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Reproductive Success of Migratory Birds in Habitat Sources and Sinks Author(s): Therese M. Donovan, Frank R. Thompson, III, John Faaborg and John R. Probst Source: Conservation Biology, Vol. 9, No. 6 (Dec., 1995), pp. 1380-1395 Published by: Wiley for Society for Conservation Biology Stable URL: http://www.jstor.org/stable/2387184 . Accessed: 10/12/2013 16:29
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Reproductive Success ofMigratory inHabitat Birds Sources andSinks


THERESE M. DONOVAN,* FRANK R. THOMPSON, III,t JOHN FAABORG,* AND JOHN R. PROBSTt
ofBiologicalSciences,University *Division of Missouri, Columbia,MO 65211, U.S.A. ForestExperiment of Missouri, MO 65211, U.S.A. tNorthCentral Station, University Columbia, ForestExperiment P.O. Box 898, Rhinelander, U.S.A. WI 54501-0898, Station, tNorthCentral

Abstract: Fragmentationof breedinghabitat in NorthAmerica has been implicatedin thedecline offorestnesting, Neotropicalmigrantbirds.We used a comparativeapproach to examine theeffects offragmentation on three forest-nesting Ovenbird(Seiurusaurocapillus), Red-eyedVireo(Vireoolivaceus),and Wood migrants: Thrush(Hylocichlamustelina).We surveyedbirds and monitoredreproductive success on 28 studyplots in fragmentedand contiguousforestsin two midwesternregions.Distributionof individuals betweenfragmentedand contiguous forestsappeared to varyamong speciesand regions,but total nestfailurewas signifthan contiguous icantlyhigherinfragments forestsin bothregions for all species (p = 0.053). We attributed greaternestfailure to increased nestpredation (p = 0.093) and increased broodparasitism by theBrownheaded Cowbird (Molothrus ater, p= 0.009). In addition to greatertotal nestfailure, partial nestfailure due to cowbirdparasitism led to a reductionin thenumberofhostfledglings. Althoughthecauses of nestmortaltotal nestfailure and partial nestfailure acted in concertto reduce the ityappeared to be species specific, numberof offspring per adultfor all three specieson fragments. We used simplepopulation growthmodels to assess theviability In general, of thethree speciesinfragmented and contiguoushabitatsin bothregions. populations on fragments appeared to be population sinks and populations on continguous forestsappeared to be population sources.Assumingconstantmortality during winter, projectedgrowthindicated that without on fragmentsin both regions immigration Ovenbirdand Red-eyedVireopopulations should become extinct and Wood Thrush populations should be maintained or slightly declineonfragments. Populations ofall three species should increase in contiguoushabitats in both regionswithoutemigration.We suggestthathabitat fragmentation reduceslocal reproduction and may have rainifications for theentire population. A clear understandingof population demographydepends on examination of demographic dynamics within and among sources and sinks. We emphasize thatthelong-term viabilityof thesespeciesdepends on maintaining the breedingrange until thespatial scale at whichsource and sinkpopulalarge tractsofforestthroughout tionsinteract can be determined. de las aves migratorias en habitats fuente Ixito reproductivo y sumidero Resumen: Lafragmentaci6ndel habitat reproductivo en NorteAmerica ha estado relacionada con la disminuci6n de las aves migratorias Neotropicalesque anidan en los bosques. En este estudio usamos un metodo sobre tresaves que anidan en los bosques: Seirus comparativopara examinar el efectode la fragrnentaci6n en 28 parcelas en aurocapillus, Vireo olivaceus,y HylocichlamustelinaMonitoreamosel exito reproductivo y continuos,en dos regionesgeograficasen el medio-oeste de los Estados Unidos. La bosques fragmentados distribuci6nde los individuos entre bosques fragmentadosy contfnuos pareci6 variar entreespeciesy regiones,pero elfracaso totalde los nidosfue significativamente maisalto en losfragmentos que en los bosques los mayoresfracasosde los nicontinuos,en ambas regionespara todas las especies(p = 0.053). Atribufmos dos a la mayor depredaci6n (p = 0.093) y al mayorparasitismo por parte de Molothrus ater (P = 0.053). Ademas del mayorfracaso totalde los nidos,losfracasosparciales, debidos alparasitismoporparte de Moloaddress. Therese Current Donovan, NorthCentralForestExperiment Station,1-26Agriculture Building, University ofMissouri,Columbia,MO 65211, US.A. Manuscriptsubmitted August19, 1994; revisedmanuscript acceptedApril3, 1995. 1380
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thrus ater,condujerona una reducci6nen el ntimerode volantonesde los huespedes.Si bien las causas de la mortalidaden los nidosparecieron ser especificas para cada especie,losfracasos totales y parciales de los nidos actuaron en forma conjunta reduciendoel ntimerode descendientes por adulto en losfragmentos para las tresespecies.Usamos modelossimplesde crecimiento poblacionalpara evaluar la viabilidad de las tresespecies en babitatsfragmentados y continuosen ambas regiones.En general,las poblaciones de losfragmentosparecieron ser poblaciones sumideroy las poblaciones en los bosques continuosparecieron ser poblacionesfuente.Asumiendo una mortalidad constantedurante el invierno,el crecimiento proyectado indic6 que en ausencia de inmigraci6n, las poblaciones de S. aurocapillus y V. olivaceusdeberfanextinguirse en los fragmentosen ambas regiones, y las poblaciones de H. mustelina deberfanmantenerseo declinar levemente en losfragmentos. Las poblaciones de las tresespeciesdeberfanincrementar en los habitatscontinuosen ambas regionesen ausencia de migraci6n.Sugerimosque lafragmentaci6ndel bhbitatreducela reproducci6n a una escala local y podria tenerramificaciones para las poblaciones a escalas espaciales mayores.La clara de la demografia comprensi6n poblacional, depende del analises de la dinamica demograficadentroy entre las fuentesy los sumideros.Enfatizamosque, hasta que se pueda determinar la escala espacial a la cual las poblacionesfuentey sumiderointeractuan, la viabilidad a largoplazo de estas especiesdepende del mantenimientode grandes areas de bosques a lo largoy ancho del area reproductiva.

Introduction
Many species of forest-nesting, Neotropical migrant songbirdshave undergone significant population declinessince 1966 (Robbinset al. 1989). Threenonexclusive hypothesesmay explain these declines in migrant passerines (Askins et al. 1990; Wilcove & Robinson 1990; Finch 1991; Martin1992; Faaborget al. in press; Sherry & Holmes 1993): (1) increasedmortality as they migrate between theirwintering groundsin the Caribbean, Mexico, and CentralAmericaand theirbreeding groundsin NorthAmerica;(2) increased mortality on wintering because of anthropogenic grounds changesin habitat (e.g., tropical or otherfactors; deforestation) and or fitness on breeding grounds be(3) reducedfecundity cause of anthropogenic or loss or habitat fragmentation otherfactors. The habitat fragmentation hypothesis provides a framework forpredicting or rechanges in bird distribution productive success based on local habitat and landscape patterns (Harris1984). Supportof the fragmentation hypothesisrequiresthat(1) fragmentation affects species distribution across habitats and thatthisshift in distribution resultsin populationdecline on local, regional,or range-wide scales, or (2) fragmentation decreases proor survival of birdsnesting in fragmented ductivity habitatsand thisreproductive/survival in results dysfunction population decline on local, regional, or range-wide scales (Robbins1979). is expectedto influence Fragmentation those negatively species thathistorically have inhabited largecontiguous ofhabitat tracts witha relatively low edge density (Temin reproductive ple & Cary1988). The reduction success in the midwestern UnitedStatesis relatedto the amount of forestcover and degree of fragmentation within a landscape (Robinson et al. 1995). In small fragmented habitats birds experience increased nest predation

(Hoover et al. in press; Paton 1994), increased brood parasitism by the Brown-headed Cowbird (Molothrus ater, Robinsonet al. 1993), and decreased pairing success (Gibbs& Faaborg1990; Villard et al. 1993; Van Horn et al. in press). Together,these mechanismsmay limit the reproductive success of forest-nesting migrants (Faaborget al., in press) and mayresultin lower recruitment of individuals into the breedingpopulation.This lower recruitment may negatively influencefuturedemographictrendsat the local scale (Sherry& Holmes 1992) and mayexplainwhy mostlong-term declines of forest-nesting migrants appear in fragmented habitats (Askins et al. 1990). To assess the fragmentation effectsand the habitat size needed to maintain one mustconsider populations, the "area in which youngcan be produced in sufficient numbersto replace adult attrition under the poorest conditions"(Robbins 1979). If reproduction does not compensateforadult mortality, then local populations are sinks:They will eitherbecome extinctor will be maintainedvia constant immigration frombirds produced elsewhere(Brown& Kodric-Brown 1977; Pulliam if reproduction 1988). Alternatively, equals or exceeds adultmortality, thenlocal populationsare sources:They will sustainthemselvesor export excess individuals to otherareas (Pulliam1988). Many studies have used indirectmethods to assess population viability including:utilizingartificial nests (reviewedby Paton 1994); documenting presenceor absence of young produced on habitatsof varyingsize (Porneluziet al. 1993); and documenting the ratioofjuveniles to adultson forests of varying size (Bollinger& Linder 1994; Donovan 1994). However, such indirect methodsare insufficient to uncoverthe causes of reproductivedysfunction thatultimately limit populationson fragmented and contiguous forests. Because habitatfeaturesmayaffect and survival and therefore reproduction

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Minnesota Wisconsin

United States
Missouri . km20 5001

fragmented
X

and contiguousforestsin these two geographic locations;and (3) assess populationviability by usingsimple populationgrowthmodels forbirdpopulationson fragmentedand contiguous in thetwo geographic forests locations.

Study Species
We studied three forest-nesting, Neotropical migrant the Ovenbird songbirds: (SeiurusaurocapiUlus), Red-eyed Vireo (Vireo olivaceus), and Wood Thrush(Hylocichla mustelina). All threespecies breed in U.S. midwestern and eastern deciduous forests.Breeding BJrdSurvey (BBS) data from1966 to 1992 indicatethatthe Wood Thrushhas undergonea significant, range-wide decline, the Red-eyed Vireo has undergonea significant, rangewide increase,and the Ovenbirdhas undergonea nonsignificantrange-wide increase (Peterjohn & Sauer 1994). Breeding BirdSurvey trends in theMidwest reflect range-widetrends (Thompson et al. 1993). As yet, whetherthe differential changes in population trends are relatedto differential responsesto habitat fragmentationor to othercauses is not clear.

contiguous
km

Figure 1. Location offourstudyareas andforest cover in themidwestern UnitedStates.Circlesare examples of JO-km radius landscapes surroundinga fragmented and contiguousstudy plot,shading depicts forestcover. alterthe demography of a population,directexaminationoffitness to habitat componentsin relation patterns at local sitesand acrossregionsis needed (Martin1992). A few studies have examined directly species-specific characteristics of reproductive success of forest birdsin either fragmented (Gibbs & Faaborg 1990; Robinson 1992; Roth & Johnson 1993) or contiguous habitats (Holmes & Sherry 1992); even fewerstudieshave examined these characteristics in both fragmented and contiguouslandscapes (Villardet al. 1993; Hoover et al. in press). Yet, thisknowledgeis requiredto test the fragmentation and to manageviable populations hypothesis (Martin 1992). We testedwhetherfragmentation affects the distribution,reproduction, and viability of threeforest-nesting, bird species in two midwestern Neotropicalmigratory regions.Our objectiveswere to (1) documentdistribution and abundance of forest-breeding, Neotropicalmigrantbirdsin fragmented and contiguousforests in two geographic locations; (2) document the reproductive success and factorsthat limitreproductive success of forest-breeding birdsin fragmented Neotropicalmigrant

Methods
Area Study We studiedthe threespecies' distribution and reproductivesuccess in Missouri (MO) and Wisconsin/Minnesota (WI/MN) in 1991 through1993 (Fig. 1, Table 1). We selected these regionsbecause theycoincide with the center (WI/MN) and southwestern edge (MO) of the Ovenbird'sbreedingrange.The former location is also the northwestern edge of the Wood Thrushbreeding locationis the westerncenrangewhereas the Missouri ter of itsbreeding range.Bothregionsare west-centrally located in the Red-eyed Vireo's broad breedingdistribution.

Table1. Meanlocation, habitat description, andlandscape statistics ofthe28 sitesstudied inWisconsin/Minnesota (WI/MN) andMissouri (MO) in fagmented andcontiguous landscapes.
Landscape Region No. of UTM coordinates sites (Easting, Northing) Forest (%o) cover (ha)* Mean patch size (ha)* No. of patches (ha)* Edge density
(m/ha)*

Mean core area index (%o)*

Contiguous WI/MN Fragmented WI/MN Contiguous Fragmented MO MO

6 6 8 9

5125960, 643090 5029620,519786 666762,4107719 577473, 4311684

92.84 (1.75) 31.31 (12.91) 95.06 (2.00) 32.03 (10.27)

29175 (549.5) 550.49 (613.01) 26793.75 (4890.09) 674.57 (511.92)

1 (0) 26.50 (11.40) 1 (0) 20 (7.88)

3.60 (0.54) 7.31 (1.80) 2.66 (0.82) 8.49 (1.49)

70.62 (3-15) 10.85 (15.14) 75.62 (4.41) 7.59 (4.04)

*AIIstatistics are based on mean (standard deviation) of habitat characteristics withina 10-kmradius surroundingeach site.

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each regionwe examinedbirddemography Within in forest fragments in an agricultural landscape and in contiguousforests. For clarity, we refer to each landscape type(fragmented or contiguous)and region(MO or WI/ MN) combinationas a studyarea (n = 4; Fig. 1). We studiedsix to nine forest plots in each studyarea and treated these plots as replicatesin distribution analyses. Ninefragmented study plotswere in central Missouri and eightcontiguous study plotswere within theheavily forested Ozarks in southeastMissouri.In WI/MNsix fragmentedstudy plotswere in the lowerSt.CroixRiver valley,and six contiguous study plotswerewithin theheavily forested ChequameganNationalForest(WI). We studied all WI/MNplots in 1991 and 1992. In Missouri we studied nine fragmented plots and six contiguous forest plotsin 1991; in 1992 we studiedeightplotswithin fragmented and eight plots withincontiguousforests.In 1993 we located and monitorednests on a subset of area to increasethenestsamplesize. plotsin each study Allstudy 22 ha. Because our plotswere approximately primary objectivewas to measurereproductive success in fragments and contiguous forests, we located our plotsin fragments largeenoughto insurethatarea-sensitive species would be present and that populations would consistofbothmalesand females (Table 1). A forest tract was definedas a fragment based on severalcriteria:size (ha), edge density (m ofedge per ha), and perofforest centcore area (percentage >250 m from edge). tract forest Thus,although fragment rangedfrom181 ha to 1872 ha, all fragments had similar edge densityand percentcore areas (Table 1). Averagepatch size within fragmented landscapes was 550.5 ha (WI/MN) and forests in WI/ 674.57 ha (MO, Table 1). The contiguous MN and MO averaged29,175 ha and 26,794 ha, respectively (Table 1). mature We located study plotswithin relatively (> 40 forestin MO and maple-basswoodyears) oak-hickory in WI/MN. birchforest We selectedWI/MN plotswitha strong componentof oak (Quercus rubra) to minimize of treegeneraamongsites.Despite differheterogeneity all forests ences in forest composition, appeared to be homogeneousin structure (Donovan 1994). In both reoffragmented gionsthe non-forested portion landscapes cool-seasonpastureand row crops. was predominantly andAbundance Distribution locatedon a 150-m We surveyed birdabundanceatpoints were 300 X 600 m where possigrid.Griddimensions dimension and were estabble, givenforest constraints, lishedaroundrandomly selectedpoints.We surveyed 15 located morethan70 m from pointsat gridintersections a habitat/forest ecotonal edge because we were interforest estedin sampling species,notedge species. We visitedeach pointfourtimeson each plot during the breeding season and counted forest-nesting, Neo-

tropicalmigrants and Brown-headed Cowbirds during obcounts(Verner1988). Threeto fourdifferent 10-min servers conductedsurveys at each pointin each yearto minimizeobserverbias (Vernerand Milne 1989). Surveys began after territories were establishedand ended when mostnestshad fledged(May 5 through June15 in 2 in WI/MN). MO; May28 through July In each 10-minute countwe recordedbirddetections within50- and 70-mfixedradiuscirclesas well as total (unlimited distance)detections. We computedthe mean number of detectionsin each distance class for each plot in each year.Plotmeanswere based on 15 counting times a season(n = pointsthat were censusedfour within 60 10-min counts). We used the unlimiteddistance class in analysesto maximizedetections and ensurethat territorial birdsnearedges were adequatelysampled.Although the distribution patternswere similarfor all three distance classes, we also present results from within50 m and 70 m radiiforbetween-study comparisons. We compared mean bird abundance on fragmented and contiguous forests usinga repeatedmeasuresanalysis of variance,with landscape (fragmented or contiguand year ous) and region(MO or WI/MN)as maineffects as a repeatedeffect. Plotswithina landscapewere used as the error term. We used a Type I error rateof 0.1 for of committing a all analysesto reduce the probability are not differType II error (i.e., concludingthateffects entwhen theyactually are). Reproduction nestson a subsetofthe study We locatedand monitored plots (four to five plots in each studyarea) in 1991 and causefailure through1993 to assess reproductive in fragmented and contiguousforests specificmortality in both regions.Nests that were located when active and were low enough (<10 m) to be monitored by direct observationwere included in analyses.All active and Wood Thrush nestswere includedin analOvenbird yses, but canopy-nesting Red-eyed Vireos were ex142 Ovenbirdnests,62 Red-eyed cluded. We monitored Vireonests,and 123 Wood Thrushnests.For each nest, we recorded the number of host and cowbird eggs, chicks, and fledglings every3 to 5 days. We recorded each nest's fateas successful(nests thatfledgedat least one host species) or failed(nests thatfledgedno host failednestsas: depredated(nests species). We classified in which nest, eggs, or nestlingsdisappeared); abanin nest); failed doned (eggs or nestlings leftunattended because of stochasticevents(weatheror otherrandom on a nest); or paraeventssuch as inadvertent trampling sitized by cowbirds (no host young fledgedalthough Because many cowbirdsmayhave fledgedsuccessfully). in that they fledged parasitizednests were successftil beboth cowbird and host young, we distinguished

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tween the proportionof nests that failed because of failure)and the propor(parasitism cowbirdparasitism cowbirdeggs or young(parasittionof nestscontaining ismrate). Success andNesting Mortality DailyNest success on a per nest basis using We calculatednesting (1975) method.These estimatesconsider the Mayfield thefirst ofcause.We considered allnestfailures regardless (ifthe nest day of egg-laying day as the first observation egg was laid) or the day the was located beforethe first the first nest was located (if the nest was located after or failure egg was laid). We observednestsuntilfledging occurred.Foreach species we computeddailymortality dividedby the totalnumas the totalnumberof failures days pooled across all nests within ber of observation foreach each studyarea. We calculated daily survival We species in each studyarea as (1 - dailymortality). the throughout was constant assumedthatdailysurvival nest cycle (Mayfield1975) and calculated nestingsucrateraisedto the exponentthat cess as the dailysurvival reflects the lengthof the nestingcycle (Ehrlichet al. thata nestwill givestheprobability 1988). This estimate fledgeat least surviveits entirecycle and successfully as (1 - nesting one nestling. We calculatednest failure bias occurs ifthe mean clutchsize success). A potential and durationof the nestingcycle varies between fragand between WI/MNand mentsand contiguousforests because of MO. Clutchsize comparisonswere difficult Cowbirds removal Brown-headed host by egg frequent nests on fragments many and because on fragments were located aftertheyhad been parasitized.In addithe exact lengthof tion,we were not able to determine the nestingcycle because we did not monitornests ofnestdaily.However,we do not believeour estimates Ehrlich the data used by because ing success are biased clutch and nest cycle the mean et al. (1988) represent acrossmanystudies. duration For each species we testedwhetherdailynest mortalamong the four studyareas using a Chiity differed (Sauer & Wilcomparisons withmultiple Squareanalysis additional comparisons two liams 1989). We performed of interest: the daily foreach to test directly questions forests versus on fragments contiguous nest mortality in MO versus WI. We used a and daily nest mortality Type I errorrate of 0.1 forall analysesto reduce the a Type II error. ofcommitting probability To determinethe mechanismsthat limitreproducwe and contiguousforests, tive success in fragmented causeinto partitionedthe total daily nest mortality how specificcomponents.For example, to determine we nestpredationsolelyinfluenced dailynestmortality, summed the total numberof nest failuresbecause of predationand dividedthatnumberby the totalnumber we summed days. In the same manner, of observation

events, stochastic caused by abandonment, nestfailures daily nest and calculated cause-specific and parasitism foreach of the fourstudyareas. For estimates mortality dailynest cause-specific each species we testedwhether differed among the fourpopulations,among mortality and among regions and contiguousforests, fragmented using a Chi-Squareanalysiswith multiplecomparisons P valWe considered report). unpublished (Hines& Sauer, theseanalyses. for ues less thanor equal to 0.10 significant Success andNesting Patterns ofDailySurvival objective was to test whetherfragmentaOur primary of and viability the reproduction affected tion negatively vartheseeffects birdsand whether migrant Neotropical We analyzedeach ofthe threeforest-nestied regionally. birds separately.Because ing, Neotropical migratory power, low statistical theseseparatetestshave relatively test (Sokal & Rohlf we used a combined probabilities 1981) to determinewhether the general patternsof across landscapes and regions. differed daily mortality testson This testutilizesa seriesof separatesignificance hypothdifferent setsofdata thattestthesame scientific thetestson theindividual species provide esis. Although significant, a P-valuethatmayor maynot be statistically from separatetestsof signifithe combinedprobabilities a generalizedpatternthatwould cance may illuminate detectedby separateanalyses(Sokal not be statistically & Rohlf1981). As such,we testedwhetherdailymortalof Neotropicalmigrants mortality ityand cause-specific (Ovenbird, Wood Thrush, and Red-eyedVireo comamonglandscapes and regionsand conbined) differed less thanor equal to 0.10 significant. sideredP-values The Mayfield estimatesof daily survivaldo not acone count for partialnest losses (i.e., a nest fledging fouryoung; youngis scored the same as a nest fledging nestloss may Yet, partial both are scored as successful). populationsby reducingthe affect Neotropicalmigrant numberof nestlings per successfulnest. Because cowbirds oftenremove host eggs in nests they parasitize (Lowther 1993), the mean number of host fledglings between successfulneststhatare parasitized maydiffer We calcuand successfulneststhatare not parasitized. relatedparasitism ratesof each species in all fourstudy gions as the percentage of nests containingcowbird fledgedhost eggs or young.For nests thatsuccessftilly per we comparedthemeannumberoffledglings young, in parasitizednest to the mean number of fledglings foreach species. nestsusinga t-test non-parasitized Assessment Source-Sink whether each of thefourpopuWe soughtto determine itself withoutemigration lationsstudiedcould maintain We used three demographiccompoor immigration.

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status: (1) the nents to assess population source-sink producedper adultfemean numberoffemaleoffspring (2) the probamale per yearin a population(fecundity); to reproduce;and thatthose offspring will survive bility rate of adults(Pulliam 1988). The first (3) the survival of young the recruitment two componentsdetermine of young into the breedingpopulation.If recruitment intoa closed breedingpopulationdoes not compensate then the (1 - adult survivorship), for adult mortality of if recruitment population is a sink. Alternatively, youngintothe breedingpopulationexceeds adultmora source (Pultality, then the populationis potentially liam 1988). If = meannumberoffemale 1 - adultsurvivorship X juvenilesurvival, offspring/female/year then the populationwill replace itself(Ricklefs1973). per We calculatedthe mean numberof femaleoffspring femaleper year fromour data and used estimatesof from If and juvenilesurvival the literature. adultsurvival (1 - adult survival)divided by juvenile survivalwas greater than the numberof observed femaleoffspring per female,we concluded the populationwas a sink. if (1 - adult survival)divided by juvenile Conversely, survival was less than the numberof observed female we concluded the populationwas per female, offspring a potential source. To determine whetherthe fourpopulationsstudied were sources or sinks,we made the following assumpthe mean numberof femalefledgtions in calculating lingsproduced per femaleper year.These assumptions marked a trade-off betweenintensively studying represent across populations studying versus extensively populations a largegeographicscale. First, because all threespecies breedersver(first-time studieddo not show age-related we could not in plumage, sus olderbreeders)differences timebreedersdiffered assess iffecundity per yearoffirst forall from olderbreeders;we assumedequal fecundity adultage classes. Second, we assumedthatour speciesspecific estimatesof nestingsuccess and nest failure the breedthroughout method)were constant (Mayfield of ingseason. Last,we assumedthatthe averagenumber femaleyoungthatfledgedper successfulnest was conthe breedingseason. These lasttwo asstantthroughout sumptionswere made because we monitorednests the breedingseason and consideredthese throughout statistics. as season-long estimates We calculatedthe mean numberof femaleyoungper the mean numberof female adultfemaleper yearfrom: nest,populationspecific youngproducedper successful renesofnestsuccess,the number ofpotential estimates and the numberof potentialbroods per tingattempts, season. Forexample,we assumedthatfemaleOvenbirds could fledgea maximum of one brood per yearduring our studyand thatfemaleswould renestonce if their

the mean first nestingattemptfailed.We determined numberof femaleyoungper successfulnest and calculated how manyyoung 100 adult femaleswould prothe breedingseason. Nestingsuccess duce throughout An was 0.27 forOvenbird populationsin MO fragments. averageof 1.58 femaleyoungwere fledgedper successso 27 fulnest (assuminga 50:50 sex ratiooffledglings), femalesproduced 42.66 femaleyoung and 73 females atproduced no young.All 73 femaleswitha failedfirst temptrenested,27% of which produced 31.14 female young.Thus,74.1 femaleyoungwere producedper 100 adultfemalesor 0.74 femaleyoungper adultfemaleper year. The mean numberof femaleyoungper adult female Vireos and Wood per yearwas calculatedforRed-eyed manner,but we assumed thatan Thrushesin a similar oftwo broodsper adultfemalecould fledgea maximum in press) and thatfemaleswould renestif year(Martin, or second nestingattempts failed(Robinson, theirfirst In this manner,we utilized personal communication). nestingsuccess estimatesand the population-specific areach ofthe fourstudy fledging rateswe derivedfrom eas. We assumedthatall juvenilesof all species had a 0.31 of surviving breedingseato the following probability are son. Althoughdirect estimatesof this probability lacking(because juvenilebirdsdispersewidely,Greenratescan be inwood & Harvey1982), juvenilesurvival and estimated based on data foradultmortality directly (1980) femaleproductivity 1973). Greenberg (Rickleffs thatjuvenile and Temple and Cary(1988) hypothesized survival was 0.31, or approximately 50% of adult surand fecundity calculations. vival,based on adultsurvival forthe Wood This estimateagrees with computations Vireo (Noon & Sauer 1992) and Thrushand Red-eyed thatmodeledpopuhas been used byotherinvestigators of forest-interior birds(Temple & Cary lationdynamics 1988; Thompson 1993, Howe et al. 1991). Because our status was to makerelative, goal in assessingsource-sink of the populationsin the four not absolutecomparisons is adequate to addressour areas,the 0.31 estimate study goal. estimates adult survival We obtained species-specific that an adult will survivefrom one (the probability severalsourcesin the season to the next) from breeding were obtainedfrom Because these estimates literature. othersources and applied to the populationswe studon inferences ied, cautionshouldbe used when drawing rates of populationchange in our sites. We used each survivalestimateto assess source-sink species-specific statusin each studyregion.We used the mean of the in of adult survival publishedestimates species-specific our population models. Althoughthese survivalestimates varyfromstudyto study,they are adequate to of populationgrowthin our make relative comparisons populations. fourstudy

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Models Population Projection Source-sink status does not completelydescribe the of a species because it does not populationdemography in reproductionand indicate the relativedifferences time.We will changethrough how populationnumbers used the following two-stagepopulation projection time model to assess change in populationsize through or emiwithno immigration givencurrent reproduction gration:
NA (t + 1) (NAt X SAt) + (NAt X Ft X SJt),

basedon Table2. Repeated measure analysis ofvariance results Thrushes mean detections ofOvenbirds, Red-eyed Vireos, andWood infragmented andcontiguous forests (landscape effect, L) in Wisconsin/Minnesota andMissouri (region effect, R) in 1991and 1992 (year effect, Y).
Species Ovenbird Effect R L Y RXL RXY LXY RXLXY R L Y RXL RXY LXY RXLXY R L Y RXL RXY LXY RXLXY DF 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 F 85.62 0.08 71.16 0.00 50.94 1.70 20.16 9.96 70.80 55.73 29.68 0.40 3.26 3.57 4.44 8.11 5.91 0.20 6.60 0.34 0.07 P 0.0001 0.7736 0.0001 0.9769 0.0001 0.2052 0.0002 0.0041 0.0001 0.0001 0.0001 0.5328 0.0847 0.0723 0.0452 0.0087 0.0246 0.6585 0.0183 0.5671 0.7954

where t = year of simulation, NA = numberof adult is thenumber ofbreeding adultfeNAt breeding females, males in year t, SAtis the survivalof breedingfemale of adultsfrom yeart to yeart + 1, Ftis themeannumber femaleoffspring per adultfemalein yeart, and Sitis the in year in yeart to breeding survival ofjuvenileoffspring t + 1. For each species we used the mean numberof birdscensusedper 10-minute count X 1000 as a starting and propopulation size (for ease of interpretation) jected populationgrowthor decline over 20 years.Our relative popprovidea reasonable, pointcountestimates foreach ofthefourstudy arulationsize ofadultfemales males are predominantly counted eas. Although singing with point counts,mistnet capture data indicatethat and contiguousforestsin the sex ratios on fragments (Donovan significantly WI/MNand MO do not differ 1994; Porneluzi, unpublished,respectively).For the

Red-eyed Vireo

Wood Thrush

population growth models we used the average of the published adult survival estimates for each species. Annual adult survival averaged 0.62, 0.56, and 0.67 for Ovenbirds, Red-eyed Vireos, and Wood Thrushes, respec-

on distance 50 m,70 m,andunlimited ofselected specieswith census deviation (SD) per 10-minute andstandard Table3. Meandetections inWisconsin/Minnesota andMissouri (MO) in 1991and 1992. (WI/MN) (Cont)forests fr-agmented (Frag)andcontiguous
Species Ovenbird Region MO Landscape Frag Year N Mean (<50 m) 0.21 SD (<50 m) 0.10 Mean (<70 m) 0.45 SD (<70 m) Mean (All) 0.69 SD (All) 0.32

Cont WI/MN Frag Cont


Red-eyed Vireo MO Frag Cont

1992 1991 1992 1991 1992 1991


1992 1991 1992 1991 1992

1991

8 6 8 6 6 6
6 9 8 6 8 6

0.35 0.30 0.19 0.75 1.08 0.64


1.28 0.25 0.51 1.38 0.99 0.73

0.16 0.22 0.13 0.11 0.33 0.19


0.37 0.10 0.20 0.35 0.21

0.60 0.56 0.36 1.43 1.68 1.21


2.11 0.37 0.73 2.22 1.69

0.24 0.33 0.22 0.22 0.45 0.41


0.51 0.13 0.29 0.28 0.24

0.20

0.91 0.69 0.74 2.16 2.71 1.78


3.00 0.46 0.94 2.32 2.71

0.35 0.38 0.42 0.58 0.72 0.45


0.68 0.18 0.34 0.29 0.42

WI/MN

Frag Cont

1991 1992
1991 1992

1992

1991

0.69 0.94 1.34


0.29 0.13

6 6
9 8

0.07 0.36
0.12 0.11

0.21

0.34

1.32
1.18

0.42
0.23

1.66
1.97

0.61
0.40

1.51 2.05
0.39 0.23

0.12 0.48
0.18 0.16

1.78 2.60
0.52 0.41

0.24 0.50
0.26 0.28

Wood Thrush

MO

Frag

Cont WI/MN Frag


Cont

1991 1992 1991 1992 1992


1991

6 8 6 6 6
6

0.10 0.06 0.08 0.07 0.15


0.04

0.05 0.05 0.05 0.03


0.04

0.21 0.09 0.17 0.13 0.06


0.11

0.11 0.07 0.12 0.08 0.10


0.05

0.30 0.21 0.33 0.34 0.08


0.09

0.12 0.17 0.15 0.28 0.12


0.06

Conservation Biology Volume 9, No. 6, December 1995

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etal. Donovan

Bird andSinks Sources

1387

t~~~~~~
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t N V"
O .
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Conservation Biology Volume9, No 6, December 1995

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1388

andSinks Bird Sources

Donovan etal.

(nesting success)or fail a nest willsuccessfully fledge andtheprobability that nestsurvival, ofdaily variance nestsurvival, Table5. Daily Thrushes. Vireos, andWood ofOvenbirds, Red-eyed (nestfailure)
Daily survival variance 0.0004 0.0001 0.0002 0.00005 0.011 0.0001 0.0002 0.0002 0.0001 0.0001 0.0002 0.0001 Young per successful nesting attempte 1.58 2.15 1.65 2.00 0 1.57 0.83 1.43 1.07 1.51 1.21 1.50 Fecundity per adult female per year" 0.74 1.32 0.59 1.35 0 1.46 0.63 2.00 0.85 1.75 1.12 2.40

Species Ovenbird

Regiona MO WI/MN

Landscapeb Frag Cont Frag Cont Frag Cont Frag Cont Frag Cont Frag Cont

Nc 15 40 36 51 2 34 13 13 36 56 22 9

Obs. daysd 118.0 364.5 303.5 596.5 12.5 311.0 208.0 161.5 417.0 545.5 226.0 165.0

Daily mortality 0.051 0.038 0.063 0.034 0.160 0.042 0.048 0.025 0.046 0.031 0.040 0.018

Daily survival 0.949 0.962 0.937 0.966 0.940 0.958 0.952 0.975 0.954 0.969 0.960 0.982

Nest failure 0.730 0.620 0.803 0.579 0.991 0.686 0.735 0.495 0.732 0.586 0.681 0.399

Red-eyed Vireo

MO WI/MN

Wood Thrush

MO WI/MN

(WI/MN). aRegionsare Missouri (MO) and Wisconsin/Minnesota bLandscapesare fragmented(Frag) or contiguous(Cont). CN= totalnestsample size. dObs.Days = thetotal numberof observationdayspooled across all nestswithinthestudyarea. eYoungper successful nestingattempt. per successful nestingattempt= themean numberoffemalefledglings fledgedper adultfemaleper year. successfully fFecundity per adultfemaleper year = themean numberoffemale offspring

of all We used a value of 0.31 forjuvenilesurvival tively. threespecies in the populationgrowthmodels and our values in the population growth calculated fecundity models.

Results
andAbundance Distribution were moreabundantin WI/MNthanMO (reOvenbirds between fragbut abundancedid not differ gion effect), Table (landscape effect, mentedand contiguousforests 2). In MO detectionsaveraged 0.80 and 0.72 in fragwhereasin forests respectively, mentedand contiguous WI/MN detections averaged 2.44 and 2.39 on frag(Table 3). mented and contiguousforestsrespectively in 1992 than1991 espewere moreabundant Ovenbirds contiguousplots (region X landcially in the WI/MvN Tables 2 and 3), where mean scape X yearinteraction, detectionsincreasedfrom1.78 to 3.00 detectionsper 10-mincount. Vireoabundancevariedwiththe regionand Red-eyed Table 2). In landscape (region X landscape interaction, 1.82 and 2.19 detecvireo abundance averaged WI/MN, and contiguousforests tions per count on fragmented (Table 3). In MO, however, vireo abunrespectively dance dropped froman averageof 2.52 detectionsper per plots to 0.70 detections count on contiguousforest Vireosincreased (Table 3). Red-eyed counton fragments Taareas (yeareffect, from 1991 to 1992 in all fourstudy bles 2 and 3). Wood Thrushes were more abundant in MO than

than Table 2) and on fragments WI/MN(region effect, Table 2). In MO, contiguousforests(landscape effect, abundanceaveraged0.47 and 0.26 in fragWood Thrush (Table 3). respectively mentedand contiguousforests, theirabundancein WI/MNaveraged0.34 and Similarly, respectively and contiguousforests, 0.09 in fragmented (Table 3). Abundancedeclinedfrom1991 to 1992 (year Table 2), but this decline was most apparentin effect, Table 2) where abunMO (region X year interaction, 0.52 to 0.41 (Table 3). dance decreasedfrom Success Reproductive ifdailynest analysesused to determine The Chi-Square differed dailynest mortality mortality and cause-specific and contiguous fragmented amongthefourpopulations, tests. powerful and regionswere not statistically forests, For example, when the Type I errorrate was 0.1, the testto detecta 0.02 difference power of the Chi-Square among populationsis 0.12. Thus, if in daily mortality rates,we had a mortality had different populationstruly and a high thesedifferences ofdetecting low probability did thattruedifferences (0.88) ofconcluding probability of non-signifinot exist (Type II error).Interpretation should be made in in daily mortality cant differences nonrate.Even statistically ofthehighType II error light in significant can be biologically differences significant of nesting success and populationchange. terms
OVENBIRD

sigof Ovenbirdnestsdid not differ Dailynestmortality among the fourstudyareas (Table 4). In MO nificantly

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Donovan etal.

Bird Sources andSinks

1389

*Parasitism rte

Men Cowbirds/cnus

Non-parasidnest

Paraizednest

0.9 0.8 0.7 0.6 0.5OA


0.3

o.g
.

~~~~~~~~~~~~~a) a
1 4.) 'M5

63
19 54714

0.2 01 0

n=~~~~~~~~~~50 =31
n=40

3-~~~~ : 20

0.9 0.8 0.7?$ 0.60.5 .0.3 0.2 0.20.1


0-1

b)

/~~~~~~~~~ffzi \
n=1 ;_ \ n1 n=11-Sj2L j:12

Ovenbird

Vlreo WoodThrush Red-eyed

{;

Figure3. The mean numberand standard deviation of Ovenbird, Red-eyedVireo,and Wood Thrushfledglingsin non-parasitized and parasitized neststhatsucat least one hostyoung. Nestsamples cessfullyfledged are combinedacross all studyareas. Dailynestmortality caused by cowbirdparasitism and stochastic eventsdid not differ amongthe fourstudyareas (Table 4). Failureby parasitism was a factoronlyin fragments, where 16.7 and 5.27% of dailymortality was to parasitism in MO and WI/MNrespectively attributed (Table 4). Failurebecause of stochasticevents was recordedonlyin WI/MN, for5.27 and 5.06% of accounting on fragments and contiguousforests dailymortality (Table 4). These estimates were not different from the MO populations. 1 out Parasitism rateswere low in contiguousforests; of 40 and 2 out of 50 nestswere parasitized in MO and 8 out WI/MN forests, respectively (Fig. 2). On fragments of 12 and 6 out of 31 nestswere parasitized in MO and WI/MN,respectively (Fig. 2). Across all studyareas 71 nests successfully fledged at least one Ovenbird. Of in those nests the mean numberof Ovenbirdfledglings nestswas lower (2.00) than the mean numparasitized in non-parasitized nests(4.16, fledglings ,berofOvenbird T = 5.41, P = .0001, Fig. 3).
RED-EYED VIREO

0.9 0.8 0.7 0.60.50.40.30.1 0


0.241=

09-

..

>aiE
-

~~~~~~~~c) c

n-I

n-55 MO-Unfr MO-Prag .WI-Unff WI-Frag

Site
Figure 2. Parasitism rate, or theproportion of nests containing cowbird eggs oryoung in each study area, and the mean number of cowbirds detected per 10min census in each study site for Ovenbird (a), Redeyed Vireo (b), and Wood Thrush (c).

nest failurewas 0.73 and 0.62 on fragmentsand contiguous sites, respectively (Table 5). Similarly, in WI/MN nest failurewas 0.80 and 0.58 on fragmentsand contiguous sites, respectively (Table 5). These apparently large differencesin nest failure reflect approximately a 0.025 difference in daily mortality between fragmented and contiguous forests(Table 5). Daily nest mortalitybecause of nest depredation accounted forthe majorityof nest failures(failure to fledge at least one host offspring)in all four study areas but did not differ between regions or landscapes (Table 4). Daily mortalitybecause of nest abandonment was the second largest factorin nest failuresand differedamong the four study areas (Table 4). Abandonment caused failure in all study areas except the MO fragmentsand did not differ

betweenregionsor landscapes(Table 4).

of Red-eyed Vireo nests did not difDaily nest mortality fersignificantly amongthe fourstudyregions(Table 4). In MO, nestfailure was 0.99 and 0.68 on fragments and contiguoussites respectively (Table 5). In WI/MNnest failure was 0.74 and 0.49 on fragments and contiguous sitesrespectively (Table 5). because ofnestdepredation accounted Dailymortality forthe majority in all fourstudy (250%) of nestfailures areas but did not differ between regionsor landscapes (Table 4). Nest predationaccounted for 50% of daily

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1390

Bird Sources andSinks

Donovan etal.

mortality in WI/MN contiguousforestsand >80% of areas (Table 4). in the otherthreestudy dailymortality did not because of nest abandonment Daily mortality differ among the fourstudyareas or between regions was greater in contiguous (Table 4). Nest abandonment landscapes(Table 4) where landscapesthanfragmented accountedfor7.66 and failures because ofabandonment in MO and WI/MNcontiguous 50% of daily mortality (Table 4). habitats, respectively appeared Nest failure because of cowbird parasitism where 19.96%ofdailymortalfragments, onlyin WI/MN (Table 4). Dailymortality to parasitism itywas attributed among the four because of parasitismdid not differ to or betweenlandscapes study areas,between regions, to stochastic was not attributed (Table 4). Nest failure areas (Table 4). eventsin anyofthefourstudy in Vireo nestswere parasitized Three of 28 Red-Eyed MO contiguous forests; and 0 out of 11 nestswere paraforests sitizedin WI/MN (Fig. 2). In MO fragcontiguous mentsonly2 vireonestswere located,one ofwhichwas 8 out of 12 nestswere In WI/MN fragments parasitized. parasitized (Fig. 2). Across all studyareas 24 nests sucat leastone vireoyoung.Of thosenests fledged cessfully in parasitized nests the mean numberofvireofledglings fromthe mean number significantly (2.4) did not differ in non-parasitized nests (3.0; T = of vireo fledglings 1.12,P = 0.277, Fig.3).
WOOD THRUSH

forests were not detectedon contiguous birdparasitism in WI/MN or in MO (Table 4). ofWood Thrushnestswas low to absentin Parasitism 1 of 55 nestsin MO and 0 of 9 nests contiguous forests: cowbirdeggsor young.Conversely, in WI/MN contained in MO 29 out of 36 and 8 out of 18 nests in fragments and WI/MNcontainedcowbird eggs or young(Fig. 2). fledgedat Across all studyareas, 61 nests successfully least one Wood Thrush young. Of those, parasitized fewer(2.07) Wood Thrushes nestsfledgedsignificantly nests (2.94, T = 2.9, P = 0.0049, than non-parasitized Fig. 3). Success DailySurvival andNesting testindicatedthat Resultsof the combinedprobabilities ofnestswas morecloselyassociatedwith dailymortality was landscape than with region. Daily nest mortality but did than contiguousforests, greateron fragments because notdiffer amongregions(Table 4). Nestfailures of predationand parasitism were greateron fragments due to predawhereas failures than contiguousforests, tion and parasitism did not differ between regions(Taand stochastic ble 4). Failuresbecause of abandonment eventsdid not differ betweenlandscapesor regions(Table 4). Migrant Viability Neotropical Adult survivalestimatesfor the Ovenbirdvarygreatly across studies (Fig. 4), rangingfrom 0.845 (Roberts 1971) to 0.540 (Savidge& Davis 1974). When the adult value of 0.845 was used (and juvenile survival survival is 0.31), all four studypopulations produced enough youngto sustainthe populations(Fig. 4a). Alternatively, value of0.540 was used, none of when theadultsurvival the populationsproduced enough youngto sustainthe populationsand all four populationswas classifiedas sinks(Fig. 4a). When the mean adult survival was used of was 0.31, recruitment (0.623) and juvenile survival 1.2 femalefledglings per yearwas requiredforpopulation maintenance. Mean femalefledglings per adult female per yearwas 1.32 and 1.35 in MO and WI/MNconand 0.74 and 0.59 in MO and forests respectively tiguous (Fig. 4a, Table 5). Berespectively WI/MNfragments success was lower forOvenbirdsin fragcause fledging in bothMO and WI/MN in fragments ments, populations were morelikely to be sinksthanpopulationsin contigWhen the mean demographic values were uous forests. used in population growthmodels, Ovenbirdpopulain both regionsdeclined to near extions in fragments tinction within20 yearswithoutimmigration (Fig. 4d). in both MO Ovenbirdpopulationsin contiguousforests and Wl/MNincreasedwithin20 yearsand mayfunction as populationsources(Fig. 4d). Vireo estimates forthe Red-eyed Annualadultsurvival

ofWood Thrushnestsdid not differ Dailynestmortality among the fourstudyareas or between resignifi'cantly differed gions (Table 4). However,dailynest mortality was between landscapes (Table 4). In MO nest failure and contiguousplots re0.73 and 0.59 on fragmented was 0.68 and 0.40 and in WI/MNnestfailure spectively, on fragmented and contiguousplots, respectively (TaFailure because of nest predationaccounted for at in all fourstudy areas but did least 56% ofdailymortality not differ amongpopulations(Table 4). Failurebecause of abandonment accounted for 11 to 33.5% of daily morforin bothWI/MN areasand MO contiguous study tality estsbutwas notdetected in MO fragments (Table 4). Nest failures because of stochasticeventswere detected in and accounted MO contiguous and WI/MNfragments for 5.8 and 11.I% of dailymortality (Table 4). Failures and stochasticeventsdid not caused by abandonment or bediffer areas,betweenregions, study amongthefour tweenlandscapes 4). (Table differed amongthe Failuredue to cowbirdparasitism had higherdaily fourstudyareas (Table 4). Fragments thancontiguousforests mortality because of parasitism accounted for 31.6 parasitism (Table 4). On fragments and 22.4% of dailymortality in MO and WI/MNrespecfailures because of cowtively(Table 4). Alternatively,
ble 5).

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Donovan etal.
32.5 2/ 1.5
*

Bird Sources andSinks

1391

a)

~~~~~~~~~~~50004500
4000/

Source

3500 3000MCandW-C * 25000W 2000

0.5

*A)~~M-F100
W-F )0W-F

1500-

W-F M

0.st
0

t
0.5
1 1.5

Sink
2.5 b) / 3

500 1 9000

M5
10 15 20 e)

2.5
r

Source
{ {/
+*A )W-

< '0

~~~~~ ~~~~~~7000_ ~
6000-

8000-

)~~~~~~~~~~~~~000
/W WF C MS5

3.(J(J 1.5M

)-C / +1 1.5 1W-F

000 2000 I IrI,IIII, 2.


2.5 3 8000

Figure4. Thesource sink status


o

"a
;t

(
0 3

oIII0.O.S

/Sink

iooo1

1,,,,, 1.54 2 2

10

1S

20

Vireos (a), Red-eyed of Ovenbirds (b), and Wood Thrushes(c) based on juvenile survivorship =
rate 0.311:Differentpublished

Source
2.5 2 *

c) )M-C * }M-C /0AT0


AT)C V )W-F A v JM-F

7000/
/ 5000 4003

f)

estimatesof adult mortality are shown by different symbols. For timateof adult survivorship (0.845) of Ovenbirds, allfour

example, using Roberts (1971) es-

1.5
1 {

* {
* */

40-(

3~1~J0002000-

0.5

01
0 0.5
1

W-F M-F

W-

M-C

Sink
1.5
2

1000-

0
3

Y
S 10 1S 20

,
Year

2.5

Adult Survival Mortality/Juvenile

souri contiguous;W-C= WisconM-F sin/Minnesota contiguous; Missourifragmented W-F Wisconsin/Minnesota fragmented) are classifiedas potential sources.Resultsofpopulation simulations for theOvenbird Vireo(e), and (d), Red-eyed Wood Thrush(f) in thefour studyareas. Initialpopulation size (year 1) was derivedfrom
a
1000.

studypopulations (M-C = Mis-

* *
A A +

*
*

1971 Roberts, in press Faaborgand Arendt, Savidge and Davis, 1974 Hann, 1937 Nicholset al., 1981

etal.,1981 Nichols Roth and Johnson, 1993


Robinson, unpubl.

Massachusettes PuertoRico Pennsylvania Michigan Maryland

0.845

ADULT SURVIVAL

0.57 0.54 0.537 NewHampshire -

0.53 0.59

0.64
-

Delaware
Illinois

0.57 -

0.58
0.80

ranged from0.53 to 0.59 (mean survivalestimate = was 0.31, the con0.56, Fig. 4b). Whenjuvenilesurvival were classitiguouspopulationsin bothMO and WI/MN whereas fragfiedas potentialsources forall estimates, mentedpopulationsin MO and WI/MNwere classified as sinks (Fig. 4b). When the mean adult survivalwas was 0.31, recruitment used (0.56) and juvenilesurvival per adultfemaleper yearwas of 1.42 femalefledglings Mean femalefledgneeded to balance adult mortality. lingsper yearwas 1.46 and 2.0 in MO and WI/MNconand 0 and 0.63 in MO and WI/MNfragtiguousforests (Fig. 4b, Table 5). Because fledging ments,respectively success was lower for Red-eyedVireos in fragments, in both MO and WI/MNwere populationsin fragments more likelyto be sinksthan populationsin contiguous

values forests(Fig. 4b). When the mean demographic models,populationsin were used in populationgrowth in both regionsdeclined to near extinction fragments within20 years,whereaspopulationsin contiguousfortimeand poests in MO and WI/MNincreasedthrough serveas sources(Fig. 4e). tentially Annual adult survivalrates for Wood Thrushvaried 1993) to 0.80 (Robinson,unfrom 0.58 (Roth& Johnson published). When the adult survivalrate of 0.80 was used, all fourpopulationsproduced enough young to (Fig. 4c). When the adult compensateforadultmortality in bothMO and WI/ of0.58 was used, fragments survival as sinkswhereas contiguousforests MN were classified as sources(Fig. 4c). When in MO and WI were classified value of 0.67 and juvenilesurthe averageadultsurvival

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1392

Bird Sources andSinks

Donovan etal.

vival value of 0.31 were used, recruitment of 1.06 female offspring per adultfemaleper yearwas needed to compensate for adult mortality (Fig. 4). Mean female fledglings per yearwas 1.75 and 2.4 in MO and WI/MN contiguous forests and 0.85 and 1.12 in MO and WI/MN fragments respectively (Fig. 4c, Table 5). Populationsin contiguousforests in both MO and WI/MNwere classified as sources, whereas populations in WI/MNfragments apparently broke even and populationsin MO fragments were sinks. Because fledgingsuccess was lower forWood Thrushesin fragments, populationsin fragments in both MO and WI/MNwere more likelyto be sinksthan populationsin contiguousforests. When we modeledpopulation withthesedemographic growth values and withoutimmigration or emigration, Wood Thrush populations in contiguous forests in bothregions dramatically increasedthrough time,whereasgrowthin to population was low relative WI/MN fragments growth in contiguoushabitat, and growthin MO fragments was slightly negative (Fig. 4f).

Discussion
Because our primary goal was to documentreproductivesuccess in fragments and contiguous forests in two we purposelystudiedfairly to regions, large fragments avoid the greatly reduced densities(Wennyet al. 1993) and ratesofpairing success (Gibbs & Faaborg1990; Van Horn et al. in press) foundin smallerfragments in the Midwest. Thus givenlargefragment size, we did not exaffected would be significantly pect thatdistribution by Yet our focal species were distributed fragmentation. and contiguoushabitats: differently among fragmented Ovenbird distribution did not differ in eitherMO or WI/ MN; Red-eyedVireos were less abundantin MO fragmentsbutnot on WI/MN and Wood Thrushes fragments; in both MO and WI/MN were moreabundant fragments comparedto contiguous habitats. Without of disspecies-specific knowledgeofpatterns it is difficult to explain these persal or site faithfulness, differences and contigdistributional betweenfragments reflect variation in uous forests. They may geographical the distribution of required resources (Brown 1984). this variationmay resultfromdispersal Alternatively, and distance differences relatedto source-sink dynamics to majorsourceareas (Maurer& Villard1994). Ifthefew young produced on fragments emigrateto breed elsefrom subwhere, dispersal contiguousto fragmented to is maintain populations necessary fragment population size. By contrast, within-forest dispersalmay predominate in contiguousforestpopulations,which may be and less variablethan movementbetween fragmented contiguous subpopulations. and are able Because birdsare highly vagileorganisms to colonize fragments farfromtheirnatal site, our re-

sultssuggestthatdistributional differences maynot reflecta local population'sability to sustain itself (Pulliam 1988). Ifpopulations consist of a pool of immigrants that have rescued fragments fromextinction (Brown & Kodric-Brown 1977), occurrenceand density may be misleading indicatorsof habitat quality and productivity (Van Horn 1983; Pulliam1988). Thus,documentation of reproductive success and dispersalare needed to provide insightsinto why populations are distributed as theyare on fragmented and contiguous forests. Although cause and effect was not established, our reofbreeding sults support thehypothesis that fragmentation habitatreduces the reproductive success of forest-nestare large ing migratory birds even when fragments fromthe inde(>500 ha). The combined probabilities pendent tests for each species showed that the daily nest mortality was greaterin fragmented habitatsthan These differences contiguous habitats. in dailynestmortality were greater betweenlandscapesthanbetweenreUnited gions, suggesting that,withinthe midwestern States,habitat patterns affect migrant birdreproduction morethandifferences in geographic location.Increased nestpredation and parasitism in fragments caused an increase in dailymortality. We suggestthatlandscape featuresin fragmented habitatsmay be more suitablefor predatorand parasite populationsthan in contiguous landscapes (Andren 1992; Thompson et al. in press; Donovan et al. in press). Independentanalysesof all three species show that tended to be higher(0.10 < p < dailynest mortality 0.22) in fragments comparedto contiguoushabitatsin both MO and WI/MN. The nonsignificant landscape trends can be interpreted in two wayswhen analyzedon a per-speciesbasis. First,Mayfield daily nest mortality and nest failuremay trulydiffer between the populations,but our sample sizes and observationdays may have been too smallto detectthesedifferences (i.e., the of 0.02 in dailynest of detecting a difference probability survival was 0.12; Type II errorrate = 0.88). Alternadiffer tively, Mayfield dailynest survival may not truly We measureddailynestsurbetweenthesepopulations. vivalas the probability thata nest would fledgeat least one host young,ignoring partialnest loss. The number of nests thatfailedto fledgeat least one host youngin and contiguous factmaynot differ betweenfragmented that partial forests.However, our resultsdemonstrate can influence thepopnestloss from cowbirdparasitism to maintain itself time ulation'sability through by reducof female the Cowbirds oftenrenumber ing fledglings. move a subsetof host eggs and replace themwiththeir own. Thus,removalof hosteggs by cowbirdsoften only a nest,but these effects would not be partially destroys detected when nestingsuccess is calculated on a per nestbasis. Despite the problems withsmallsamplesizes and nestingsuccess calculations,we believe the patterns detected are real. When we combined our data

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Donovan et al.

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therelationship withothersin the region, betweennestcorlevels were significantly ing success and parasitism indices forour focal relatedwith severalfragmentation migratory birds species and severalotherforest-nesting (Robinsonet al. 1995). Althoughfewer young were produced per adult in the mechafragments compared to contiguousforests, apsuccess in fragments nismsthatlower reproductive Predationreduces Ovenpeared to be species-specific. bird and Red-eyed Vireo nestingsuccess in fragments, and cowbird parasitism reduced the mean numberof cowbirdparanest.Bycontrast, fledglings per successful reduces Wood Thrushnestingsucsitismsignificantly nest theprobability thata Wood Thrush cess, decreasing and decreasing would fledgeat least one host offspring nests.Thus, in successful the mean numberof offspring fragmentation negativelyaffectedreproductionin all of thisreduction varthreespecies, but the mechanism ied amongspecies. Our resultssuggestthatfragmentation reduces both fledgeat least theprobability thata nestwillsuccessfully and the mean numberof offspring one host offspring birdpopproducedper nest.We suggestthatmigratory as a ulationsin the breedingseason maybe structured networkof sources and sinks,where more productive subsidize or rescue unproducsource areas effectively tive sink areas (Pulliam 1988; Brown & Kodric-Brown 1977; Stacey& Taper 1991; Rolstad1991). If thisis so, the relevantdemographicunit for long-term manageof subpopulations that mentofthesespecies is a system are linked by dispersingindividuals(Hanski & Gilpin et al. 1992; Harrison 1994). 1991; Villard of some migrant If demography bird populationsis in a source-sink thenappropriate longfashion, regulated efforts theconsequences term conservation mustconsider a largeproportion ofsource-sink Numerically, dynamics. of a population may reside withinsinks at any given time, and althoughsinks may not persistindefinitely, to overall population significantly they can contribute size and longevity (Pulliam1988; Howe et al. 1991). Geto a more diversegene sinksmay contribute netically, is 1987), but this benefit pool (Lande & Barrowclough exand emigrants are not successftilly lost ifimmigrants (Howe et al. 1991). changedamongsubpopulations in thatconstant Sinksmaybe a detriment immigration fromsources to sinks(fragments) mayexerta negative on thelarger influence population (Davis & Howe 1991). sinksmaybe draining sources,resultDemographically, ing in widespreaddecline of some species across their of Ovengeographic ranges.Thus,a clearunderstanding andWood Thrush demogVireo, population bird, Red-eyed ofdemographic dynamics raphy dependson examination and amongsubpopulations thatare linkedby diswithin has significant persal. Because habitat fragmentation for local Ovenbird,Red-eyed population ramifications Vireo,and Wood Thrushpopulationseven on relatively

largefragments, we suggestthatthe long-term viability of these species depends on maintaining heavilyforthe breedingrange until ested landscapes throughout the geographicscale at which source and sinkpopulacan be determined. tionsinteract

Acknowledgments
ofmany theassistance We gratefully acknowledge people dataused in thismanuscript, A. who gathered including: A. Buerkle, D. Cooper,J. Dunham, Anders, D. Burhans, D. Fink,C. Freeman, M. Guzy,J. Hayes,R. Kennedy,K. Loraff, S. Negri, J.Porath, P. Porneluzi, J.Preebee,J.VoK. Ryerson, G. Shauer,A. R. Weisbrod gel, D. Novinger, K. Winters, and the undergraduate interns from the Misof Conservation. David Larsonhelped souriDepartment and W. Dijak providedadcalculatelandscape statistics, ditionalGIS assistance.GaryKrauseprovidedmanysuggestions regarding statistical analyses,and this manuscriptgreatly benefited by the helpfulcommentsof C. and J. Weaver. Galen, P. Jones,M. Ryan,R. Semlitsch, Fundingwas providedby the U.S. ForestServiceNorth CentralForest Experiment Station,the MinnesotaDeof Natural partment ResourcesNongameWildlife Fund, the University of MissouriDivision of Biological Sciences, and the NBS Global ChangeProgram. Literature Cited
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