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BREEDING SEASON HABITAT USE and ECOLOGY of MALE NORTHERN PYGMY&Hyphen;OWLS

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BREEDING SEASON HABITAT USE and ECOLOGY of MALE NORTHERN PYGMY&Hyphen;OWLS j. RaptorRes'. 37(2):117-124 ¸ 2003 The Raptor ResearchFoundation, Inc. BREEDING SEASON HABITAT USE AND ECOLOGY OF MALE NORTHERN PYGMY-OWLS ALAN R. GIESE 1 Departmentof Fisheries and Wildlife,Oregon State University, Corvallis, OR 97551 U.S.A. Emc D. FORSMAN U.S.Forest Service, Pacific Northwest Research Station, 3200 SWJeffersonWay, Corvallis, OR 97331 U.S.A. ABSTRACT.--Weexamined habitat use,nest location,diet, and activitypatterns of radio-markedNorthern Pygmy-Owls(Glaucidium gnoma) during four breedingseasons (1994-97) in fragmentedforests on the OlympicPeninsula, Washington. We observedforaging in all availablevegetation categories, but patterns of use were non-random.Structurally diverse and older forestswere most heavilyused, openingsand patchesof saplingsreceived the leastuse, and use of edge wasintermediate. We locatedeight nests,all in woodpecker cavitiesin patches of structurally-diversetbrest. We found no clear evidence that nests were locatednear edges.Northern Pygmy-Owlswere diurnally active,and male owlsdelivered food to femalesduring nest establishmentand incubation,and to both femalesand nestlingsduring brooding. Date of fiedging varied tkom mid-Juneto mid-July.Fledging was synchronous,and minimum estimates of brood size ranged from 1-5. Diet included a mix of small birds, mammals and insects.Our results suggestthat the creationof openingsby clear-cutlogging is unlikely to benefit Northern Pygmy-Owls, and that the replacementof structurallydiverse forests with uniform forestsmay be detrimental. KEYWOP, DS: NorthernPygmy-Owl; Glaucidium gnoma;fragmentation; habitat use. ESTACION REPRODUCTIVA, USO DE HABITAT Y ECOLOGIA DEL BUHO PIGMEO MACHO DEL NORTE RESUMEN.•Examinamosel uso de hibitat, la localizaci6nde nidos,la dietay los patronesde la actividadde bfihos pigmeos(Glaucidium gnoma), durante cuatro estacionesreproductoras (1994-97) en bosquesfrag- mentadosen la peninsulaOlympic, en Washington.Observamos el forrajeo en todaslas categoriasde ve- getaci6ndisponibles, cuyos patrones de uso no estabandeterminados pot el azar.Los bosquesmils viejos y estructuralmentediversos fileron los m•s usados,los clarosy parchesde •rboles fueron los mucho menos usados,la utilizaci6nde bordesfue intermedio.Localizamos ocho nidos,todos en cavidadesde carpinteros en parchesde bosquesestructuralmente diversos. No encontramosninguna evidenciaclara que los nidos estuvieransituados cerca de los bordes.Los bfihospigmeos del norte fueron mis activosdurante el dia, los bfihosmachos entregaron el alimentoalas hembrasdurante el establecimientode nidosy la incubaci6n, tanto a hembrasy pichonesdurante el empollamiento.La fecha de crecimientode plumas en que los polluelosdeNan abandonarel nido vari6 a parfir de mediadosde junio hastamediados de julio. E1 creci- miento de plumasfue sincr6nicoy las estimacionesminimas del tamafiode la nidadafiuctuaron entre 1 y 5. La dieta incluy6una mezclade pijaros,mamifkros e insectospequefios. Nuestros resultados sugieren que espoco probableque la creaci6nde clarospot la tala beneficieal bfiho pigmeodel norte y que el reemplazo de bosquesestructuralmente diversos pot bosquesuniformes puede set perjudicial. [Traducci6n de C6sar Mirquez] Holt et al. (1990) labeled the Northern Pygmy- westernforests remains scant,and published infor- Owl (Glaucidium gnoma) North America's least- mation is often equivocal.Yet western forestshave studied owl. Thirteen years later, detailed infor- experienced decades of intense resource extrac- mation for this locally-common inhabitant of tion pressure(Parry et al. 1983) and it is presently impossibleto estimatehow suchchanges might af- • Presentaddress: Department of Biology,Arizona State fect Northern Pygmy-Owls.Here, we report on University,Tempe, AZ 85287-4601U.S.A.; e-mail address: Northern Pygmy-Owl habitat use and ecology in [email protected] forestsheavily fragmented by clear-cutlogging. 117 118 GmsEAND FORSMAN VOL. 37, NO. 2 The range of the Northern Pygmy-Owlextends many owls each day as possible(typically two to eight) from Alaskato Central America, includingforests and beginningwhere we had finished the day before Thus, eachowl was located at all timesof the day,because from the west coast to the Rocky Mountains a new owl typicallytopped the list each day. Location (Johnsgard1988). They are diurnally active,use times ranged from about an hour before sunriseto about cavitynests, and are generalistpredators of small an hour after sunset. birds,mammals, and insects(Bent 1938,Johnsgard We classifiedhabitat into five vegetationcategories 1988). Northern Pygmy-Owlshave been commonly basedon structuralcharacteristics. The Open-Saplingcat- egoryincluded areas dominated by smallconifer saplings sightedin or near openings,leading to statements (1-4 m tall) intermixed with extensiveareas (>25% cov- that they preferentiallyforaged in openings(Bent er) of bare ground and shrub cover,mostly on recent 1938, AOU 1983, Johnsgard 1988), nested near clear-cuts.The EarlyStem Exclusion category was dominat- edges (Webb 1982, Reynolds et al. 1989), and ed by youngconifers (5-15 m tall) with few openingsin the overstoryand with dense, overlappinglimbs in the might benefit from partial forestclearing (Johns- understory.The Late StemExclusion category was domi- gard 1988). In contrast,others have suggested that natedby medium-sizedconifers (typically 20-40 cm DBH partial forestclearing may be detrimentalto North- and >15 m tall) with a closedcanopy and open under- ern Pygmy-Owls(Marshall 1992). story.The StructurallyDiverse category was characterized Hayward and Garton (1988) used call-response by a wide varietyof tree sizesand a multilayeredcanopy. It consistedprimarily of matureand older forests(typi- surveysto examine resourcepartitioning by small cally >80 yr), but alsoincluded mid-aged stands charac- forest owls in Idaho and concluded that the North- terizedby high structuraldiversity. Finally, the Edgecat- ern Pygmy-Owlwas a habitat generalist.Although egory included all areas within 30 m of an intersection their workis the mostextensive study to date,many between a patch of Open-Sapling,and any of the other questionsremain about habitat useby this species. categories.Thirty meterswas used because pilot-study ob- servationssuggested that foraging flights by Northern We usedradiotelemetry to studyhabitat use for two Pygmy-Owlsrarely exceededthis distance.A foraging behaviors,foraging and nesting.We were specifi- flight wasdefined as any flight which includedor ended callyinterested in whetherpartial forest clearing with an attempt to capture prey. Becausepatchiness in m•ght benefit this species.We alsodescribe home the studyarea waspredominantly the result of clear-cut range size, diet, activitypatterns, nest characteris- logging, patcheswere generallyeasy to delineateand classify.We delineatedpatch boundariesusing ARC/ tics,and nestingbehavior. For the purposesof this INFO (ESRI, Redlands,CA) and digital orthophoto- paper,we hereafteruse 'Northern Pygmy-Owl'and graphs.We visitedeach patch and basedvegetation clas- 'owl' synonymously. sifications for both used and available cover on visual inspection. METHODS We usedcompositional analysis (Aebischer et al. 1993) The studyarea was a mixture of federal,state, and pri- and programRESELECT (Leban 1994; available at http. vate landson the northwestcorner of the OlympicPen- //ces.iisc.ernet.in/hpg/envis/resdocl 120.html) to com- •nsula,Washington. The area was hilly to mountainous pare relativeuse among vegetation categories. Composi- with elevationsranging from 50-1350 m. Mean annual tional analysistreats the individualas the samplingunit, precipitationranged from 150-250 cm. Natural vegeta- accountsfor the unit-sumconstraint of proportions,and tion wasdominated by coniferousforests of westernhem- allowsunique availabilityfor each individual.We usedan lock (Tsugaheterophylla), Douglas-fir (Pseudotsuga menzie- F-testto determine study-widedeviation from random us- szz),western red-cedar (Thuja plicata),and silverfir (Abies age,and pairwiset-tests for differentialuse between veg- amabilis). Valley bottoms typically included variable etation categories. amountsof bigleaf maple (Acermacrophyllum) and red al- Use ratios were calculatedby dividing proportionate der (Alnusrubra) (Hendersonet al. 1989). Even-aged use by proportionate availability.Proportionate use was patchesof primarily Douglas-firresulted from logging definedas the proportionof locationsfor a givenowl in and silviculture. a given vegetation category.All locations were deter- We locatedowls by walkinglogging roads while vocally mined by homing to an owl with a hand-held receiver •m•tatingtheir calls.Surveys were conducted during April until the owl was locatedvisually, or until triangulation and May 1994-97,and were not designedto samplethe indicated that the owl was directly overhead.Locations study area equitably. Rather, we concentrated on areas were mappedin the field on aerial photographsand sub- wherewe had observedowls previously, and surveyed oth- sequentlydigitized using digital orthophotographs.Lo- er areasless intensively. When found, owlswere captured cations of owls <50 m from an active nest were not in- •n mistnets, marked with bandsissued by U.S. Geological cluded in the analysesbecause of the possibilitythat SurveyBird Banding Laboratory;and fitted with back- thoselocations represented nest activityrather than for- pack transmitters(Model BD-2G ca. 2.5 g with harness, aging. Removal of such locationswould introduce bias if Holohil SystemsLtd., Carp, Ontario,
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