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Prey of Breeding Northern Goshawks in Washington

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Prey of Breeding Northern Goshawks in Washington j. RaptorRes. 32(4):297-305 ¸ 1998 The Raptor ResearchFoundation, Inc. PREY OF BREEDING NORTHERN GOSHAWKS IN WASHINGTON JAMESW. WATSON,DAVID W. HAYSAND SEAN P. FINN1 WashingtonDepartment of Fish and Wildlife,600 CapitolWay N., Olympia,WA 98501-1091 U.S.A. PAUL MEEHAN-MARTIN 2 ResearchExperiment Station, 3625 93rd Ave. S.W., Olympia,WA 98502 U.S.A. ABSTRACT.--Weidentified 936 prey from food remains and pellets collected at 82 Northern Goshawk (Accipitergentilis) nest sitesin Washingtonfrom 1986-96. Mammals and birds constitutedhalf of the prey by frequencyand biomassthroughout Washington, although birds were more prevalent(P = 0.050) in the diet of goshawksnesting in the Olympicand Cascademountains of westernWashington (53%), than in the Cascadesof easternWashington (47%). Douglas'squirrels (Tamiasciurus douglasii), grouse (Dendragapusobscurus and Bonasaumbellus), and snowshoehares (Lepusamericanus) were jointly the most frequentlyrepresented prey on the westside (41%) and eastside (54%). Grouseand snowshoehares accountedfor the overwhelmingmajority of prey biomassin theserespective areas (76% and 80%). Relativeto other Northern Goshawkpopulations, goshawks in Washingtonappeared to prey on species from a similar number of genera,but they had a smallerfood-niche breadth and they took larger-sized birds primarily due to their high consumptionof grouse.Northern Goshawksin westernWashington took prey in more equal numbersthan thoseon the eastside. Potential bias from examinationof prey remainswhen comparedto pelletsreinforced the need for inclusionof observationson prey deliveries at nestswhen determining the diet of nesting Northern Goshawks. KEYWORDS: NorthernGoshawk; Accipiter gentilis;diet;, food habits;grouse, hare,, Washington. Presasdel azor en reproducci6n en Washington RESUMEN.--Identificamos936 presasde restosde comida y egagr6pilasrecolectadas en 82 nidos de Acdpitergentills en Washingtondesde 1986-96. Los mamiferos y las avesconstituyeron la mitad de las presaspot frecuenciay biomasaen Washington,aunque las avesprevalecieron (P = 0.050) en la dieta de las montafiasOlympic y Cascadedel oestede Washington(53%), al contrario de las Cascadedel esteWashington (47%). Tamiasdurusdouglasii, Dendragapus obscurus, Bonasa umbellus y Lepusamericanus fueron en conjunto las presasmas representadasen el oeste (41%) y este (54%). Dendragapusobscurus y Lepusamericanus representaron la mayoriade la biomasade presasenestas areas respectivas (76% y 80%). Con relacitn a otras poblacionesde azoresdel notre, los azoresde Washingtonaparentemente depredaron a especiesde un nfimero similar de gtneros, pero tuvieron nichos de alimentacitn de menor tamafio y una media mayor en el peso de las aves,lo cual es el resultadodel alto consumode Dendragrapusobscurus y Bonasa umbellus. Sin embargo,los azoresdel norte en el oestede Washington tuvieron un uso mas equitativode presasque los del este. E1 sesgopotencial del ex/tmen de los restos de presascomparado con las egagrtpilas refuerza la necesidadde incluir observacionespara la deter- minacitn de la dieta de los azores del norte en anidacitn. [Traducci6n de Ctsar Marquez] In the Pacific Northwest, the association of 1994, Woodbridge and Detrich 1994) may be re- Northern Goshawks(Accipiter gentilis) with mature lated to the structural characteristics of stands that forests (Bull and Hohmann 1994, Hargis et al. optimize the availabilityof goshawkprey (Reynolds et al. 1992). Even where preferred prey are abun- dant, structural characteristics of habitat such as • Presentaddress: Raptor ResearchCenter, Department of Biology,Boise State University,1910 UniversityDrive, tree density and understory may reduce prey avail- Boise, ID 83725 U.S.A. ability thereby affecting habitat selection and dis- 2 Presentaddress: Snohomish County Public Works, 2930 tribution of nests (Beier and Drennan 1997, De- Wetmore Ave., Suite 101, Everett, WA 98201 U.S.A. Stefanoand McCloskey1997). 297 298 WATSON ET AL. VOL. 32, NO. 4 In Washington,nesting Northern Goshawksare sampled>2 yr). Most nest treeswere in late successional distributed east and west of the Cascade Moun- forests and were located in national forests (N = 58), national parks (N = 11), private timberland (N = 9), and tains, where there are different climates, forest state land (N = 4). In ownershipsother than national communitiesand potentiallydifferent prey species parks, landscapessurrounding nests and within nesting and prey abundance.In westernWashington, mar- territories were potentially subjectto forest management. itime influences at nest sitesin the Olympic Moun- The actual degree of timber harvestwithin these terri- tories was unknown. tains may further influence prey and diets of nest- The climate in westernWashington is characterizedby ing Northern Goshawks.Diets in this area couldbe mild, wet winters and warm, dry summers. Forests are very different from that of goshawksnesting inland predominantlyDouglas-fir (Pseudotsuga menziesii), western in the west CascadeMountains. In the CoastRange hemlock ( Tsugaheterophylla), and westernredcedar ( Thu- ja plicata).Sitka spruce (Piceasitchensis) is presentat a few of Oregon, for example,dense understory vegeta- lower elevation nest siteson the west side of the Olympic tion and high rainfall are believedto contributeto Peninsula. Forests in the Cascadesof eastern Washing- low populationsof nestinggoshawks due to their ton, a region with cool wintersand hot, dry summers,are negativeeffects on prey availability(Reynolds and dominated by standsof Douglas-fir,ponderosa pine (Pi- Wight 1978, DeStefano and McCloskey1997). nusponderosa), and westernhemlock. For each collection of prey remains,we attempted to There is also the potential for dietary differences identify the minimum number of individualsrepresented in Northern Goshawksnesting in managed stands to specieslevel from pooled occurrencesin food remains in national forestsand private timberland, where and pellets. Matched hair and feather samplesfrom prey timber harvestmay influenceprey availability,and and pelletswere consideredto representthe same indi- vidual, whereas counts of pooled bones and flight feath- in national parks,where there is no timber harvest ers allowed for identification of >1 individual. When we (Crocker-Bedford 1990). found skeletal remains of snowshoehares (Lepusameri- Prey frequency,prey biomass,and food-niche canus)and grouse (Dendragapusobscurus and Bonasaum- breadth are commonlyused to quantify raptor di- bellus),we estimatedthe age of individualsthrough size ets (Marti 1987). Methods of identifying raptor comparisonswith museumspecimens and the degree of bone fusion at joints. Ages of these specieswere used to prey, including direct observation,examination of estimate their respectivecontributions to prey biomass. prey remains,and pellet examinationprovide data Solid remains (e.g., skeletal remains, beaks) were iden- on diet, but each is subject to potential biases tified from museum specimens(University of Washing- (Marti 1987). Here, we examine prey speciesiden- ton, Seattle, or University of Wisconsin,Madison) or tified in prey remainsand pelletsof nestingNorth- identificationkeys (Olsen 1964, 1972). Fur and feathers were matched to museum specimensor descriptionsin ern Goshawksthroughout Washington,with specif- field guides (Peterson 1947, Burt and Grossenheider ic objectivesto contrast prey frequency,biomass 1976). Arthropods were excluded from data analyses. and food-niche breadth for populations of North- Our suspicion was that many insectsfound in prey re- ern Goshawks east and west of the Cascade Moun- mains, particularlybeetles and ants,were consumedin- directly when small mammals and grouse were eaten. tain crest; contrastmajor prey groups among sub- Presence of ants was correlated (r2 = 0.18, P = 0.001) regions of these populations, and in areas with the occurrence of Douglas' squirrels (Tamiasciurus potentially subject to timber harvest and those douglasii)and chipmunks(Tamias spp.) in prey remains, without harvest;identify differencesin prey species and stomach contents of three whole carcassesof Doug- or food-niche breadth peculiar to Washington las' squirrelsand chipmunkscontained numerous beetle Northern Goshawks relative to other areas in shell fragments. Several pellets contained insect frag- ments mixed with fir needles,seeds, and grouseremains. North America; and identify biasesassociated with We reported mammalian and avian prey by frequency identification of prey from remains or pellets. and biomass. Biomass estimates were derived from aver- age weightsof speciesfrom Reynoldsand Meslow(1984) STUDY AREA AND METHODS and other publishedsources (Table 1). Weightswere de- Preyremains and pelletswere collectedat 38 Northern rived for juvenile and adult age classesof snowshoehares Goshawk nest sites in western Washington (16 in the and grouse. For prey for which we could estimatebio- Olympic Mountains,22 in the CascadeMountains) from mass(i.e., not including unidentified birds or mammals), 1986-96, and at 44 nest sites in the Cascades of eastern we calculated the mean weight of avian prey (MWAP), Washington(17 in the centralCascades, 27 in northern mean weight of mammalian prey (MWMP) and mean Cascades)from 1992-96 (Fig. 1). Prey and pellets were weight of total prey (MWTP). Food-nichebreadth was collected from nests, under nest trees, and at plucking calculatedfor prey genera using the following equation posts.Most remainswere collectedincidentally during (Levins 1968): breeding surveysfrom the nestling stage
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