The Auk 111(3):652-660, 1994

CAUSE AND EFFECT IN POPULATION DECLINES OF

MIGRATORY BIRDS

JOHN H. RAPPOLEAND MARY VICTORIA MCDONALD •

Conservation
andResearch
Center,NationalZoological
Park,Smithsonian
Institution,
FrontRoyal,Virginia22630,USA

AnsTRt•CT.--Population declineshave been reportedfor nearly one-third of all Nearctic

migrantbirds that winter in the Neotropics.Speculativeexplanationsfor thesereportshave
beenpresentedemphasizingthe importanceof eventsoccurringduring one portionof the
annualcycleversusanother.Theseexplanationsaredifficultto testdirectly.However,certain
predictionscanbe maderegardingthe characteristics of populationscontrolledby breeding-
ground factorsasopposedto thoseassociated with migrationor wintering-groundphenom-
ena. Thesecharacteristics can be measuredfor and testedexperimentallyon a species-by-
speciesbasis.We present14 suchpredictions,and review datacurrentlyavailableto assess
their relevanceto observeddeclines.Basedon thesedata, populationsof many speciesof
Nearcticmigrantsappearto be controlledby wintering-groundevents.Received 30 September
1993, accepted
5 January1994.

DECLINESIN AT LEAST109 speciesof Nearctic In this paper, we do not evaluate the evidence
migrantshave been reported by one or more for declines per se. Rather, we make several
survey methods(Droeõe and Sauer 1989, Mor- predictionsregarding demographicaspectsof
ton and Greenberg 1989, Terborgh 1989, Rob- migratory bird populations based on the as-
bins et al. 1989a, Askins et al. 1990, DeGraaf and sumption that these populationsare declining
Rappolein press?.These figures are causefor asa resultof breeding-groundfactors.Available
concerndespiteconflictingregionalpopulation information from our own work or the litera-
trends (Sauer and Droege 1992), and despite ture is presented to aid in evaluation of each
arguments that at least some purported evi- prediction. In the interest of conciseness,we do
denceof declinemaybe the resultof procedural not attempt to provide a comprehensive liter-
errors (e.g. survey design and analysisflaws; ature review; instead, we refer the reader to
Hutto 1988, Jameset al. 1990, 1992, Rappole et reviews of the subjectcontainedin Terborgh
al. 1993a). If the declines are real, what are the (1989), Hagan and Johnston (1992), Morton
causes? (1992), and DeGraaf and Rappole (in press).
Measurementof mortality rates for different After presentingour predictionsand brief ar-
stagesin the annualcycleand their relationship guments,we concludethat wintering-ground
to reproductiveratesis the key to determining habitat alteration,not breeding-groundfactors,
when populationregulationis takingplace,and provides the best explanation for the observed
to understandingthe origin of migratory-bird declinesin many Nearctic avian migrant spe-
declines. However, this measurement is diffi- cies.Finally, we encouragecriticalteevaluation
cult enough for game specieswhere a variety of our predictionsand conclusions,and we pro-
of natalityand mortalitydatais available(Nich- vide' somesuggestionsregarding further tests
ols 1991); such measurementsare considerably of the hypothesis.
moredifficult or impossibleto achievefor song-
birds. Nevertheless, migratory-bird popula- HYPOTHESIS AND PREDICTIONS
tions are likely to have different characteristics
depending upon whether population regula- The hypothesis to be evaluated is that pop-
tion is a breeding versusa nonbreeding phe- ulations of Nearctic avian migrants are declin-
nomenon. These characteristics can be mea- ing as a result of breeding-groundevents.We
sured. discuss14 predictionsbasedon this hypothesis.
Prediction1. Winteringsitesfor migrantsshould
• Current address:Departmentof Biology,Univer- not belimited.Migrantsshouldhavetheirchoiceof
sity of Central Arkansas,Conway,Arkansas72035, optimalwinterhabitatsites,and shouldnot occupy
USA. suboptimalsites.--Thisprediction is derived from
652
July1994] Population
Declines
ofMigratory
the population models of Fretwell (1972:89),
which state that habitats of lower suitability
will be occupied only when the "basic suit-
ability" of optimal habitat declinesdue to over-
crowdingto the point where it equalsthe "basic
suitability" of suboptimalhabitat. If breeding
habitatwere limiting migrant populations,then
more optimal winter habitat should be available
than necessary,and individuals should be high-
ly selectivewith regard to habitat on the win-
tering ground--avoiding suboptimal habitats
(Brown 1969, Fretwell and Lucas 1970, Krohn
1992).
Severalstudiesindicatethat migrants,in fact,
are not restrictedto a few, apparently optimal
habitats on the wintering ground (Karr 1976,
Hutto 1980,Waide 1980,Greenberg1992,Lynch
1992). Migrants are found in a wide variety of
habitats, including those that are apparently
suboptimal(seereviews and data in Rappoleet
al. 1983:7-14,Rappoleet al. 1989,Askinset al.
1990).
Observationof this phenomenonhas caused
many observersto conclude that most migra-
tory birds are habitat generalistscapableof tol-
erating a wide range of winter habitats (e.g.
Morse 1971, Petit et al. 1993). An alternative
explanationis that more birds are presentdur-
ing the winter period than the declining
amountsof optimal winter habitatscan support
(Rappoleand Morton 1985),forcing individuals
to use suboptimal winter habitats.
Prediction
2. If breeding habitatsarelimiting,ap-
parentlysuitablebutmarginalbreeding habitats(i.e.
lowerin relativefitnessthan optimalbreeding hab-
itats)shouldappearfilled with individuals of both
sexesattempting to breed,regardlessof thepressure
from nestpredation,parasitism, or similarfragmen-
tationeffects,sincethe principalalternativewould
betoforego
breeding
altogether
(zerofitness).-Con-
trary to this prediction, abandonment rather
than occupationof breeding habitatby Nearctic
migrants has occurred. This abandonment has
been identified as "the area effect" in which
small or even medium-sizedsites(up to 500 ha
for some species) with apparently suitable
breeding habitat are not occupied (Galli et al.
1976, Robbins et al. 1989b, Askins et al. 1990,
Freemark and Collins 1992, McShea et al. un-
publ. manuscript).
An explanation,basedon the assumptionthat
populationsare controlledby wintering-ground
events, would be that the "area effect" results
from the fact that too few breeding adultsare
Birds 653
returning to fill all available breeding sites
(Wilcove and Terborgh 1984,Askinset aL 1990:
26, DeGraafand Rappolein press).If this were
the case(i.e. if fewer birds were returning than
necessaryto fill all available breeding space),
onewould predictthatbreedinghabitatswould
be abandonedin a regular, predictablefashion
from leastto most favorable,as suggestedby
the Brown-Fretwell-Lucas model of habitat use
(Brown 1969, Fretwell and Lucas 1970, Krohn
1992). In other words, the "area effect" would
be the result of fewer birds returning to breed
than the habitat can support.
Prediction3. Migratorybird declinesshouldnot
beobserved in breeding
habitatsthatareundisturbed,
and presumably optimal.--Contrary to this pre-
diction, long-termstudieshave beenperformed
in apparentlyundisturbedbreedinghabitatsthat
have found declinesin Nearcticmigrants(e.g.
Hall 1984,Marshall 1988,Holmesand Sherry
1988).Thesestudiesarebasedon countsof sing-
ing males, a number that can vary markedly
depending on the number of unpaired males
in the population (Rappole and Waggerman
1986, Rappole et al. 1993a, Gibbs and Wenny
1993).If wintering-groundfactorscausea large
reduction in number of adults returning to
breeding territories, this decline should cause
a reductionin number of unpaired males(since
more breeding territories would be open to
them), resulting in decline of the amount of
song, and a concomitantdecline in the popu-
lation estimatesbasedon songcounts.
A 15-year study of a banded population of
KentuckyWarblers(Oporornis formosus) in ma-
ture oak forest,initiated by E. S. Morton in Vir-
ginia, has assessedterritory occupancyand
singing behavior (McDonald and Morton un-
pubL manuscript).Attributes of optimal Ken-
tucky Warbler territories have been character-
ized (McShea et al. unpubl. manuscript),and
the proportionsof available optimal versusoc-
cupied areasnoted for each year. In 1979, the
first year of the study, 60 (89%) of the 73 ter-
ritories used at one time or another during the
15-year period were occupied.By 1993, only 36
(45%)of the apparentlysuitableterritorieswere
occupied.This decline is highly significant (r2
= 0.81). Yet, rates of predation, brood parasit-
ism,and numberof young successfully fledged
per pair have remained unchanged over the
years.
Prediction
4. Declinesshouldnot occurin species
wherenoapparent
change
hasoccurred
in breeding
654 R•PPOLE
A•r• McDoN^Lr• [Auk, Vol. 111

habitat.--Use of remote sensingeventually will species.Holmes and Sherry (1992) performed

make the comparativeassessmentof absolute sucha studyand found that AmericanRedstarts
amount of breeding versuswintering habitats (Setophagaruticilla) and Black-throated Blue
possible(Green et al. 1987, Powell et al. 1992, Warblers (Dendroica caerulescens)showed a
Homer et al. 1993,Rappoleet al. in press).Lack- higher degreeof site fidelity to the wintering
ing the information necessaryto completesuch ground than to the breeding ground, contrary
a comparisonat present, researchershave ex- to Prediction 5. However, they provide an al-
amined subsetsof Nearctic migrants for which ternative explanation, maintaining that breed-
declines have been recorded and which winter ing-ground habitatsvary markedly in quality
in lowland wet forest, one of the most threat- from year to year, and low return rates reflect
enedtropicalhabitats.Thesecomparisons show movementsof competitiveindividualsto high-
a strong correlation, not with lossor degrada- er-quality sites.
tion of breeding habitat, but between observed We disagreewith this explanationfor higher
declines and loss of winter habitat (Robbins et return rates to wintering than to breeding
al. 1989a, Askins et al. 1990:41-44). groundsfor the following reasons:First, there
I.• fact, there is debate over whether or not is no evidenceto indicatethat the missingbirds
breeding habitat has declined at all for many havemovedto "higher-quality"sites.The birds
forest-relatedmigrantsin recentdecades.Avail- are simply not present.Second,if breeding hab-
able data indicate that breeding habitat for a itat were limiting, individuals would not have
number of forest-relatedspecieshas increased the option of moving to higher-quality sites,
during this period in many parts of North since these likely would already be occupied.
America (Birch and Wharton 1982, Powell and Third, there is no reasonto believe,a priori,that
Rappole1986).In contrast,wintering habitatfor breedinghabitatsare any more or lessvariable
the Wood Thrush (Hylocichlamustelina)has de- on an annual basisthan wintering habitats.
clined by 23%in southernBelize, 73%in north- Prediction6. The proportionof youngbirdsal-
eastern Costa Rice., and 95% in southern Mexico lowedto enter the breedingpopulationshouldde-
since pre-Columbian times (Rappole et al. in creaseas theamountof qualitybreeding habitatde-
press).Even for those speciesfor which scrub creasesrelative to quality winter habitat.--If
or second-growthhabitatsare optimal, habitat breedinghabitatwere limiting, one would pre-
availability in many parts of the tropicsis be- dict that available breeding siteswould be oc-
coming increasinglydifficult. Shadecoffeeand cupied mainly by older, experiencedindivid-
cacaoplantationsandshiftingagriculture,which uals. Older birds would return to successful
once provided considerablehabitat for these wintering sitesand show higher survival and
birds, have given way in many areasto mono- return rates, due to experienceand site domi-
cultures(G. Stiles pets. comm.). nance,than youngerbirds.However, if winter
Prediction5. Springreturnratesby adultsto un- habitat were limiting, both older and younger
altered,optimalbreeding sitesshouldbehigherthan birdswould be forcedinto lower-qualitywinter
return rates by adults to unalteredwinter sitesin habitatswhere age and experiencewould pro-
optimalhabitat.--If adult birds are equally faith- vide lessof a competitiveedge becausedensity-
ful to both destinationsof the migratory jour- independent factors would more likely be
ney, lower return rateswould be expectedfrom operative.Indeed, if winter habitatswere de-
birdsreturningfrom the sitemostlikely to have clining rapidly, then the experience of older
beenaltered;that is, if breeding-sitequality has birds in sites formerly optimal but now de-
been reduced (reducing survivorship), birds graded would be nullified and might even be
should show a relatively lower rate of return disadvantageous.This negative effect on older
to winter site (than breeding site) and, if the birds would result in an overall decrease in the
wintering site has been altered, there should ageof the population,an outcomethat hasbeen
be lower return rates to the breeding site. observedby Sherry and Holmes (1992, 1993).
Site fidelity has been documentedfor many However, they explain this phenomenonasthe
speciesof migrantsfor both the breedingand result of cyclic variation in production of off-
winter seasons(Rappoleet al. 1983,Mabey and spring from year to year based on food avail-
Morton 1992),but seldomhasthis phenomenon ability and on other breeding-groundphenom-
beenexaminedduring the sametime periodfor ena.
both breeding and wintering populationsof a Prediction7. Thenumberof nonbreedingmalesin
July1994] Population
Declines
ofMigratory
Birds 655

the breedingpopulationshouldbe high.--Non- Wood Thrushes in Veracruz rain forest and sec-
paired males have long been recognized as a ond growth. They found that,while many Wood
characteristic aspectof breedingpopulationsfor Thrushesoccupiedsecondgrowth habitats,they
many speciesof birds (Hensley and Cope 1951, sufferedhigher mortalityratesin thesehabitats
von Haartman 1971, Rappole et al. 1977). These and continually attempted to locate territories,
males fall into four categories:males holding moving as wanderers (floaters) through occu-
territory in apparently optimal habitat; males pied territories.Rappoleand Morton (1985)and
holding territory in suboptimal habitat; wan- Rappole et al. (1992) found that, while many
derers(floaters)moving over large distancesin speciesof forest-related migrants were found
searchof unoccupiedterritories;and "lurkers" in low second growth, the turnover rates in
(McDonald et al. unpubl. manuscript),unpaired these marginal habitats appeared to be much
males that do not move over large areas, but greater than in primary forest. Rappole and
stay on or near the territories of one or two Warner (1980) performedremoval experiments
paired males. If breeding habitat is limiting, on Hooded Warblers (Wilsoniacitrina) in Vera-
there should be large numbersof individuals cruz and found that removed territory holders
in the latter three categories.However, if win- were replaced by wanderers.
ter habitat is limiting, fewer individuals should Prediction9. The numberof breeding
individuals
return north to breed than the available breed- in populationsof migrants
in optimalhabitatsshould
ing habitat could accommodate,and a decline notfluctuateappreciablywithpredator /preyandcli-
should be observed in the number of nonbreed- maticcycles onthebreeding groundbecause theywill
ers in the population becausean abundanceof bebuffered bytheeffectof havingexcess numbersof
breeding habitat is available in which they can potentialbreedersin the population.--A conse-
settle and establish territories. quenceof having more winter habitatavailable
This phenomenonhas been observedin the than breeding habitat should result in having
aforementioned Kentucky Warbler study in many more individuals in breeding condition
Virginia. The total number of male Kentucky than the breeding habitat can support. How-
Warblers (both territorial and nonterritorial) ever, if winter habitatwere limiting, one would
capturedon the study site annually declined predict that fewer breederswould be available
from 80 to 50 individuals between 1980 and than amountof breeding habitat,and droughts,
1993(McDonaldet al. unpubl. manuscript);over cold snaps, and similar factors affecting mor-
the sametime span,the number of individuals tality would translateinto directmeasurableef-
occupyingsuboptimalsites declined from an fects on the number of breeding individuals.
estimated 20 individuals to 0, and the total This phenomenonhasbeen observedby Blake
number of lurkers declined from about 10 to 5 et al. (1992),who found that regionalmigratory-
individuals (based on intensive multiyear in- bird breeding abundancesin the Midwest were
vestigation of a breeding population of color- affected negatively by drought. Rodenhouse
bandedbirds using netting, playback,and ob- (1992) constructed a model to demonstrate the
servation). potential effectsof climatic changeon warbler
ComparingDNA-basedmarkersor usingoth- populationsin which he predicted preciselev-
er molecular-genetictechniquescould provide els of population response to specific climatic
additional information on this phenomenon.If fluctuations.This model presumesthat the pop-
populations are declining below the carrying ulation is below breeding-habitat carrying ca-
capacityof optimal breeding habitat, then rates pacity and, therefore,annual recruitmentinto
of extrapair fertilization not attributable to the breeding population bears some direct re-
neighbors should decline as well. lationship to fledging success. If breeding hab-
Prediction 8. There should be little or no evidence itat were the critical factor limiting population
of floaters(wanderers) in winteringpopulations.- size, this result would not be expected.
In contrastto this prediction, those studiesin Prediction10. Thenumbers of territorialindivid-
which territorial competitionfor winter habitat ualsin optimalwinter habitatshouldshowsharp
has been studied in recent years have docu- annualfluctuations.--Ifthere were more optimal
mented evidence of floaters (or "wanderers" winter habitat available than necessaryto sup-
sensu Wunderle 1992; Winker et al. 1990, Stacier port the population,there shouldbe significant,
1992, Wunderle 1992). Rappole et al. (1989) observablechangesin numbers of individuals
measuredmortality rates and movementsof in winter habitat as the numbers of migrants
656 RAPPOLœ
ANt)McDOIqALt) [Auk, Vol. 111

decline asa result of breeding-seasonphenom- kins et al. 1990:2, Blake et al. 1992, Litwin and
ena. In fact, this effect has not been observed. Smith 1992). However, Johnstonand Hagan
Where studied, wintering migratory-bird den- (1992) reported someinstancesin which tem-
sities in optimal habitats have remained re- perateresidentpopulationchangeswere posi-
markably stableacrossyearsand latitude (Rap- tively correlatedwith migrant trends.
pole in press),indicating the presenceof large Prediction
13. Declinesin Nearcticmigrants
should
buffer populationsin suboptimalhabitats. not be paralleledby changesin nonmigrantpopu-
Prediction11. Alterationofbreeding
habitatcould lationsoccupying the samewinteringhabitats.--If
bringsomespecies with similarecological
require- lossof breedinghabitatis causingthe decline
mentsintocompetition, forcinggeneticor competi- in Nearcticmigrants,then a comparisonof re-
tivereplacement (1980) gionalpopulationtrendsof wintering migrants
of oneof thespecies.--Gill
has documented evidence of increased inter- and tropicalresidentsoccupyingthe samehab-
breedingof populationsof Golden-wingedand itats should show declinesin migrant popula-
Blue-wingedwarblers(Vermivora and tions while resident populationsremain stable.
chrysoptera
V. pinus)over the past half century, resulting However,if lossof wintering habitatis the cause
in gradualdisappearance of the Golden-winged of migrantdeclines,thenpopulationsof species
Warbler.This phenomenonhasbeen explained in both groupsshouldremain stablein optimal
as the result of recentbreeding-groundcontact tropicalhabitatsor, alternatively,migrantsand
betweenthe two species.However, there is an residentsrequiring a given habitat should de-
alternativeexplanation.Interbreedingmayhave cline in parallel if that habitat is degraded.Ev-
increased as an indirect result of the fact that idence for the latter occurrence was found in a
these two specieshave very different winter- 21-yearstudy in CostaRica (Stiles 1990).
habitatusepatterns.The Blue-wingedWarbler Prediction
14. Theratioof songbirdsmigratingin
winters "in a variety of brushyareas,scruband fall compared to springshouldincrease
overtime.-
openwoodland"along the Gulf and Caribbean If carryingcapacityof breedingversuswinter-
coastsof Middle America (AOU 1983:602), hab- ing habitats were equal, the total number of
itatsthat are not threatenedin mostpartsof its birds in fall should be much larger than the
range.In contrast,the Golden-wingedWarbler number of birds in springasa resultof the large
winters in lowland wet tropical forests,a hab- number of offspring added to the population
itat that hasundergoneconsiderablealteration during the breeding period because,by spring
(Rappoleet al. 1993b).If Golden-wingedWar- the number will have been reducedby six to
biers have been seriouslyreducedin numbers eight monthsof mortality factorsactingon the
asa resultof winter-habitatloss,they may have population. However, if overall bird popula-
been forced to searchfor mates in the closely tions are declining as a result of breeding-sea-
related Blue-wingedWarbler population,a sit- sonphenomena,the numberof birds heading '
uationanalogousto what hasoccurredbetween southin fall shoulddecline more sharplythan
gray wolves (Canislupus)and coyotes(C. latrans; the number of birds returning north in spring.
Wayne et al. 1992). Measurement of these kinds of trends for mi-
Prediction
12. Declines should gratorysongbirdsasa groupwould be difficult
in Nearcticmigrants
be paralleledby changesin temperate,nonmigrant or impossiblebasedon small samplesat a few
populations occupyingthesamebreeding habitats.- migration stopoversites due to the confound-
If fragmentationcausesseveredegradationof ing effectsof weather (Moore et al. 1993).How-
breeding habitat as a result of loss of micro- ever, it may be possibleto assesstrends in rel-
habitator increasedexposureto predationand ative numbers of migrants using rigorous
cowbird parasitism (Askin'set al. 1990), then samplingprotocolsand censusingor netting for
residentspeciesusing thesesamehabitatsthat extended periods at several permanent sites
have ecologicalrequirementssimilar to those throughouteachmigrationseasonfor a number
of migrants(e.g.opennestingbirds)shouldde- of years.As yet, this has not been done.
cline in fragmentsin a manner similar to mi- Another technique that may prove to be a
gratoryspecies.Contraryto thisprediction,most feasiblemethod for assessing fall/spring pop-
breeding-groundstudieshave foundthat,while ulationratiosfor migrantswould be to compare
migratory speciesdeclined on study sites,res- the volume of migrating birds near the Gulf
ident populationsdid not decline (Lynch and Coastusing weather-radarfilms, which record
Whitcomb 1978, Briggsand Criswell 1979, As- flocksof migrantsin nocturnalflight. Changes
July1994] Population
Declines
ofMigratory
Birds 657

in migration volume could be analyzed for a moval experimentscan be used to test for the
period coveringseveraldecadesusing this pro- presence of wanderers (floaters) on both the
cedure, providing estimates of amounts of breeding and wintering ground in habitats of
changein fall versusspring volume over time. different quality. Return ratescan be calculated
Analysesto date provide evidencesuggesting for bandedbirdsat breedingand wintering sites.
that absolutenumbersof migrating birds had As valuablelong-term and in-depth studieson
declined significantly between the years 1957 breeding,stopover,andwintering-groundpop-
and 1990 (Gauthreaux 1992). This finding, of ulationscontinueto accumulatedataon repro-
course,is not useful for determining when in ductive success,return rates,and survivorship,
the life cycle the declines are occurring. That it should be possibleto constructmeaningful
determinationwill requirelong-termfall/spring and comprehensive life tables. Significant
ratio comparisonsfrom specificsites. changesin recruitment rates or in proportions
of different age classesshould help to elucidate
CONCLUSIONS relevant demographictrends in these popula-
tions that mere countscannotprovide. Another
Our chief aim is to move the debate on mi- use of life tableswould be comparisonsof de-
gratory-bird demographicsand conservation mographictrends between breeding and win-
from the realm of speculationbasedon infer- tering grounds for a given species,or among
encesinto one in which hypothetico-deductive ecologicallysimilar species(asin Robbinset al.
principlescanbe applied. The formulationand 1989a).Further testsof the predictionsmadein
examinationof the predictionspresentedhere- this paper could involve comparisonof tem-
in are meantasa stepin this direction.Fourteen perate and tropical resident populations with
predictionshave been made basedon the hy- thoseof migrants.Foodsupplementscould be
pothesisthat Nearctic migrant population de- provided on breeding territories to test Sherry
clines have occurredas a result of changesto and Holmes' (1992) idea that food availability
breeding-habitatamount or quality. Examina- ("habitat quality") is the factor driving adult
tion of thesepredictionsbasedon the literature return rates, rather than decreased overwinter
doesnot support the hypothesis.Alteration of survival. With the increasing power of geo-
wintering-ground habitat provides the most- graphic information systems,it may even be
parsimoniousexplanationfor the observedde- possiblebefore too long to calculateamounts
mographic characteristics.However, there is of breedingand wintering habitaton a species-
sureto be discussionover the appropriatepre- by-speciesbasis(Homer et al. 1993,Rappolein
dictions. Deduction of reasonablepredictions press). Other tests will become obvious when
that can be used to discriminate between the the questionis viewed from the perspectiveof
competinghypothesesshouldprove to be a fer- testable predictions based on likely alterna-
tile field for debateand experimentation. tives. If our tentative conclusions are correct,
For someof the aboveexplanations,there are the focus of conservationactivities for many
alternative explanationsbased on breeding- speciesof migratory birds should be on factors
habitatlimitation (e.g.Holmesand Sherry 1992, affectingnonbreeding-seasonsurvival.
Sherry and Holmes 1992),or a combinationof
breeding/wintering population limitation ACKNOWLEDGMENTS
(Sherry and Holmes 1993). In addition, it must
be recognizedthat not all migratoryspeciescan We thank S. Derrickson, W. J. McShea, E. S. Morton,
J.D. Nichols,G. V. N. Powell, F. G. Stiles,A. R. Tipton,
be expectedto respondin the sameway, and
K. Winker, and J.M. Wunderlefor helpful comments
that different factorsmay be at work in different
and discussionsof the ideas in this paper.
regions of North America or in the Tropics
(O'Connor 1992). For instance, trends are not
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lend themselveswell to further testing. Re- North America. Curr. Ornithol. 7:1-57.
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ANDMcDONALD [Auk, Vol. 111

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