New Techniques for Time-Activity Studies of Avian Flocks in View-Restricted Habitats
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j. Field Ornithol., 60(3):388-396 NEW TECHNIQUES FOR TIME-ACTIVITY STUDIES OF AVIAN FLOCKS IN VIEW-RESTRICTED HABITATS MICHAEL P. LOSITO,• RALPH E. MIRARCHI, AND GUY A. BALDASSARRE 1 Departmentof Zoologyand WildlifeScience AlabamaAgricultural Experiment Station Auburn University,Alabama 36849-5414 USA Abstract.--Focal-animal sampling was comparedto a newly developedtechnique, focal- switchsampling, to evaluatetime budgetsof Mourning Doves(Zenaida macroura) in habitats where the field of view was restricted.Focal-switch sampling included a formula employed to weighthabitat use and testfor restrictionsof habitatstructure on behavior,and a standard waiting period to decidewhen to end sampling or continue pursuit of lost flocks. Focal- animal samplingbiased estimates of activebehavior downward, whereas estimates of inactive behaviorwere similar for both methods.Focal-switch sampling increased research efficiency by 12%. The standardwaiting period saved24% of samplesfrom prematuretermination and reducedobserver bias by maintaining equal sampling effort per flock. Weighting of habitat use also reducedsampling bias. Focal-switch sampling is recommendedfor use in conjunctionwith focal-animalsampling when samplingin view-restrictedhabitats. NUEVAS TP•CNICASPARA EL ESTUDIO DE ACTIVIDADES DE CONGREGACIONES DE AVES EN HABITATS CON VISIBILIDAD RESTRINGIDA Resumen.--Lat6cnica de observaci6ndirecta de animales(focal-animal) es comparada con un nuevom6todo en dondese evaluan los presupuestosde actividadesen Zenaidamacroura en habitatsen dondeel campode visi6nestaba restringido. La nuevat6cnica (cambio-focal) incluyeuna f6rmulapara "pesar"la utilizaci6nde habitaty medirlas restriccionesque imponenla estructuradel habitaten la conducta.Incluye ademfis,un periodode espera estandarizadoque permite al observadordecidir cuando terminar susobservaciones o conti- nuar las mismas.E1 muestreo de un soloanimal (focal-animal)tiene un sesgopara estimar la conductaactira, mientras que los estimados para conducta inactiva rueton similares para ambosm6todos. E1 nuevo m6todo increment6 la eficienciadel estudioen un 12%.E1 periodo de esperaestandarizado evit6 que se terminara prematuramente con el 24% de lasmuestras observadasy redujo el sesgoal mantenetel mismoesfuerzo de muestreopot agregaci6nde aves.Se recomienda la utilizaci6ndel nuevo m6todo en conjunci6n con el previamentedescrito cuandose muestreeen habitatdonde la visibilidadpueda set obstruida. There are many effectiveand widely used methodsto study activity budgetsof avian flocks (e.g., Altmann 1974, Baldassarreet al. 1987). Most studieshave focusedon speciesthat congregatein open habitats (e.g., shorelineand open water) where large numbersof individualscan be sampledreadily (Maxson and Bernstein1984, Paulus 1984a, Quinlan and Baldassarre1984, Tamisier 1976). However, problemsof visibility biasarise in studiesof speciesin densehabitats (Bradley 1985), because the investigatoris confrontedwith samplingconstraints that often cause data variability and biases(Kessel 1976, Verner 1965). • Currentaddress: Environmental and ForestBiology, College of EnvironmentalScience and Forestry,State University of New York,Syracuse, New York 13210. 388 Vol.60, No. 3 NewTime-Activity Methods [389 Focal-animalsampling is the observationof an individualfor a pre- determinedperiod of time (Altmann 1974). It may be the best method for detailedanalysis of behaviorand the only methodavailable to sample small numbersof individuals.Focal-animal samplingis thereforeappro- priate for studiesof speciesin densevegetation where the number of potentialsample individuals is frequentlyreduced by vegetation. A major, unresolvedproblem with focal-animalsampling occurs when focal individualsdisappear from view (hereafterreferred to as "out-of- view";see Lehner 1979). For example,Verner (1965) arbitrarilyrecorded Marsh Wren (Cistothoruspalustris) out-of-view data as feeding,whereas Maxson (1977) broadlymeasured Ruffed Grouse(Bonasa umbellus) ac- tivity by radiotelemetry.Other observers(e.g., Paulus 1984b, Rave 1987) had biasedresults because they were unableto focal-sampleflying birds. Suchlimitations may discouragetime-activity studies of flockingupland species.Accordingly, techniques must be developedto addressproblems associatedwith: (1) focal individuals disappearingfrom view; (2) the amount of time a focal individual should be observed;(3) the spatial distributionof flock individualsamong heterogeneoushabitats; and (4) the eliminationof observer-expectancybias when a sampleflock departs beforethe terminationof the sampleperiod. We reporta new technique,focal-switch sampling, developed to address thesesampling problems during an activity-budgetstudy of Mourning Dove (Zenaida macroura) flocks in northern Alabama. We also report someresults of the activity study by relating behavioralfrequencies to broad habitat categorieswhile employingboth focal-animal and focal- switch sampling. METHODS Focal-switchsampling was testedagainst focal-animal sampling using radio-fitted hatching-year(HY) Mourning Doves to lead observersto dove flocks on the Swan Creek Wildlife Management Area, Wheeler National Wildlife Refuge, and surroundingareas in northern Alabama. Dove flockswere locatedfrom July to September1987. Radio-marked individualswere not observedfor behavioraldata to eliminate potential bias inducedby the radio transmitter.Flocks were sampledduring three observationperiods (OPi = sunriseto 3.5 h after sunrise,OP2 = 3.5 h after sunriseto 3.5 h before sunset,OP3 = 3.5 h before sunsetto sunset) to block-out variation causedby differing periods of Mourning Dove activity (Deuver and Fatora 1968). The closestreadily visible individual in the flock was selectedas the first bird to be sampled(Altmann 1974). Two observers(one per method) collecteddata on eachsample flock. Each methodwas usedby all four observersduring the study.The total data include30.5 h (OP1 -- 10 h, 10 min; OP2 = 10 h; OP3 = 10 h, 20 min). Behavioral categorieswere: feeding,locomotion, comfort, resting, alert, and out-of-view. Focal-switchsampling, unlike focal-animalsampling, allowed switch- ing to a new focal individual (nearestneighbor) if the original focal 390] M. P. Lositoet al. J. Field Ornithol. Summer 1989 T^BLE 1. Demonstrationof weighting focal samplesaccording to habitat use (Y•) where nA = 50, nB = 25, nc = 25, N = 100, T = 60 min, and f= 5 min.a Focal-unit (sample {s}; no sample {--}) Habitat 1 2 3 4 5 6 7 8 9 10 11 12 A s -- -- s -- -- s -- -- s s s B -- s -- -- s -- -- s .... C -- -- s -- -- s -- -- s -- -- -- a y, = (n,/N) x (T/J) where: Y, = numberof focal-units(f) sampledin habitat i; n, = number of individuals in habitat z; N = total number of individuals in flock; T = duration of sampleperiod;f = duration focal-unit. Then YA= n,•/N x T/f = 50/100 x 60/5 = 6, YB=3, and Yc = 3. individual went out-of-view. To avoid observer-expectancybias (Balph and Romesburg1986), two consecutiveout-of-view recordingswere al- lowed before focal-switching.Focal-switching also was restrictedto in- dividuals in the same habitat as the out-of-view individual. This controlled the effectsof habitat structureon behavior.For example, if a focal in- dividual in habitat A moved to habitat B and was lost from view, focal- switchingwas allowed only on individualsin habitat B. The null hypothesistested was that the two methodsresulted in the collectionof the sameamount of data per behavioralcategory during a given unit time. We used Hotelling's multivariate t-tests (SAS Institute, Inc. 1985) to determinethe overall differencein resultsobtained by use of the two methods.Because the data were normally distributed,we used pairedt-tests (SAS Institute,Inc. 1985) to comparebehavioral frequencies betweenmethods for each observationperiod. A probability level of 0.05 was used for all statistical tests. The sampleperiod (T), the maximum time limit for samplinga flock, was dividedinto focal-units(f) where: T = 60 min, andf = 5 min. The focal-unitwas the predeterminedtime within which behaviorof the focal individual(s)was sampledinstantaneously every 20 s (Altmann 1974) as dictatedby a tape recorderused as a metronome(Paulus 1984a, 1984b). The number of completefocal-units (focal-switching not needed)versus the number of incompletefocal-units (focal-switchingnecessary) was testedfor equalityusing paired t-tests(SAS Institute, Inc. 1985). To beginsampling a flock,the observersestimated visually the number of individualsusing major habitat types (e.g., upland field, forestcanopy, willow thicket) and focal-unitswere distributedproportionately (Table 1). Habitat weightingwas testedby comparingactivities sampled in perch (e.g., forestcanopy + willow thicket canopy)versus non-perch habitats (e.g.,upland field + mudflat). Unpaired t-tests(SAS Institute,Inc. 1985) were usedto testthe null hypothesisthat the two habitat structuresaffected behaviorequally. When a sampleflock departed before termination of the sampleperiod, we allowed a standardperiod of 10 min for a new samplingsituation to Vol.60, No. 3 NewTime-Activity Methods [391 TABLE2. Number of instantaneousrecordings (mean + SE) of each Mourning Dove behaviorcollected using focal-animaland focal-switchmethods simultaneously during eachobservation period, northern Alabama, 1987. Method b Observation Behavioral category perioda Focal-animal Focal-switch Locomotion 1 4.5 + 1.1' 9.5 + 2.3* 2 2.6 + 1.3' 5.2 + 1.9' 3 6.9 _+ 1.4' 11.9 _+ 2.4* Feeding 1 21.0 + 5.0* 32.3 + 8.5* 2 1.4 + 0.5 4.5 + 2.3 3 38.6 + 8.0 49.0 ___11.0 Resting 1 13.8 + 3.1 12.3 +