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Female Site Fidelity and Polygyny in the Blackpoll Warbler (Dendroica Striata)

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Female Site Fidelity and Polygyny in the Blackpoll Warbler (Dendroica Striata) FEMALE SITE FIDELITY AND POLYGYNY IN THE BLACKPOLL WARBLER (DENDROICA STRIATA) BONITA C. ELIASON JamesFord Bell Museum of NaturalHistory, Department of Ecologyand Behavioral Biology, Universityof Minnesota,Minneapolis, Minnesota 55455 USA ABSTR^CT.--From1980 to 1982,8-30% of male BlackpollWarblers (Dendroica striata) studied on Kent Island, New Brunswick,were bigamouseach year (2 = 16.8%).I testedwhether differencesin the qualityof the breedingsituation can give rise to polygynousmatings. Five territory parametersand two male parameterswere usedas measuresof quality. Male arrival timeswere correlatedwith malemating status and the reproductivesuccess of monogamous females.Territories of bigamousmales had morelarge conifersthan did thoseof monoga- mous males.Territory parametersdid not appear to influence female choices. Strongsite attachment by femalesmay lead them to matebigamously when few malesare availableearly in the seasonnear the females'former nest sites. The reproductivesuccess of secondaryfemales was not differentfrom that of monogamousfemales. Any costsof mating bigamouslymay be offsetby increasedreproductive success associated with early nest ini- tiation and competitiveadvantages conferred by site dominance. Site attachmentand return patternscan accountfor the incidenceof polygyny in this population,but probablycould not give rise to high levelsof polygynyor large harems. More informationis neededto evaluatethe influenceof site fidelity on mating decisionsof femalesin other polygynouspasserine species. Received 5 November 1985, accepted I May 1986. RESEARCHon the evolution of polygyny in In the original formulationof the polygyny passerinebirds has focusedon speciesin which threshold model, male quality and territory polygynyis common(see Wittenberger 1979, quality were assumedto be highly correlated Vehrenkamp and Bradbury 1984 for reviews). (Orians 1969). Weatherhead and Robertson Relatively little attention has been directed at (1977, 1979) suggestedthat when male quality understandingthe occurrenceof polygynyat and territory quality are not highly correlated, low levels in primarily monogamousspecies females might choose mates based on male (Ford 1983).I attemptedto identify factorsthat quality alone.Although the detailsof their the- causelow levels of polygyny in a population ory have been disputed(Heisler 1981, Searcy of the BlackpollWarbler (Dendroicastriata). and Yasukawa1981, Wittenberger 1981), their Early researchon mating systemsin territo- ideas stimulated studies of the role of male rial passerine birds demonstrated that the characteristicsas well asterritory parametersin number of matesacquired by a male was cor- influencing female mate choice. related with the resources he defended. This I evaluatedthe role of five territory-quality led to the formulationof the polygynythresh- parametersand two male-qualityparameters in old model, which explainshow differencesin determining mating statusin a population of territory quality can give rise to polygyny in the Blackpoll Warbler. The mating status of territorial passerinebirds (Verner 1964, Verner malescould be explainedby male and territory and Willson 1966, Orians 1969). The model was parameters,but femalesettling datescould not. incorporatedinto a moregeneral theory on the The reproductive successof monogamousfe- evolutionof matingsystems (Emlen and Oring males was correlated with male arrival dates. 1977).In the model,female fitness is presumed The polygyny thresholdmodel assumesthat to be a functionof accessto resources.Bigamy femalesare free to mate with any male. This occurswhen differencesin territoryquality are implies that they survey all opportunities large enough that a female can have higher availableat the time of settling before choos- fitnessif sheshares resources on a high-quality ing. In reality, the tendency to return to a fa- territory of a mated male than if she matesmo- miliar site often constrains the choices of both nogamouslyon a low-qualityterritory. males and females that have bred previously 782 The Auk 103: 782-790. October 1986 October1986] FemaleSite Fidelity and Polygyny 783 (Hinde 1956, Greenwood 1980, Oring 1982). 35 male-yearsof data was obtained.These data rep- Female Blackpoll Warblers tend to breed near resent 25 different males and 22 different females. To their former nest sites. Local sex ratios are distinguishthe matesof bigamousmales, I term the sometimesskewed by differential returns of first female to initiate her nest the "primary" female males and females. Given these two factors, the and the secondthe "secondary"female. If differencesin the quality of the breeding situa- benefitsof early nest initiation may causefe- tion give rise to polygynous matings, variation in males to mate with mated males. I assess the measuresof quality should explain (1) the division costsand benefitsto femalesof mating biga- of malesinto mating-statuscategories, (2) the settling mouslyunder these circumstances.Finally, I patterns of the first females on all territories, and (3) suggestthat high nest successmay make the the reproductive successof females within mating- island a favorableplace to breed, enhancing statuscategories (Vehrenkamp and Bradbury 1984). the importanceof site tenacity in maintaining To get sample sizes large enough to test these pre- a breeding spot there. dictions,it was necessaryto pool the information for 1980-1982. STUDY AREA AND METHODS Beginning in 1980, I used the following parame- ters as measuresof territory quality: (1) territory size, The study was conductedin a 35-ha spruce-firfor- (2) coniferdensity, (3) total conifers,(4) large conifer est on Kent Island, New Brunswick,during May- density, and (5) total large conifers.The birds forage August 1979-1982. Kent is an 80-ha island located 20 at midcanopy in conifers up to 10 m tall. Microlepi- km southeast of the Maine-New Brunswick border dopteralarvae that are inside needlesand closedbuds in the Bayof Fundy.The vegetationof the islandhas are an important part of the diet (Eliason unpubl. been describedby McCain (1975).Blackpoll Warblers data). Becauseof these factors, direct assessmentof used only the forestedparts of the island. food on eachterritory proved to be beyond the scope Birdswere netted and marked with unique com- of this project.Male age and arrival dateswere used binations of three plastic color bands and one num- as measuresof male quality. bered aluminum band. Blackpoll Warblers are sex- I determinedterritory size by tracing the outlines ually dichromatic,and therefore males and females of the territories with a plane planimeter. For this were distinguishedreadily. During the 4 yearsof the analysisI used territory maps for the 2-week period study41 adultsand 173nestlings were color-banded. when mostfemales settled on territories.In this pop- For the purpose of analysis, "old" birds are those ulation, more than 90%of foraging by BlackpollWar- known to be 2 years old or older. Four males and 1 biers was done in conifers(Eliason unpubl. data), so female that were banded before 1979 provided some conifer density on the territory was taken as a mea- known-age birds at the outset.Fourteen birds band- sure of availableforaging space.Tree densitieswere ed as nestlingsreturned in subsequentyears, provid- determined by counting trees (by speciesand size ing a group of known-ageyearlings. class)in 10 x 10-m plots. The number of plots used Arrival dates of both sexeswere determined by to sample each territory was proportional to the size patrolling the studyarea daily, looking and listening of the territory, so that sampling intensities were for birds. I use the term "settling date" for the first equal(about 6% of territory area).The third param- day a bird was seen on a territory on which it sub- eter, total number of conifers per territory, is the sequently bred. Territory boundaries were deter- productof territory size and coniferdensity. Forag- mined by noting male singing postsand sitesof in- ing observationsalso indicated that coniferssmaller teractionbetween males. Points were plottedon maps than 5 m tall and 8 cm in diameterat breastheight of the studyarea made from aerialphotographs. Each were usedinfrequently by the birds. The large co- week maps of all territories were drawn by connect- nifer parameterswere calculatedby including only ing observationpoints accumulated during the week. trees above that minimum size. After the territoryestablishment period, I visitedeach Male mating-statuscategories were: bigamous, mo- territory at least once every 2 days for a minimum of nogamous,or unmated. Logisticregression was used 1-2 h to determinemating relationships,monitor in- to constructthe best possiblemodel to explain vari- teractionsbetween mates,and follow reproductive ancein the mating statusof males.The five territory- chronology. quality parametersand two male-qualityparameters A maximum of 15 males held territories on the were used as potential explanatoryvariables. island in any year. The first field seasonwas devoted An explanatoryregression model was first sought to a feasibilitystudy during which the breedingbi- for the responsecategories of mated vs. unmated. ology of 5 males and their mates was studied. The The analysiswas then repeatedwithin the matedcat- breeding habitat is dense,and nesting femalesare egoryfor bigamousvs. monogamous.The procedure secretive. To be reliable, I concentrated on the activ- was performedwith the computerprogram LOGIT ities of 8-12 malesper year in 1980-1982.A total of (K. Larntz and S. Weisberg,Dept.
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