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Nesting Ecology of Scrub Jays in Chico, California

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Nesting Ecology of Scrub Jays in Chico, California NESTING ECOLOGY OF SCRUB JAYS IN CHICO, CALIFORNIA LYMAN V. RITTER, PSW Forestand Range ExperimentStation, ForestrySciences Laboratory,2081 E. SierraAve., Fresno,California 93710 Twelve races of the Scrub Jay (Aphelocoma coerulescens)occupy a geographicrange extendingfrom southernMexico northwardover mostof westernNorth America to southernWashington and Idaho. Another race is isolated in central Florida (American Ornithologists'Union 1957). Good quantitativeinformation on nestingecology is availableonly for the Florida race(A. c. coerulescens;Woolfenden 1973, 1975, Stallcupand Woolfenden 1978). Atwood (1978, 1980b) describedthe breedingbiology of the Santa Cruz IslandScrub Jay (A. c. insularis),an insularpopulation, but presented few quantitativedata on nestingsuccess. Anecdotal information on nestingby other races can be found in Bent (1946), Hardy (1961), Brown (1963), Stewart et al. (1972) and Verbeek (1973). This paper documentsbasic reproductiveparameters of ScrubJays (A. c. superciliosa)in the Sacramento Valley of northern California. STUDY AREA AND METHODS I conductedthe study along upper Lindo Channel and Big Chico Creek, partlywithin BidwellPark in the northeasternsection of Chico, Butte County, California. Valley Oak woodland typified the habitat, with an overstoryof Valley Oak (Quercus Iobata), California Sycamore (Platanus racemosa), BlackWalnut (Juglanshindsii), Box Elder (Acernegundo), Interior Live Oak (Quercus wislizenii)and California Laurel (Umbellularia californica). The understory consisted of California Wild Grape (Vitis californica), Blue Elderberry($ambucus mexicana), Poison Oak (Toxicodendrondiversiloba) and California Blackberry (Rubus vitifolius).Valley Oak woodlands are characteristicallyheterogeneous, with oaks dispersedin groveswith interven- ing openings.Data were alsocollected from other nestingsites in the suburbs of Chico. All studylocations were at an elevationof about 70 m (230 ft). Jays were capturedduring the autumn and early winter monthsin ground trapsbaited with acornsfrom Valley Oaks. Each capturedjay was weighed, aged (Pitelka 1945), banded, and marked with patagialflags to facilitatein- dividual recognition(see Hester 1963). The sizesof the territoryand home rangewere determinedby plotting;oca- tionson a field map and connectingthe outermostpoints, to form the largest polygonpossible, as describedby Odum and Kuenzler (1955). Areas within the polygonswere measuredwith a planimeter.The locationsplotted while determiningthe sizeof territorieswere sitesof boundarydisputes during the breedingseason. In the case of home range measurements,the pair's loca- tionsthroughout the year were plotted. Observationswere made on 119 nestsfrom January 1971 throughJune 1974. Nestswere locatedby searchinglikely spotsand by observingnest- buildingor food-carryingbehavior. Nest height and nestingsubstrate were recorded at each active nest. I also recorded the date of initiation of each Western Birds: 14:147-158, 1983 147 SCRUB JAY NESTING ECOLOGY clutch, clutchsize, incubationperiod, numbersof eggshatched and young fledged,and fledgingperiod or causeof failureof a clutch.To obtainthese data, I visited each nest at 3- to 5-day intervalsand watched selectednests from blinds. Incubation periods were determined only for nests whose historieswere carefullyobserved from beginningto end. Nest visitswere kept briefto minimizealtering the normal nestingactivity or nestpredation. Nestingsuccess was calculated using "egg-day" and "nestling-day"as units of exposure(Mayfield 1975). Calculationsof nestingsuccess assumed a 5-day egg-layingperiod, an 18-day incubationperiod and a 16-daynestling period. When calculating"egg-day" and "nestling-day,"losses were assumedto have occurredmidway through the intervalbetween visits to the nest. Johnson's (1979) methodwas usedto developstandard errors of daily mortalityrates of eggsand nestlings.Furthermore, his ratio test was usedto statisticallycom- pare mortalityrates between stages of the nestingcycle, betweeninitial and renest attempts,and between years. Hatching successwas determinedfor neststhat were found beforehatching and that remainedundisturbed through this phase of the incubationperiod. Nest failure was generallyclassified as causedby eitherpredation or weather,depending on the appearanceof the nest and its contents. RESULTS AND DISCUSSION SOCIAL INTERACTIONS AND TERRITORIALITY Thirty-sixadult jays and 66 birdsof the year were banded and marked, mostlyin 1970-1972. Trapping and field observationsof socialinteractions were concentratedalong upper Lindo Channel, whereone or both members of sixadjacent pairs of territoryholders were marked. BreedingScrub Jays in Chico appearto be resident,monogamous for life, and to recognizediscrete territorialboundaries only duringthe breedingseason. All activityduring the breedingperiod was confinedto the defendedarea, meetingcriteria of the type "A" territoryof Hinde (1956). Three pairs of ScrubJays, which were markedin the fall of 1970 and observedregularly until June 1974, occupied the generalvicinity of their capturelocations throughout that period. Two of those pairs held territorieswith almost identical boundariesthrough four breedingseasons. Other studieshave suggestedthat ScrubJays in the West maintainperma- nent pair bondsand year-roundterritories, although "territory" is not defined in thesestudies (Pitelka 1951; Hardy 1961; Brown 1963; Verbeek 1973; At- wood 1978, 1980a, 1980b). FloridaScrub Jays exhibit cooperative breeding and have year-roundterritories, with size dependingon the numberin the family unit (Stallcup and Woolfenden 1978, Woolfenden and Fitzpatrick 1978). The intensityof territorialdisputes during the nonbreedingperiod seems to differ among localities.Verbeek (1973) found intense territorial defensethroughout the year in Scrub Jays (A. c. californica)in Monterey County, California. Atwood (1980a, 1980b) noted a relaxationin territory defenseduring the nonbreedingseason among Santa Cruz IslandScrub Jays. And among Scrub Jays (A. c. oocleptica)in Berkeley, California, Brown (1963) describeda dominancehierarchy in which, during the nonbreeding 148 SCRUB JAY NESTING ECOLOGY season,territory holders were dominant over all otherjays that occurred in the area of their breedingterritory. Duringautumn and winter,Scrub Jays in Chicosometimes form loose ag- gregationsof up to 10 birds,which may be partlyfamilial. For example, three nestlingsfrom the samenest that were banded during the springof 1971 were recapturedwithin 100 m of the nestthe following October to December.First- yearbirds seemed to makeup a largeproportion of flockmembers. Aggrega-• tionsof jays often were observedin old territoriesof known breedersand were looselyassociated with the resident pair. The nonbreedingranges of adjacent territoryholders, where one or bothbirds of the residentpairs were marked, oftenoverlapped, but rarelywere they observedto aggregatelonger than about 15 minutes.Flocks of jays, as noted by Atwood (1980a), were not observedduring the breedingseason. Aggressiveencounters were frequentduring the nonbreedingseason, usuallyover food. It wascommon to seea jaychase and supplant another that wascarrying food. Suchsupplanting flights typically followed circular paths, ratherthan linear flights typical of mostterritorial disputes. Approximateterritory size in westernmainland and Santa Cruz Island ScrubJay populationswas 2 to 3 ha (Atwood1980b). Florida Scrub Jays unassistedby helpershad territoriesof approximately8 ha. Fiveterritories of ScrubJays in Chicoaveraged 2.2 ha (s.d. = 0.84, range = 1.0 to 3.1). Dur- ing the nonbreedingperiod, marked pairs were seenmost often in the area de'fendedduring the breedingseason. However, considerableextension of theirmovements occurred during the nonbreeding season. The home ranges of two pairsthroughout a 24-monthperiod covered 4.9 ha and 5.4 ha. NEST-SITE PREFERENCE Four plant species,California Wild Grape, Blue Elderberry,Interior Live Oak and Coffeeberry,provided nest cover for 84% of all nests.Use of these coverspecies suggests that concealmentof the nestwas a primaryfactor in selectionof plantspecies for nestplacement. Nest-building typically occurred beforemost species of deciduousshrubs and treeshad renewedtheir foliage. More nestswere builtin CaliforniaWild Grape vinesthan in any other cover species,and althoughthis is a deciduousspecies, branch tangles provided protectionfrom possiblepredation before new foliageprovided concealment (but see sectionon nestingsuccess). Blue Elderberryis among the firstof the deciduousplants to renewits foliage in the spring,and it wastypically in full leaf by the time jays builtnests in it. InteriorLive Oak and Coffeeberryare evergreen.All 19 nestsfound in suburbanhabitat were placedin evergreen shrubs and trees. Variationamong nesting sites was great. Nestswere placedin terminal br.anches, the forksof branches,the forksof treetrunks, on lateralbranches, and in vines. Figure1 showsthe percentages of nestsin variousheight intervals above the ground.Taking the 4 yearstogether, just over 50% of the nestswere placed from 2 to 4 m abovethe ground.The relativelylarge percentage of nests foundabove 6 m in 1972 maybe due to the samplingvariation present with smallsample sizes. The heightof 119 nestsranged from 0.6 to 15.2 m, witha 149 SCRUB JAY NESTING ECOLOGY mean of 3.4 rn (s.d. = 1.94 rn). The mean nestheights were not significantly different between years (ANOVA). I found no significantdifference in nest heightof first(• = 2.8 rn, s.d. = 1.14)and second(•
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