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Nest Predation and Nest-Site Selection of a Western Population of the Hermit Thrush ’

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Nest Predation and Nest-Site Selection of a Western Population of the Hermit Thrush ’ The Condor90~5 1-57 0 The CooperOrnithological Society 1988 NEST PREDATION AND NEST-SITE SELECTION OF A WESTERN POPULATION OF THE HERMIT THRUSH ’ THOMAS E. MARTIN AND JAMES J. ROPER Department ofZoology, Arizona State University,Tempe, AZ 85287 Abstract. Audubon’sHermit Thrushes(Cutharus guttatus auduboni) in centralArizona havea low nestingsuccess (7 to 20%)due almost exclusively to nestpredation. We examine the siteschosen for nestingand comparethem to nonusesites randomly selected within the vegetationtypes associated with nests.Hermit Thrush nest sitesdiffer from nonusesites primarily in that nest siteshave more small (l- to 3-m tall) white firs (Abiesconcolor) in the patch(5-m radiuscircle) surrounding the nest.Hermit Thrushesnest almost exclusively in smallwhite firs and theydo not foragein or near them.Hermit Thrushesmay selectnest sitesthat have a largenumber of otherpotential nest sites(i.e., small white firs) near the nest becausepredation risk is therebyreduced. Indeed, nestswith a high probability of predationwere surrounded by a lowerdensity of smallwhite firs than moresuccessful nests. However,low predationnests also were more concealed than high predation nests. Nest- site selection appears to be a function of characteristicsin the immediate vicinity of the nest (concealment,overhead cover, nest orientation), but also on a larger scalesurrounding the nest. Considerationofnest-site selectionon this largerscale may castlight on the question of whether nest sites limit territory and habitat selectionby birds. Key words: Daily mortality; nestconcealment; nest orientation; nest predation; nest-site selection;nest-patch selection; nesting success. INTRODUCTION Nest-site selection is closely tied to fitness be- Best 1985). However, the nest patch may be causeof the effectson offspring production (e.g., equally important to selectionof sitesfor nesting. see Martin, in press a). Consequently, nest-site Nonrandom selection of nest patches has only choice should be molded by nest-site character- been examined a few times (i.e., MacKenzie and istics that influence the number and quality of Sealy 198 1, Clark et al. 1983, Petersen and Best young that can be successfullyfledged. Habitat 1985) and none of these studies attempted to characteristicsthat influence probability of nest relate vegetation characteristicsof the nest patch predation may be particularly important because to nesting success.Yet, studiesin aquatic systems nest predation often is the primary source of provide a basis for expecting foliage density to nesting mortality for a wide range of bird species influence predation probability at a scaleas large (Ricklefs 1969). or larger than the nest patch; increasesin vege- Nests may be affectedby habitat at two spatial tation density in foraging patches of aquatic scales:(1) the nest site (characteristicswithin the predators often reduce predation risk by con- immediate vicinity of the nest) and (2) the nest cealing prey or inhibiting predator search effi- patch (characteristics of the habitat patch sur- ciency (e.g., Crowder and Cooper 1982, Ander- rounding the nest). Previous work has focused son 1984, Cook and Streams 1984, Leber 1985). on the nest site, examining effects of overhead Indeed, Bowman and Harris (1980) found rac- cover on energy costs (e.g., Calder 1973, Wals- coon (Procyon lotor) foraging efficiency de- berg and King 1978, Walsberg 1981), nest ori- creased,search time increased,and fewer clutch- entation relative to solar exposure (e.g., Giesen es of bird eggswere found in enclosures where et al. 1980, Schafer 1980, Cannings and Threlfall understory foliage density was artificially in- 198 1, Zerba and Morton 1983), and effects of creased.Thus, foliage density in the nest patch nestconcealment on probability ofpredation (e.g., may impede random and intentional nest dis- Keppie and Herzog 1978, Nolan 1978, Best and covery by inhibiting transmission of chemical, auditory, or visual cues. An alternative hypoth- esis that may operate simultaneously or inde- ’ ’ Received 13 February 1987. Final acceptance8 pendently is that predation probability may de- July 1987. creasewith increasesin density of the particular 52 THOMAS E. MARTIN AND JAMES J. ROPER foliage types that are used as nest sites; such in- ence of parents, eggs,nestlings) of each nest were creasesmay reflect the number of potential nest recorded every 3 to 5 days. Nests that fledged at sitesthat predators must examine which reduces least one young were considered successful.Ob- their chancesof finding the actual nest (Martin, servations of fledging, fledglingsnear the nest, or unpubl.). These alternatives can be addressedby parents feeding new fledglingsin the general area examining effectsof the nest patch on probability of the nest were taken as evidence of a successful of nest predation and by specifically examining nest. Depredation was assumedwhen the nest or predation probability as a function of numbers eggsor nestlings (when too young to fledge) dis- of potential nest sites surrounding nests. appeared. Although most nestswere found prior In this paper, we present data on nesting suc- to onset of incubation, some nests were not and, cessand nest-site and patch choice of Audubon’s so, nest successand mortality were calculated Hermit Thrush (Catharusguttatus auduboni) and using the Mayfield method (Mayfield 196 1, 1975) then examine nesting success and predation as modified by Johnson (1979) and Hensler and probability relative to the numbers of potential Nichols (198 1). Half the number of days between nest sites and other habitat characteristics as- subsequent visits over which a nest was depre- sociated with actual nests. As we will show, Au- dated was added to the number of previous days dubon’s Hermit Thrush is particularly appro- the nest survived to obtain the total number of priate for this analysis becausenest-tree selection days a nest survived. Tests of differencesin nest- in central Arizona is highly specific,which allows ing successwere conducted using the z-test de- reasonableestimates of numbers of potential nest scribed in Hensler and Nichols (198 1). sites. Four nestsin 1985 and nine nestsin 1986 that were found by observing parents were never vis- STUDY AREA AND METHODS ited more closely than 10 m to check effects of Study sitesare drainagesdominated by big tooth human visitation on probability of predation be- maple (Acer grandidentatum) in the understory causesuch effects can sometimes obscurethe im- and located on the Mogollon Rim in Central portanceof nest concealment (Westmoreland and Arizona at 2,300 m elevation. These drainages Best 1985). These nests were checked from 10 vary in area and numbers of coexisting bird m or more using binoculars and observations of specieswith a total of 29 speciesrecorded (Mar- parental activity at the nest to determine whether tin, in press b). These sites have a mixed over- or not the nest was active. When no activity was story with ponderosa pine (Pinus ponderosa), found, the nest was approached to verify pre- white fir (Abiesconcolor), douglas-fir (Pseudotsu- dation. ga menziesii), white pine (Pinus strobiformis), Nest-site characteristics were measured after quaking aspen (Populus tremuloides),and Gam- termination of nesting. Plant speciesused as the be1 oak (Quercus gambelii). Saplings of canopy nesting substrate, height of the nest above the trees, plus maple and New Mexican locust (Ro- ground and height of the nest tree were measured binia neomexicana)are the dominant understory by meter stick, or by ocular estimation in the woody species(see Martin [in pressb] for further three casesof large trees. Orientation of the nest description). These drainages contrast with sur- relative to the main stem was recorded in 45” rounding forest which is primarily characterized octants. Nest concealment was indexed by esti- by open ponderosa pine with Gambel oak in the mating percent foliage cover in a 25-cm circle subcanopy and little understory vegetation. centered on the nest from a distance of 1 m from Red squirrels(Tamiasciurus hudsonicus), gray- above and from the side in each of the four car- neck chipmunks (Eutamias cinereicollis), long- dinal directions. Minimum (MINSC) and aver- tailed weasels (Mustela frenata), House Wrens age (AVESC) side cover were used for analyses. (Troglodytes aedon), and Steller’s Jays (Cyano- Habitat characteristicswithin a 5-m radius cir- citta stelleri) are present as possible nest preda- cle around each nest were measured at all nests tors on Hermit Thrushes (Martin, unpubl. data). in 1985 and 1986 and for a few nests in 1984. From mid-May to early July in 1984 through Included within this sampling were a few nests 1986, 15 maple drainageswere searchedfor Her- that were found in the first 2 weeks of the breed- mit Thrush nests. Nests were located by observ- ing seasonbut which never contained eggs.These ing parents with nesting material or by simply nests were probably depredated before being searchingthe vegetation. Date and status (pres- found by human observers.However, thesenests NEST-SITE SELECTION 53 were assigned a status of “unknown” and not provided more adequate sample sizes for anal- included in analysesof habitat characteristicsrel- yses. ative to nesting success.Habitat variables mea- Analysis of variance was used to test univar- suredwithin the circlesincluded numbers ofwhite iate differences in habitat variables between firs between 1 and 3 m tall (WFSM) because groups. Variables that discriminated between Hermit Thrushes almost
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