals with the territory. Much interest- characterizationof body-weight changesfrom mid- ing informationis presentedhere, especiallybecause winter through the breeding season. These data, densities of food items were measured. Surviving however, are lumped averagesof weights of birds pairs tended to occupythe sameterritory (80%),while capturedin eachmonth. Energy is the currencyused males with new mates tended to follow females to 759 760 Reviews [Auk, Vol. 100 their old territories. Territory size did not decrease female) probably promoted brood-mixing as well as with an increasein food supply and ducks seemed the tendency for young to move away from adults to settle only where suitablefood was present.Ter- and follow other pairs. There was no consistency ritories are of further interest in Shelducks because among years on which adults accompaniedcreches they do not containthe nest site nor are they the site other than that they were themselvesbirds which of pair formation or brood rearing. They are, instead, had hatchedyoung. The significanceof brood-mix- an area in which the female can feed intensely with- ing still needs study. Chapter 8 continues the dis- out being disturbedby conspecifics(71% of her time cussionwith respect to duckling survival. Approxi- during the pre-egg-layingphase and 80-90% during mately35% of the young fledged,but this washighly incubation recesses). While these data are convinc- variable (9-66%). Most losses occurred in the first ing, it certainly would be nice to have more infor- week of life and were inversely related to density mation on the role of stored reservesversus daily (although not consistentlyso). There was some evi- intake to meet the energy demandsof reproduction. dencethat a lower proportion of ducklings in mixed Even though Pattersonconcludes that territorial be- broodssurvived than ducklingsin single broodsand havior has been selectedfor becauseit provides an that lack of brooding during inclement weather was undisturbed feeding area for the female, he admits a contributing factor. A discussionof costsand ben- that territoriesmay function in limiting density in efits of single versus mixed broods in terms of risk local populations, but notes that this aspect needs to predation, interaction with weather, and freedom more quantitative study. for adults to feed and migrate points out possibilities Chapter5 detailsShelduck behavior associated with deservingof researchbut doesnot answer the ques- searchingfor nest sites,which were usuallyburrows tions in a satisfactorymanner. Of interest here were in sand dunesaway from the estuarinemud-flat ter- data revealing that "donor" parents were dominant ritories. Shelducksare again unusual in that they are over "recipients" in 12 of 14 cases.There were no semi-gregariousduring this activity and are attracted differences in survival of adults that hatched broods, to other birds on the ground in the nesting area just fledged broods,or did not hatch broods.After duck- a relatively short distancefrom the territory, in which lings fledged,adults departedon the molt migration, they are highly defensive(although agonismis evi- with males often leaving first. dent when two or more pairs come together in the The annual cycle is brought full circle with a dis- nesting area, they clearly are attractedto eachother). cussion of recruitment and the nonterritorial flock in Discussion of costs and benefits of this behavior tends Chapter 9. This section highlights implications for to be teleologically phrased and indicates the need the study of gene flow and the complexityof social for more study (antipredator advantagesrelated to organizationthat can only be observedin wild pop- vigilance and/or learning of the location of "good" ulationswith marked individuals.Although therewas nest sites). Because Shelduck nest sites were so dif- unity in the sex ratio in the population, only 26% of ficult to find and the birds tended to desert easily, returning yearlings were male. Acquisition of terri- this aspectof the studywas frustratinglydifficult and, tories by age classeswas as follows: none as year- correspondingly,weak in comparisonto other data. lings, hAat 2 yr of age, • at 3 yr of age, and all that Egg laying and incubationare treated in Chapter were going to have territories did so by age 4; ap- 6. Clutches were initiated over an extended period proximately20% of older birds were not territorial (mid-April to mid-June) and there was no evidence in any given year. From there the discussionof em- of renesting. Again, teleologicalphrasing is evident igration goeson in much
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