Wood Warbler Populations in the Yolla Bolly Mountains of California
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WOOD WARBLER POPULATIONS IN THE YOLLA BOLLY MOUNTAINS OF CALIFORNIA RUSSELL GREENBERG, 270 KelloggAve., Palo Alto, California 94301 TODD KEELER-WOLF, P.O. Box 866, Corning, California 96021 VIRGINIA KEELER-WOLF, P.O. Box 866, Corning, California 96021 Long-term studiesof bird populationsin California mountainsare conspicuouslyabsent from the literature. While distribution of summer residentspecies is generallywell known, population changesthrough the summerand fall need study. This is certainly the casein the Yol- la Bolly Mountains of the interior north coastranges of California. Hemphill (1952) presentsa fairly complete account of speciesoccur- rence in the southernYolla Bolly Mountainsduring the summer. A banding project carried out in the Yolla Bollys 15 June to 6 October 1973 afforded us the opportunity to observeseasonal changes in bird populations. This paper is a summary of data collected on a closely interrelatedgroup of species,the wood warblers(Parulidae). LOCATION OF STUDY The baseof our researchactivity was a cabinat Howell'sCamp (ele- vation 6200 feet, USGSAnthony Pk. quadrangle)in the Mendocino NationalForest 10 mileseast of the southeastcorner of the Yolla Bolly- MiddleEel WildernessArea, and 30 airlinemiles west of Corning,Te- hamaCounty. Thissite is locatedon the firstmajor north-south ridge west of the SacramentoValley. Steeptopography is a prominentfea- ture of the Yolla Bollysand the risefrom the valleyfloor to Howell's Campis rapid. Thereis a broadzone of interdigitationbetween yellow pineforest and chaparral.The ridgetop aroundHowell's Camp is cov- ered by yellow pine forest dominatedby White Fir (Abies concolor), with IncenseCedar (Libocedrus decurrens), Ponderosa Pine (Pinus pon- derosa), SugarPine (Pinus lambertiana), and mats of Ceanotbuscordu- latus, Ribes lobbii and R. roezlii with some Bitter Cherry (Prunus emarginata).On the slopesof the ridgegrow Ponderosa Pine, Douglas- Fir (Pseudotsugamenziesii), Black Oak (Quercuskelloggii) and Oregon Oak (Q. garryana). Chaparraloccurs to 4,500 feet elevationon the slopesof ThomesCreek Canyonseveral miles to the south. The bandingsite is locatedon a meadowwith two largethickets of MountainAlder (Alnustenuifolia) and scatteredelderberry (Sambucus caerulea)shrubs. A springin the meadowfills a smallreservoir. To the Western Birds 5: 81-90, 1974 81 WARBLER POPULATIONS west lie the high ridgesof the Yolla Bollyswith severalpeaks near 8,000 feet above sealevel. ExtensiveRed Fir (Abiesmagnifica var. shastensis) forestswith Jeffrey Pine (Pinusjeffreyi), White Pine (Pinusmonticola) and locally Foxtail Pine (Pinusbalfouriana) is a major habitat not pre- sent closeto Howell's Camp. High elevationvegetation including White Fir dominatedyellow pine and Red Fir community types is referred to as the boreal zone (after Hemphill 1952). Plant speciesand commun- ity-type classificationare based on Munz(1968). A detailedvegetational analysisof the regionis providedby Keeler-Wolfand Keeler-Wolf (1974). METHODS One mist-net was operated an averageof eight hours per day on 35 days between 2 July and 5 October. During this period nearly 1500 birds were caught;964 were warblers. Due to the absenceof summer rain and scarcity of surfacewater, the spring and elderberriesof the meadowwere powerful attractersof birds..Many speciesof passerines, hummingbirds,and woodpeckerswere seen feedingon the elderberry nectar and berries. The net was located next to a drippingwater trough in an alder thicket. Birds could be seen descendingfrom the sky in numbers into the thicket. The net was nearly as high as the thicket and caught most of the birds going to drink or bathe. The banding samplewas a good representationof the speciesproportions observed in the meadow. Age of warblerswas determinedby degreeof skull ossification,plumage characters, and presenceor absenceof primary and secondarymolt. To supplementour bandingdata sight observationswere made dur- ing our stay 15 June to 5 October1973. Most observationswere made in the Howell's Camp area, but our knowledgeof the statusof many specieswas increasedby extendedtrips in 1973 and in the two previous years into the wildernessarea, includingthe Red Fir forest regionof North and South Yolla Bolly Mountains. RESULTS A summaryof speciesbanded by time period is presentedin Table 1. The warbler flocks in the meadowwere composedof mixed species whose relative abundance varied. To avoid placing undue stresson hourly fluctuationsthe data has been groupedinto more meaningful samplesizes. To illustratelong-term trends we haveorganized our data 82 83 WARBLER POPULATIONS IOO I I I' I I I I I I I I I I I I 90 YELLOW-R{ 8o uJ6o ORANGE-CROWNED c• õo 30 NASHVILLE 20 I HERMIT 0 5 II 3 13 22 29 8 I 20 I 5 JULY AUGUST SEPTEMBER OCT Figure1. Relativeabundance of fourmost frequently caught warbler species, 2 Julyo5October 1973, in YollaBolly Mtns., California. Points plotted represent meandates of bandingperiods given in Table 1. 84 WARBLER POPULATIONS into samplingperiods of irregularlength. The data cannotbe usedto estimatethe size of the populationssampled due to the transientna- ture of the population. Only four warblers(0.4%) were recaptured:1 Yellow-rumpedWarbler (Dendroica coronata auduboni) and 3 Orange- crownedWarblers (Vermivora celata). Three recaptureswere made over three weeksafter the originalcapture. It is clearthat largenomadic populationswere being sampled. In Figure 1, data from Table I has beenconverted to relativefrequency for Orange-crownedWarbler, Nash- ville Warbler (Vermivoraruficapilla), Yellow-rumpedWarbler and Her- mit Warbler (Dendroicaoccidentalis). Data pointsare plotted at mean dateof time period,given in Table 1. Thisgraphically shows the change in speciescomposition of the bandingsample over time. Trendsof the data are discussed below. SPECIES ACCOUNTS The Lutescent Orange-crownedWarbler (V. celata lutescens),a common chap- arral breeder,was presentin large numbersthroughout the boreal regionsat the beginningof the study in June. This speciesincreased in abundancethrough July and began dropping in abundanceslowly in August then rapidly in September. All of the birds bandedwere born that year and goingthrough postjuvenal molt. The percentageof birds havingcompleted postjuvenal molt (except crownwhere molt was completedlast) increasedthrough summerand fall as follows: 2-25 July, 5.6% (n=89); 30 July-14 August, 15.8% (n=95); 15-23 August,57.0% (n=79); 29 August-5October, 91.8% (n=49). Orange-crownedWarblers were abundant in scrub and thickets rangingcommonly into coniferousforests, often high in the canopy. A maximum of 50 was banded 11 August. A small number of birds which ap- pearedto be of the northern races(V. celata oresteraor V. c. celata) were present from 16 Septemberto 2 October. Orange-crownedWarblers were more abundant in 1973 than in the previoustwo years. NashvilleWarblers were heard singingin lower elevationyellow pine forest dur- ing June, but were not observedaround Howell's Camp and other alder thickets until early July. Flocksof this speciesassociated with Orange-crownedWarblers. As summerprogressed Nashville Warblers became relatively and absolutelymore numerous,but they droppedin abundanceearlier than Orange-crownedWarblers and were nearly absentin late August. NashvilleWarblers banded were all hatch- ing yearbirds in variousstages of postjuvenalmolt. The percentageof birdshaving completedpostjuvenal molt (except crown where molt was completedlast) in- creasedthrough the summeras follows: 9-24 July, 14.3% (n=14); 31 July-ll Au- gust, 44.2% (n=43); 1 August-7September, 78.3% (n=23). The maximum banded was 34 on 11 August. Yellow Warbler(Dendroica petecbia) immatures were seenand bandedin small numbersduring September and early October. A maximumof 4 wasbanded on 12 and 17 September.They were alwaysobserved in the meadowalder thickets. Audubon's Yellow-rumped Warblerswere still nesting during June and July when large flocks of Vermivorawere moving through the boreal region. The Yellow-rumpedWarbler is one of the most abundantbreeders in the borealregion. Singingwas commonin June, but stoppedalmost entirely in early July and was 85 WARBLER POPULATIONS replaced by intense nesting activity. Very few Yellow-rumpedWarblers were caughtin July; from 2-12 July only 4 adultsand 5 youngwere banded. However, 41 juvenilesand 5 adults were caughtbetween 23 July and 18 August. Late sum- mer juvenileswere in heavy body molt. After this, Yellow-rumpedWarbler re- mainedcommon in the bandingsample, but only 7 adultswere banded. During late July and throughout August large flocks of Yellow-rumpedWarblers were found, especiallyin coniferousforests in associationwith Dark-eyedJuncos (Junco byemalis oreganus)and Chipping Sparrows(Spizella passerina). Another large influx occurredin September.This, coupledwith rapid dropsin Vermivoraabun- dance,made Yellow-rumpedWarbler the dominantfall warbler,with a maximum of 83 bandedon 1 October. During the fall a few Myrtle Yellow-rumpedWarblers (D.c. coronata) were banded. Black-throatedGray Warbler (Dendroica nigrescens) is a commonnesting species in the PonderosaPine dominated yellow pine and DiggerPine (Pinussabiniana) belts, but a few were caught and seen in the White Fir dominated yellow pine forestsfrom late August through early October. We observedonly a few birds above 5,000 feet elevation before August. All birds banded were hatching-year birds with little conspicuousmolt. A maximum of three was banded on 1 October. Townsend's Warblers (Dendroica townsendi) were seen and banded in small numbersin