Comparative Study of Todies (Todidae): with Emphasis on the Puerto Rican Tody, Todus Mexicanus.--Angela K
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REVIEWS EDITED BY WALTER BOCK Comparative study of Todies (Todidae): with emphasis on the Puerto Rican Tody, Todus mexicanus.--Angela K. Kepler. 1977. Cambridge, Nuttall Ornithological Club Publ. No. 16. xiii + 190 pp. $11.75.--With the publication of this monograph, knowledge of another little-known tropical group has advanced a major step forward. Following a brief introductory chapter detailing study areas, methods, and external morphology, 14 chaptersoutline the behavior, ecology,breeding biology, distri- bution, and evolutionof the endemicGreater Antillean family. Most of the monographis concernedwith T. mexicanusin rainforestand scrub forest on Puerto Rico, but data on other speciesprovide interesting comparative insights. Kepler provides much new information and overturns a number of misconceptions about todies. Jamaica, Cuba, and Puerto Rico each have one endemic tody, while two occur in Hispaniola. As is commonin island birds, all speciesoccupy a wide range of habitats. The Jamaican speciesseems to be limited by an island-wide limestone formation which restrictssites for constructionof nest burrows. The two Hispaniolan speciesare broadly sympatricnow but apparently differentiated in isolationwhen the islandwas divided by a deep oceanictrough. They are sympatricover a 1265-maltitudinal rangeon one mountain. Todus angustirostris predominates at higher elevations while subulatus is more abundant in the lowlands. Tody behavior is the central theme of several chapters. Vocalizationsare described(including sono- grams) and their functions are discussed.Todies lack complex vocal repertoiresand thus depend on plumagedisplays and wing-rattling in both courtshipand territorial aggression.Wing-rattling is produced as air passesrapidly over the outer primaries. Among the five speciesintensity of flank display is pro- portionalto the amount of pink colorationin the flank feathers.At the extremes,mexicanus has no pink and no flank displayswhile subulatushas extensivepink flanks that are exaggeratedby puffing up the feathers and lifting the wings. A short chapterdiscusses maintenance behavior such as bathing, preening,scratching, and locomotion. Average flight distance increases(with one exception)with wing chord even in these relatively small short-winged birds. The unusually long wing of T. angustirostrismay be attributable to character di- vergenceas it flies more regularly and sits for shorter periods when sympatricwith subulatus. Todies tend to remain paired and occupy the same home range throughout the year. During the breedingseason, territories are somewhatsmaller than non-breedingseason home ranges.Territory size and population density varies with habitat but territory volume varies less among habitats than does territory area. Todies spend an unusual proportion of their time (up to 14 h/day) foraging. They are exclusively insectivorousand the most common foraging technique involves short flights to pick insectsfrom the undersideof leaves. Feeding rates and foraging efficienciesvary with habitat; both are highestin scrub. When sympatric, the Hispaniolan speciesexhibit character displacementrelative to their foraging in allopatry on the same island. I suspectthat todies are most similar ecologicallyto mainland tyrannids such as spadebills(Platyrinchus) or the somewhatlarger flatbills (Rhynchocyclus). As in most coraciforms,both sexesparticipate in the constructionof nest burrows. On Puerto Rico problemsof burrow constructionand maintenance vary among habitats and climates, apparently limiting occupationof someareas. Although stream banks are usedas nestsites in dry scrubland,they are rarely used in rainforestwhere floodingis more common. Todies lay relatively large eggs(26% of body weight) and spendless than 25% of their time incubating. Attentive periods vary from 2 min early in incubation to 16 min late in incubation (average 13.1 min). Incubation lasts 21 to 22 days. Unlike most birds, feeding rates of nestlingsincrease continually up to fledging. Young are fed insects,which increasein size through the nestlingperiod. Nestling mortality is high in unusuallyrainy periodsbecause foraging by adults is restricted.The introducedmongoose is a major factor in nest losses,especially in rainforest. Breedingis distinctlyseasonal in all habitats and more extendedin rainforestthan in scrubland.Yearly variation in rainfall period causesdisplacement of breedingseasons; drought in 1970 acceleratedbreeding in rainforest and delayed it in scrubland. One of the most interestingchapters deals with nest helpers.Many birds (jays, wrens)that have nest helpers have complex social systems,but this is not true for todies. Although the evidenceis indirect, tody helpersseem unrelated to the nest owners. Generally helpers occurlate in incubation or during the nestling period. Although the number of fledged young per adult is lower in helped neststhan in nests without helpers, both clutch sizes and fledglingsper nest are higher in helped nests. Helpers during incubationhave lesseffect on fledglingsuccess than helpersduring the nestlingperiod. 430 April 1978] Reviews 43 1 The value of studiesof single taxa is clearest in Chapters 14 and 15 on population regulation and evolutionin todies.Kepler integratesher ecologicaland behavioraldata to provide a more completeview of tody biology. Discussionof habitat requirements, weather, and insect densitiesprovides insights into selectionpressures on todies. Kepler concludesthat the most important factorsregulating mexicanus populationsin rainforestseem to be predationand food supply as affected by climate. In scrubland, nestingsites and food supply (interspecificcompetition) are most important. Evidence for the importance of interspecificcompetition is indirect at best. Kepler often interprets her results in the context of recent theoretical advances, but fails to conduct rigoroustests of hypotheses,including statistical analysis of results.Further, tabular material is inade- quate to allow suchtests. Citations to a more detailed dissertationand to manuscriptsin preparationare distracting becausethey are not generally available. Technically there are few typographicalerrors but somemaps are poorlydrawn and/orreproduced. Printing of tablesfrom originaltypescript presumably reducespublication costs without detractingfrom the monograph. This study demonstratesthe value of studiesof singletaxa, especiallywhen they go beyond the more anecdotalefforts of past decades.The contributionsof suchstudies to ornithologicalscience will continue to be substantial,especially when they includeintegration of quantitative data on a variety of aspectsof avian biology.--J^MEs R. K•RR. Ecology and energetics of contrasting social systems in Phainopepla nitens (Aves: Ptil- ogonatidae).--Glenn E. Walsberg. 1977. Berkeley, Calif., University of California Press. 63 pp. $4.50.--This researcheffort comparesthe breedingsocial systems in the Phainopeplabut is an excellent model for the approach of natural history studiesby biologists.The Phainopeplais a relatively easy speciesto observeand study, but there are aspectsof its biologythat are perplexingand difficult. It has been an enigmato avian biologistsin the Southwest.In many parts of Arizona and California the birds appear in the fall and establishterritories over areas containingmistletoe clumps, they breed in early spring, rear their young, and then disappearuntil the next fall. Walsberg, with someluck, hard work, and a broad background, has gatheredand synthesizedinfor- mation on the ecology,ethology, and physiologyof the Phainopeplawhere it breedsin one area in early springand then followedwhat he felt was part of the samepopulation to examineits breedinghabits in the summer.The different typesof territorialdefense (A and B) usedby eachpopulation in eachsituation are compared as well as the reasonsfor the evolutionary differencesresolved by the quantitative descrip- tion of the food resource,time of abundance, and availability. The selectiveadvantage of each social systemis examined by comparing the energy expendituresinvolved with defending these resourcesin each area. Some might argue that the weakest part of Walsberg'sthesis is that the California desertbreeding population is part of, or is the same as, the population that breeds in the coastal areas of southern California. I feel he is correct, and our Phainopeplamovement data in the ColoradoRiver Valley support his contention.Many questionsregarding this populationremain unanswered(do the sameindividuals breed twice per year, etc.), but the evolution of this dual breeding system is logically supported by paleoecologicalknowledge of floral shifts during the Pleistocene.By combining this information, the specificityof the Phainopepla'sfood habits (berries),and the placementand timing of theseresources, Walsberg was able to explain the movementsand the selectiveforces that directed the evolutionof these two different social systems. This work will be an excellent and exciting set of lectures to present to studentsin ornithology or vertebrate natural history. Vertebrate ecologistswill spend many interestinghours discussingand de- bating numerouspoints. Time, and now more directedeffort, will help resolvemany of the still unan- swered questions.--ROBERT D. OHMART. Ontogeny and phylogeny.--Stephen Jay Gould. 1977. Cambridge,Mass., Harvard University Press.xiv + 501 pp. $18.50.--As the title suggests,this bookis a reviewand analysisof the relationship betweenthe processesof ontogenyand phylogeny,a topicin evolutionarybiology