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Roost Site Selection of Great Horned Owls in Relation to Black Fly Activity: an Anti-Parasite Behavior?1

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Roost Site Selection of Great Horned Owls in Relation to Black Fly Activity: an Anti-Parasite Behavior?1 950 SHORT COMMUNICATIONS SAUER,J.R.,AND S. DROEGE. 1992. Geographical pat- impacts of nest predation and brood parasitism on terns in population tends of Neotropical migrants seasonal fecundity in songbirds. Conserv. Biol. in North America, p. 30±68. In J. M. Hagan III 13:46±57. and D. W. Johnston [EDS.], Ecology and conser- WOODWORTH, B. 1999. Modeling population dynamics vation of Neotropical migrant landbirds. Smith- of a songbird exposed to parasitism and predation son. Inst. Press, Washington, DC. and evaluating management options. Conserv. SCHMIDT,K.A.,AND C. J. WHELAN. 1999. The relative Biol. 13:67±76. Downloaded from https://academic.oup.com/condor/article/102/4/950/5562970 by 59662000 user on 13 February 2021 The Condor 102:950±955 q The Cooper Ornithological Society 2000 ROOST SITE SELECTION OF GREAT HORNED OWLS IN RELATION TO BLACK FLY ACTIVITY: AN ANTI-PARASITE BEHAVIOR?1 CHRISTOPH ROHNER2 AND CHARLES J. KREBS Department of Zoology, University of British Columbia, Vancouver, B.C., Canada, V6T 1Z4 D. BRUCE HUNTER Department of Pathology, Ontario Veterinary College, University of Guelph, Guelph, Ontario, Canada, N1G 2W1 DOUGLAS C. CURRIE Centre for Biodiversity and Conservation Biology, Royal Ontario Museum, 100 Queen's Park, Toronto, Ontario, Canada, M5S 2C6 Abstract. We document a shift in roosting behavior owls, Great Horned Owls, Leucocytozoon, parasitism, of Great Horned Owls (Bubo virginianus) from winter roosting behavior. and late spring to summer. During summer, Great Horned Owls roosted near the ground or exposed on Ornithophilic species of black ¯ies (Simuliidae) can open ground, whereas they chose concealed perches at reduce the ®tness of a host in a variety of ways (Mùller mid-canopy level for the rest of the year as typical for 1990, Bennett et al. 1993). For example, black ¯ies are forest owls. This shift of roosting behavior coincided a vector of Leucocytozoon, a protozoon causing avian with the emergence of ornithophilic black ¯ies, which malaria (Desser and Bennett 1993). As an example of transmit avian malaria (Leucocytozoon spp.). The shift possible consequences of disease, higher loads of Leu- in roosting behavior was consistent with measurements cocytozoon ziemanni were associated with lower clutch of parasite exposure at different habitat positions. sizes in female Tengmalm's Owls Aegolius funereus Black ¯y activity was highest at mid-canopy level, and (KorpimaÈki et al. 1993). Detrimental effects may be almost no black ¯ies were active on open ground. directly caused by anemia. Blood extraction by the ec- Ground-roosting was not caused by poorly developed toparasite alone (or in concert with the blood parasite) ¯ying capability of juveniles, because solitarily-roost- may signi®cantly contribute to anemia (Fitch et al. ing adult owls showed the same behavioral shift in a 1946, Richner et al. 1993, Hunter et al. 1997). In ad- second year of study. Black ¯ies and avian malaria are dition, there may be a cost of increased investment in widely distributed, and the effect of the vertical distri- immunological defense (Toft 1991, Richner et al. bution of these parasites in forests on roosting, nesting, 1995). and foraging of sylvatic birds deserves further study. We studied the response of Great Horned Owls Key words: black ¯ies, Bubo virginianus, forest (Bubo virginianus) to the 10-year population cycle of snowshoe hares Lepus americanus in the boreal forest from 1989±1992 (Krebs et al. 1995, Rohner 1996). Great Horned Owls are large and long-lived predators 1 Received 8 November 1999. Accepted 17 May feeding mainly on lagomorphs, they defend long-term 2000. territories, and are widely distributed in North and 2 Current address: Department of Renewable Re- South America (Voous 1988, Donazar et al. 1989, sources, University of Alberta, Edmonton, Alberta, Rohner 1995, 1997). Several demographic parameters Canada, T6G 2H1, e-mail: christoph.rohner@ in Great Horned Owls were strongly affected by the ualberta.ca population cycle of snowshoe hares (Rohner 1996). SHORT COMMUNICATIONS 951 For example, juvenile survival was very high during study were collected in 1991 and 1992. The study area the peak of the hare cycle but dropped in the ®rst year comprised 350 km2 of the Shakwak Trench, a broad of prey decline in 1991. Anemia, caused by Leucocy- glacial valley bounded by alpine areas to the northwest tozoon ziemanni and feeding by black ¯ies, was iden- and the southeast. The valley bottom averages about ti®ed as a major cause of mortality of juvenile Great 900 m above sea level and is covered mostly with Horned Owls in our study area (Rohner and Hunter spruce forest Picea glauca, shrub thickets Salix spp., 1996, Hunter et al. 1997). some aspen forest Populus tremuloides, grassy mead- In this paper, we report on the roost site selection ows with low shrub Betula glandulosa, old burns, es- of juvenile and adult Great Horned Owls in response kers, marshes, small lakes, and ponds. to varying densities of black ¯ies. Initially, we were During 1988±1992, we equipped a total of 21 adult Downloaded from https://academic.oup.com/condor/article/102/4/950/5562970 by 59662000 user on 13 February 2021 puzzled to observe that Great Horned Owls roosted on owls and 55 owlets with radio-transmitters, which the ground during the post-¯edging stage, often ex- were attached with a shoulder harness (details in Roh- posed at sites in the sun. Forest owls are known to ner 1996, 1997). Radios on the backs of owls were roost concealed in trees, and adult Great Horned Owls hidden in feathers and weighed 50 g (, 5% of body always roosted well-hidden and on perches above 4± mass, Kenward 1985). Because a position-speci®c 5 m in winter and early spring (Rohner and Doyle mercury switch was installed, an observer was able to 1992b). After learning about black ¯ies and avian ma- identify whether an owl was perched or ¯ying. laria as agents of juvenile mortality in our study sys- Obtaining data on owl roost sites was extremely dif- tem (Rohner and Hunter 1996, Hunter et al. 1997), we ®cult, because visibility was limited in the coniferous investigated the hypothesis of whether Great Horned boreal forest and because owls were elusive and did Owls modify their roosting behavior to escape these not use traditional roost sites (Rohner 1996). We lo- parasites. We predicted that exposed roost sites on the cated roost sites by carefully approaching radio- ground would be host refugia, because some studies marked owls during daytime with hand-held antenna, indicate that ornithophilic black ¯ies feeding on forest receiver, and binoculars. Once a roosting owl was spot- birds prefer dense vegetation at mid-canopy level ted, perch heights were estimated in meters, and a site where they are most likely to encounter their hosts was scored as exposed or in cover depending upon (Bennett 1960, Hunter 1990). whether dense vegetation was present or absent within Black ¯ies emerge in spring when minimum daily a radius of 1 m of the roosting owl. In order to avoid temperatures rise above freezing. Subsequently, they disturbance, most roost sites were examined later in swarm and mate over open water, then switch to nec- detail. Observations only were used when an owl had tar-feeding to gain resources for dispersal, undertake not changed position (according to the motion-sensi- dispersal or migratory ¯ights to speci®c habitats for tive switch of the radio-transmitter) when it was ap- blood-feeding, and then return to streams and rivers proached. Usually, independent evidence such as for egg-laying, with one to several generations per year feathers, feces, pellets, or claw marks con®rmed the (Bennett 1960, Sutcliffe 1986, Currie 1997). In early locations as roost sites that had been used for at least May, our study area was still covered by snow, and several hours. During 1991±1992, a total of 68 roost lake ice did not break up before the end of May. Based sites were located. All data from individual family on more southerly reports on black ¯y phenology, members were pooled and used as one data point for maximum black ¯y activity in the study area was ex- each territory to avoid pseudo-replication (Hurlbert pected in late July and early August (Bennett 1960, 1984). Hunter 1990). Ornithophilic black ¯ies were captured using the In order to test the hypothesis that Great Horned method of Bennett (1960). Live Bantam chickens Gal- Owls selected roost sites with lower activity of black lus gallus, which are commonly used as bait for at- ¯ies than elsewhere, we conducted a mensurative ex- tracting black ¯ies, were exposed in cages after their periment (Hurlbert 1984) on the distribution and activ- heads were covered with a hood to prevent the chicken ity of black ¯ies, and we collected information on roost from pecking off any black ¯ies. Exposure cages were sites of Great Horned Owls. We predicted that (1) placed on a white board of 60 3 60 cm, and then black ¯y activity in summer will be highest in vege- hoisted up to different heights in trees with the help tation at mid-canopy level and much lower at exposed of a pulley system. During each trial, we exposed the sites on the ground, (2) as black ¯ies emerge and peak cages for 20 min, and then sealed them with a col- towards summer, Great Horned Owls will shift in their lecting cage of ®ne mesh. Within 30 min after a blood use of roost sites from higher to lower perches and meal, engorged ¯ies had left the chicken and settled from more concealed to more exposed sites. An alter- on the sides of the cage. They were collected with an native hypothesis may explain ground-roosting of owls aspirator and ®nally preserved in 70% ethanol.
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