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NEST-SITE SELECTION BY SAGE THRASHERS IN SOUTHEASTERN IDAHO PDF

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GreatBasinNaturalist51(3). 1991,pp.261-266 NEST-SITE SELECTION BYSAGETHRASHERS IN SOUTHEASTERNIDAHO KrmiclliL. i^tc.scn'andl.oiiis15. Hcst" — AbstRACT. Nfstsitesseleetetll)\ SatieTiiiasiiers{Oiroscoptcsiiioiilaiiiis)insoutheasternIdahowereeliaraeter- izedandeomparedwithavailablehabitat. Mierolialjitatswitliin5niofnestshadtallerandmorea^nreiiatedsluni)sand lessbaref^roundthanthestud\ areainj^eneral. Bigsa.nebrush{Artemisiatridcntatawyomin^oisis)plantsusedlor nestingwere tallerthan averageavailable shrubs, had greaterfoliage density, were more often living, and more frequentlyhadbranehesandfoliagewithin30cmoftheground. Nestplacementwasspecificwithrespecttorelative nestheigiitanddistancefromthetopandperimeterofthesupportshrub. SageThrashersdisproportionatelyused easterlyexposuresandunderusedwesterlyexposuresfortheirnests. Keywords: SV/^cThrasher.bi>isaiichrusfi,shnihsteppc IdaJio,iicst-sitcselection. Choiceofnestsitesrepresentshabitatselec- Sti'dvAreaandMethods tionon asmall spatial scale(e.g., MacKenzie etal. 1982, StanflPerand Best 1986, Bekoffet The study area, consistingof25haofsage- asli.tes19m8a7)y.gAovvaeirlnabtilhietcyoampnodsistniitoanbiolfitbyirodfcnoems-t pblrauisnh1s1hkrumbssotuetphpeofoHnotwhee,uIpdpaehro,SnisaakdemiRniivse-r mnnities and may strncture species-habitat tered by the U.S. Department ofEnergy as relationshipsas muchasdo theavailabilityof partoftheIdahoNationalEngineeringLabo- food and otherresources (Martin 1988). Pre- ratory (INEL). Vegetation is dominated by sumbably, patterns of nest-site selection big sagebrush (Artemisia tridentata wijomin- haveevolvedasaresultofselectivepressures ^ensis), green rabbitbrush (ChrysotJiammts that have ma.\imized nesting success. In viscidifloriis), and scattered bunchgrasses. particular, predation (e.g.. Murphy 1983, Forbs are sparse and ephemeral, litter accu- cBellilmeast-eIs(lee.sg.,anPdlePsizcezmyannska198169)78,anFdermgiucsroon- bmaurlea.tiIonns19a8r0e,sfcoaunrt,6.a2n5d-hmaupclhotosf(t2h5e0gmroxun2d5i0s and Siegfried 1989) have been implicated as m) were established and gridded throughout major agents in molding nest-site selection at25-mintervalswithsteelstakesaffi.xedwith strategies. coloredplasticflagging. TheSageThrasher{Oreoscoptesmontanus) Data were collected during the breeding is a common breeding bird in sagebrush- seasonsof1980-1984. Nests were locatedby shrubsteppe communities of the western using a rope-drag technique (Petersen and United States (Wiens and Rotenberry 1981). Best 1985) to flush adults from their nests. Previous studies of nest-site selection by Nests also were discovered by observing Sage Thrashers (Reynolds and Rich 1978, adults feeding young, and many nests were Rich1978, 1980, Reynolds1981)werelimited foundincidentaltootheractivities. Eachyear in that only a few parameters (e.g., nest several nests were discovered after being height, substrate height) were investigated. abandoned, and these were included in the Ourobjectivewastoprovideamorethorough sample. Nests that had deteriorated or were analysis ofSage Thrasher nest-site selection, suspected to have been built before the cur- includingcharacterization ofnest-site micro- rentyearwereexcluded. habitat, nest substrates, and nest placement Habitat characteristics of each plot were within substrates. Further, we measured as- (Quantified in June each year by using 20 X pects ofavailable habitatwith which to com- 50-cm (juadrats (Daubenmire 1959) and line parenestsites. intercept (Canfield 1941). Each year one to 'DepartmentofBiology,MonmouthCollege.Mnrinioiitli,Illi! "Departmentof.AnimalEcolo,g\',IowaStateUni\ersit\.,\nies 261 262 GreatBasinNaturalist [Volume51 four quadrats were placed 2.5 or 5 ni from our measurements, we took the measure- each grid marker (in different locations ments of rabbitbrush, grasses, forbs, litter, each year); percent coverage ofrabhitbrush, and bare ground soon after a nest had been grasses, forbs, litter, and bare ground was located. However, thesemeasurementswere estimated. Additionally, the height of all not recorded fornests abandoned before be- sagebrush plants included totally orpartially inglocated. Datawereaveragedforeachnest withinquadratswasrecorded, andthecondi- so that nests were the observational units in tion of each sagebrush plant was noted as statistical analyses. Wealso recorded thefol- dead, 25%, 50%, 75%, or 100% living. We lowingmeasurements: (1)heightofeach nest also qualitatively estimated densityoffoliage (groundtonestrim),(2)distancefromthenest on the livingportions ofshrubsas low, inter- rimtothetopofthesupportshrub, (3)short- mediate, or high. Canopy continuity (pres- esthorizontaldistancefrom thecenterofthe ence or absence of gaps more than 20 cm nest to the perimeter ofthe support shrub, across)ofeachsagebrushplantwasrecorded, and(4)compassorientationofthenestrelative and the profile (presence or absence ofany tothecenterofthe supportshrub. Wecalcu- branches or foliage within 30 cm of the lated relative nest height as the ratio ofnest ground) ofsagebrush plants greater than 40 heighttotheheightofthesupportshruband cmtallwasnoted. expressedasapercentage. Canopy coverage and dispersion of sage- T tests and chi-square analyses were used brushwereestimatedbylineintercept. Each to compare nest-site features with those of year 10-25 samples were taken near grid the study area in general. For most t tests, markers on each plot, different grid markers variances did notdiffersignificantlybetween being used each year. These samples were the two groups. When variances differed, regularlyspacedtoprovidean even distribu- we used the t' test (Sokal and Rohlf 1969: tion of sampling effort across the plot. For 374-375), which relaxes the assumption of each sample we recorded line intercept of variancehomogeneity.Therewerefewsignif- sagebrush and distance between adjacent icant variations among years in nest-site fea- sagebrushplants thatwere interceptedalong turesusedbythrashersorinhabitatfeatures a5-mtapeextendingineachofthefourcardi- on the study area. Accordingly, data were nal compass directions. For each sample the pooledforallyears. coefficientofvariationofintershrubdistances wasusedasanindexofdispersion;thegreater Results theindex, the moreclumpedtheshrubs. We Nest-SiteMicrohabitat averaged the habitat data (exclusive ofindi- vidual shrub measurements) for each grid SageThrasherschosenestingareasinwhich markerandusedthegridmarkersasobserva- sagebrushplantswere significantlytallerand tionalunitsinstatisticalanalyses. Forindivid- moreclumped thanon thestudyareain gen- ual shrub measurements (height, condition, eral (Table 1). Percent coverages of sage- etc.),theshrubsweretheobservationalunits. brush, rabbitbrush, grasses, forbs, and litter To characterize actual nest sites, we re- within 5 m of thrasher nests were slightly cordedthesamedataforshrubs supportinga greater than those on the study area in gen- nest as for those occurring within (]uadrats. eral. Althoughnoneofthesepatternswassig- We also estimated canopy coverage and dis- nificant, their cumulative effect resulted in persionofsagebrushalonga5-mtapeextend- significantK less bare groundnearnests than ingfrom the nest in each ofthe fourcardinal on therestofthestud\-area. compassdirections. Further,werecordedthe NestSubstrates height ofeach sagebrush plant intercepted. From 1981 to 1984 we estimated coverage of All nests were located in or beneath (on rabbitbrush, grasses, forbs, litter, and bare the groimd) sagebrush plants. Shrubs se- groundin20 x 50-cm(juadratsplaced2.5and lectedfornestingaveragedsignificantlytaller 5mfromeachnestineachofthefourcardinal than those representative of the studv area directions. To minimize the potential con- {t 15.7,df=5079,p<.001). Moreover,the founding effects ofvegetation change occur- rangeofshrubsizesusedfornestingwasmuch ring between the time ofnest initiation and narrower than that of the available shrubs 1991 Sa(;i;'1'iiiv\.siii;kNKsi-srii-;Ski.kction 263 Hal)italcharactc )rSaKt''niiaslK' the SD). \arial) Nr; StiicK Sagebrushheight(cm) 49± 12(53)-' 41 ± 18(5028) Sagebnislidispersion' 86 ± 19(53) 77 ± 22(401) Sagebrushc()\erage(%) 23± 10(53) 22 ± 11(401) Rabbitbrushcoverage(%) 6±4 (34) 5± 7 (484) Grasscoverage(%) 9± 9 (34) 8 ± 9 (484) Forbcoverage(%) 4 ± 7 (34) 3 ± 5 (484) Littercoverage(%) 7 ± 3 (34) 6 ± 6 (484) Bareground(%) 50 ± 12(34) 55 ± 21(484) '^Nestingniicn,lial>italM\v 1(;;< 05,/test). ''Coenkieiitofvariationoil Further, mostavailableshrubsbearingfoliage had intermediate foliage density. Although two-thirds ofthe nest shrubs also had inter- mediate foliage density, shrubs with high foliage density were used disproportionately by thrashers as nest substrates, and shrubs with lowfoliagedensitywereusedlittle. The canopycontinuityofshrubsevidentlydidnot influence shrub selection by thrashers; the fre(iuencies with which gaps occurred in the canopies ofnest shrubs and available shrubs were nearly identical. Finally, a significantly Representative Sampk greater than expected proportion of shrubs used for nesting had branches or foliage n=5.028 within30cmoftheground. X±SD=41±18 NestPlacementWithinSubstrates Thrashersplacedtheirnestsdeepwithinor beneath shrubs (Table 3). Nest height aver- aged only slightly more than a third of the substrate height. Further, nests were placed 10 20 30 40 50 60 70 80 90 100>100 horizontally relativelyfarfrom theperimeter Height(cm) andclosetothecenteroftheshrub. Severalof Fig. 1. Frecjuency distributions of heights of Sage these measurementsare noteworthybecause Thrashernestshrubsandarepresentativesampleofsage- oftheirrelativeconstancy;coefficientsofvari- brushshrubsfromthestudyareaingeneral. ation were small for relative nest height and distances from the nest to the perimeterand (Fig. 1). Shrubs less than 50 cm tall consti- thetopoftheshrub. tuted 73% of all available shrubs; yet no The overall pattern of Sage Thrasher nest shrubs in this size range were used as nest orientations (Fig. 2) was not significantly dif- substrates. Indeed, 72% of the nests found ferentfrom auniform distribution (X" = 9.6, were in or under shrubs greater than 70 cm 7df, p = .22), buteasterly(NE, E, SE)expo- tall; shrubs in this size range composed only sures were more prevalent than westerly 7%ofallavailableshrubs. (NW, W, SW)exposures. Acomparisonofall Shrubs used for nestingby Sage Thrashers easterly orientations (combined) to all west- differedfromavailableshrubsinseveralother erly orientations was significant (X" = 6.1, respects. Nearlvall shrubs usedbythrashers Idf, p = .02). were 75% or 100% living (Table 2). This dif- feredmarkedlyfrom thedistribution ofavail- Discussion able shrubs among the condition classes, in Sage Thrashers were selective in their whichmanyshrubswerelessthan75%living. choice ofnest sites. In microhabitats chosen 264 GreatBasinNaturalist [Volume51 Table2. ComparisonsofSageThrashernestslirubswitharepresentatixesampleofsagebrushshrubsfrom the studyareaingeneral.Valuesrepresentfreciuenciesofoecurreneeandpercentages(inparentheses)ofthetotal. Nest Representati\( Variabk sample Condition <.01 Dead 1(2) 1109(22) 25%living 312(6) 50%hving 2(4) 617(12) 75%living 13(24) 709(14) 100%living 37(70) 2269(45) Foliagedensity 5.87 .05 Low 3(6) 515(13) Intermediate 35(67) 2762(71) High 14(27) 632(16) Canopycontinuity 0.01 Withgaps 22(42) 2041(41) Withoutgaps 31(58) 2959(59) Profile 4.91 .03 Full' 49(92) 1875(80) Notfull 4(8) 461(20) rfoli Table3. Aspectsofnestplacementb\'SageThrashers. Variable' 1991] SACKTllKASIIKHNKSrsriKSKLKCTION 265 obscurcMiient and cover for thrashers is suii;- differinman\'respectsfromtheaverageavail- gested also by observations oi platforms of able habitat. Third, variation in theheight oi twigs in slirul) canopies ai)ove some nests shrubschosen(coefficientofvariation 21%, (Ricli 19<S(), 1985; personalol)servation). Hicli compared to44% forthe representativesam- (1980)i)elie\edtliatsomesuclicanopieswere pleof shrubs from the studyarea)andin sev- old nests, but none that we observed ap- eral ofthe nest placement variables is small pearedso. In one instance thatweobservcnl, (Table 3). Moreover, the mean nest shrub twigs were placed in the shrubcanopy about height is similar to means reported in other oneweekafterthenestwasinitiated. studies (Revnolds and Rich 1978, Rich Aspectsofnestplacementwithinsubstrates 1980, Reynolds 1981, Castrale 1982). Sage (e.g., nest height, nestorientation) alsohave Thrashers are characteristic of most sage- been found to be related to nesting success brush-dominated rangelands in the United (e.g.. Murphy 1983, Westmoreland and Best States (Wiensand Rotenberry 1981), andbe- 1985) and microclimate amelioration (Hor- cause thrashers have evolved in sagebrush vath 1964, Rich 1978). In particular, nonran- habitat, thespecificityoftheirnest-siteselec- dom nest orientation typically is thought to tion should not be surprising. The patterns reflectresponsesbybirdstoprevailingwinds thatweobservedlikelyhavebeen moldedby or the radiative environment (e.g., Austin alonghistoryofexposuretoaparticularsuite 1976, Petersenand Best 1985, Ferguson and ofselectiveagents. Siegfried 1989). Favoringeasterly andavoid- ing westerly exposures for nests may reflect Acknowledgments attempts by thrashers to maximize exposure to the morning sun, shading from the after- WethankPaulSievertandLindaErickson- noonsun, orboth. Eastwood for assistance in data collection. It is possible, ofcourse, that factors other Several technicians of the Radiological and than piedators or microclimate accounted Environmental Sciences Laboratory (RESL) for the patterns we observed. For example, of INEL also assisted in the field, and the selection of a large shrub for nesting may RESL staff provided on-site transportation denotetheneedforstructuralsupportforthe andlodging. WethankTerrellRich, Kimberly nest.ThetendencyofSageThrasherstoselect A. With, andan anonymous reviewerforcri- microhabitats with large, clumped shrubs ticjuingearlier drafts ofthe manuscript. This mightsimplyreflectthespatialdistributionof studywasfundedbytheOfficeofHealthand areas conducive to robust sagebrush growth. Environmental Research, U.S. Department Thus, selection ofa large shrub for nesting of Energy, and is a contribution from the could, de facto, place the nest in an area INEL Radioecology-Ecology Program. Funds of large, clumped sagebrush. Or, because wereadministeredthroughtheIowaCooper- thrashersforageprimarilyontheground(per- ative Fish and Wildlife Research Unit, U.S. sonal observation), clumped shrubs perhaps Fish and Wildlife Service. This is Journal provide a favorable interspersion of shrubs PaperNo.J-13723oftheIowaAgricidtureand andopeningsforforagingnearthenest. HomeEconomicsExperimentStation,Ames, Although the determinants of nest-site Iowa, ProjectNo. 2468. selection by Sage Thrashers are not known for certain, several lines ofevidence suggest LiteratureCited that thrasher nest-site selection is strongly stereotypic. First, theuseofsagebrushplants Austin.G T. 1976. BehavioraladaptatioiLsoftheX'erdin asnestsubstratesisubiquitous. Revnoldsand tothedesert.Auk93:245-262. Rich (1978), Rich (1978, 1980), and Reynolds Bekokf.M.a.C.Scott,andD.A.Conner. 1987. Non- (1981) also found nests only in or under big randomnest-siteselectioninEveningGrosbeaks. sagebrush plants. Castrale (1982) found one Condor89:819-829. thrasher nest in a juniper ijunipcrus osteo- Belles-Isles.J.,andJ.Picman 1986. Nestinglossesand spenna) tree. To our knowledge, this is the 4ne8s3t-4s8it6e.preferencesinHouseWrens.Condor88: onlydocumentedinstanceofaSageThrasher Cankield.R H 1941.Applicationofthelineinterception nestinanythingbutsagebrush. Second, Sage methodinsamplingrangevegetation. Journalof Thrashers are specific in that their nest sites Forestry39:388-394. 266 GreatBasinNaturalist [Volume51 Castrale,J S 1982. EfiFects oftwo sagebrush control Reynolds,T D 1981.NestingoftheSageThrasher,Sage methods on nonganie birds. Journal of WikUife Sparrow, and Brewer's Sparrow in southeastern Management46:945-952. Idaho.Condor83:61-64. Daubenvmeigreeta,tiRonaFl 1a9n5a9l.ysAis.canNoopryt-hcwoevsetragSecimeentcheod33o:f Reynoldogsy,oTf.tDhe,SaangdeTThrRaischher19{7O8r.eoRsecporpotdeuscmtoinvteaneiciosl)- 43-64. W on the Snake Riverplain in southcentral Idaho. Ferguson,J.VV. H..and R Siegfried 1989. Envi- Auk95:580-582. ronmentalfactorsinfluencingnest-sitepreference RiCH.T 1978.NestplacementinSageThrashers.Wilson inWhite-browed SparrowWeavers (Plocepasser Bulletin90:303. mahali).Condor91:100-107. 1980. Nest placement in Sage Thrashers, Sage HORVATH,O. 1964. SeasonaldifferencesinRufousHum- Sparrows, and Brewer's Sparrows. Wilson Bul- mingbird nest height and their relation to nest letin92:362-368. chmate. Ecology45:235-241. 1985.ASageThrashernestwithconstructedshad- Mackenzie,D.I.,andS.G.Sealy 1981. Nestsiteselec- ingplatform.Murrelet66:18-19. MMMaaarrcttkiiennn,,vaotmnnM1zaeiasag9uTTirsolsn8yn..tnei2ieio-aE,6.tmEfstti.9.obineyZ:,blDNt1oaaaeea7E9o:ipsg4an8nl'stpIe-8s.deo-rs,.a8tlsJtgo:e4eSihmyaH.r.cteIucaAntse6lhG.bi0tn.dioa;ciReutnhnvsCoS2anaadteo2toeprrafa1-nepiaWlc2rradarnety-iteoed,s2weddrrt21egaiaaa92esetsr8n8n3trtie3a8.ido:eanlf.c'nyors3esrGnKfNa1ie.ifosesn0epnt.fs-Dco.gitt3.pbtCns2iuhfapDS1reollrne.duanuealsatetdtdf;vinahaioicotreaaaenienrnM?somaoltnEJucfracbolosantouitmhnhrlird,-d--e-d SWWSoiteKaEsAuNtLfSm,f,oedmsttiARJrrneii.taur.eosenAIiknRltps,o,sDutpaw1,ireaaaSn0Acb.2nnsadFNea:dnd,nnDoW,J7vcfiFcAJ7ieD.lrooN4rTaspRmo-oDn.oeomn7ncaRnniL8imu-nol.s0eMnBntLdc.nuoieBFoetl.istL.nlstuyeB1b.i7nt9eBen7isi6sErgn6t9nctrr.Bog9bpuyel8ipcB1:osr.Dti9gdt1u2oo8is9r36mvc8e.1ie1ae1n9-tl.oN8r2frnMe5y4Hiebs.o.2apsitnab.tr-TWrioidsiht.nsgiaeagtrntieHanesh.apcfuassfhhcbsFheacsiocrrertcteusaa5iesbct1ao.---:s-f MurphyEH,aesrMtm.eirTtnTK1hi9r8nu3gs.bhi.NredCssotnadsnuodcrcoe9ts0hs:er5a1nf-ld5vc7na.etscthienrsg.haCbointdsoorf Wray,T2ti1,o-InI4,1oa.nndneRs.tCi.ngWhsiuctcmeosrseo.fV19e7s9p.erEffSepcatrsroofwsve.geAtuak- 85:208-219.' 96:802-805. Petersen,K L.,andL B Best.1985.Nest-siteselection bySageSparrows.Condor87:217-221. Received8January1991 Fleszczynska,W. K. 1978. Microgeographicprediction Revised29Aphl1991 ofpolvgvnv in the Lark Bunting. Science 201: 935-937'. Accepted15May1991

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