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Predation on Northern Cardinal Nests: Does Choice of Nest Site Matter? ’ The Condor96~761-768 0 The CooperOrnithological Society 1994 PREDATION ON NORTHERN CARDINAL NESTS: DOES CHOICE OF NEST SITE MATTER? ’ TAMATHA S. FILLIATER,~ RANDALL BREITWISCH AND PAUL M. NEALEN~ Department of Biology, Universityof Dayton, Dayton, OH 45469-2320 Abstract. The fatesof 121 nestsof NorthernCardinals (Cardinalis cardinalis) in south- westernOhio weredetermined in 1991 and 1992.Success rate wasonly 1So/o, estimated by the Mayfield method.All failed nestswere known or suspectedto haveheen preyed upon. Severalhypotheses to explaindifferences between the locationof successfuland failednests weretested. None of thoseconsidered explained why the contentsof particularnests were taken. We proposethat a high incidenceof predationby a rich guild of nest predators precludesthe existenceof predictablysafe nest sites for cardinals.Instead, cardinals appear simply to be well-adaptedto renestrapidly in responseto the near randomnessof nest predation.A similar adaptationmay characterizemany passerines. Key words: Northern Cardinal: Cardinaliscardinalis; nesting success; nest site selection; nestpredation. INTRODUCTION Bent 1968, Rinser 1973) and builds a new nest Selection of nest site may be of crucial impor- in a different location on the territory for each tance to the reproductive successof birds. Lack nesting attempt. Females also appear to be the (1954) estimated that 75% of all eggsand nest- sex that selectsnest sites (Rinser 1973; however, lings lost from open cup nestsare taken by pred- see Laskey 1944). ators. Ricklefs (1969) estimated predation per- Every passerine studied to date displays some centagesin six passerine speciesand concluded intrapopulational variation in nest site selection. that predation accounted for 55% of egg losses Some studies have revealed patterns in this vari- and 66% of nestling losses.Many aspectsof the ation associatedwith the probability of success nesting behavior of birds appear to be adapta- (see Martin and Roper 1988) while others have tions to avoid predation of the nest contents. not (e.g., Best 1978, Barnard and Markus 1990, Many birds hide their nests or build them in Morton et al. 1993; see also Gottfried and inaccessible sites (Collias and Collias 1984). In Thompson 1978 for an experimental study). We addition, the behavior of parent birds visiting attempted to test predictions of eight hypotheses nestsis notably stealthy (Skutch 1976, Breitwisch for the placement of successful versus unsuc- et al. 1989). Active defenseagainst predatorsalso cessfulnests by cardinals. These hypothesesad- occurs in some speciesof birds, but defense en- dress lossesto predators only; lossesdue to abi- tails a risk of injury to or death of the parent. otic environmental factors are not considered Clearly, birds may be advantaged by selecting here. In addition, our assumption is that pred- sites where nests will be unlikely to be found by ators find nests by actively searching for them predators. rather than by simply watching parent birds at- Northern Cardinals (Cardinalis cardinalis) are tend to the needs of eggsor nestlings (Collias and multibrooded, and females lay up to five clutches Collias 1984, Martin 1988a). in a season,of which as many as four have been The “nest concealment” hypothesis predicts recorded as successfully fledging (Shaver and that nests that are concealed regardless of the Roberts 1930, Laskey 1944). The female usually particular plant structure will be more successful builds the entire nest (Laskey 1944, Bums 1963, than those that are not so concealed. The “nest inaccessibility” hypothesispredicts that neststhat are lessaccessible (but not necessarilybetter con- ’ ’ Received27 December1993. Accepted 29 March cealed) will be more successfulthan those that 1994. are more accessible. Inaccessible nests in this 2 Presentaddress: U.S. ForestService, Klamath Na- tional Forest,37805 Highway 97, Macdoel,CA 96058. study are defined as those over water, in thorny 3 Presentaddress: Department of Biology,Univer- vegetation,among cane (Arundinaria sp., a woody sity of Pennsylvania,Philadelphia, PA 19104-60 18. herb), or at the end of thin branches or twigs (cf. [7611 162 T. S. FILLIATER, R. BREITWISCH AND P. M. NEALEN Collias and Collias 1984). The “nest height” hy- form of predation) increasesfrom forest interior pothesis predicts that nests placed high or low to forest edge (Brittingham and Temple 1983). relative to the frequency distribution of nest The “distance to human activity” hypothesis heights in the population will be more successful predicts that nests placed closer to areas of hu- than those of more typical height. The first pre- man activity will be more successfulthan those diction (high nests are more successful)assumes that are farther away (Collias and Collias 1984). that nests built high in plant crowns are con- Human activity may discouragepredators from cealed better than nests built lower, especially as searching near heavily used trails and in other the breeding season progressesand plants leaf areas frequented by people. We added this hy- out (Nolan 1978) and, in this sense,is a version pothesisand tested it in late 199 1 and 1992 after of the nest concealment hypothesis. Such nests finding that successfulnests tended to be closer may also be found less frequently by or be less to areasof human activity than unsuccessfulnests accessibleto terrestrial predatorsthan lower nests. earlier in 199 1. The second prediction (low nests are more suc- cessful)assumes that aerial predators are of ma- jor importance and are less successfulin either STUDY SITE AND METHODS finding or accessinglow nests.The “mid-height” This study was conducted at the Aullwood Au- hypothesis predicts that nests placed at mid- dubon Center and Farm, located about 15 km height in a tree or shrub will be more successful northwest of Dayton, Ohio (39”52’N and than those placed either higher or lower in the 84”16’W) from April to August 1991 and 1992. plant. This hypothesis assumes that nests built The Aullwood property is an 80 ha sanctuary higher are more easily seen by aerial predators, with habitats in the following proportions: 26% while those built closer to the ground are located croplands and orchards, 17% mature woodlands more easily by mammals and snakes(see Alonso (dominated by beech, Fagus spp.; oaks, Quercus et al. 199 1). The “needle in a haystack” hypoth- spp.; maple, Acer spp.; and buckeye, Aesculus esis predicts that nests placed in a common spe- glabra), 17% secondary growth (e.g., ash, Frax- cies of plant will be more successfulthan those inus spp.; and maple), 16% pasture, 8% meadow, that are placed in uncommon plant species(see 5% prairie, 5% residential area, 2% pine plant- Martin and Roper 1988). If a predator restricts ings, 2% wetlands and 2% wet woods (e.g., ash its search for nests to a few appropriate plant and red maple, Acer rubrum) (J. Ritzenthaler, species, then nests in the most common plant pers. comm.); see Filliater-Lee (1992) for further specieswill be more difficult to find becausethere details on vegetation. Cardinals frequent nearly are more individual plants to search. The “rare all habitats on the property and most densely site” hypothesispredicts that nests placed in un- populate secondarygrowth and edgesof wooded commonly used plant specieswill be more suc- areas. cessful than those placed in commonly used We discovered nests by intensively searching plants. If a predator searchesplant speciescom- at least every other day and by following the monly used for nesting, then it is best to be atyp- “chipping” vocalizations of adult cardinals ical and nest elsewhere. This hypothesis differs (Lemon 1968, Montgomerie and Weatherhead from the others in that successdepends on where 1988). Nearly all nests were observed daily to other members of the population are nesting (as record progress. Many nests were observed proposed among species for communities of through 10 x 40 binoculars at a distance of 5- nesting birds by Martin 1988b). The “edge dis- 15 m. We recorded nest height and tree or shrub tance” hypothesispredicts that nestsfarther from height with a meter stick to 0.1 m where possible a habitat edge (defined here as where a closed and estimated where necessary (nests in dense habitat changesobviously to another more open multiflora rose [Rosa multiflora] or higher than habitat, e.g., forest to field) will be more suc- several meters). In addition, in late 199 1 and in cessful than those placed closer to an edge. The 1992, we estimated distance to the closest area distance to habitat edge is important because ofhuman activity to 0.5 m. Human activity areas some mammalian and avian predators actively were defined as trails (most are heavily used) and search near these edges(Gates and Gysel 1978). other areasfrequented by people. Although some Furthermore, Brown-headed Cowbird (Molo- clearings were foci of human activity, distance thrus ater) parasitism (which can be viewed as a to the nearestclearing for any particular nest may CARDINAL NEST SITES AND NEST PREDATION 763 have been different from that to the closestarea central tendencies. Kolmogorov-Smirnov of human activity. 2-sample tests (D,,, value reported) were used We recorded nest visibility from six vantage to compare shapes of frequency distributions points (above, below, and from four horizontal (Siegel and Castellan 198 1). Spearman rank cor- directions [N, S, E, W]) at a distance of l-2 m relations (r, value reported) were used for tests using the following classification scheme: “visi- of monotonic relationship. Results are reported ble” (visible from five or six vantage points), as significant if they are associatedwith an alpha “ambiguous” (visible from three or four), and value ofP < 0.05. All testswere two-tailed. Means “not visible” (visible from none to two). Al- f SD are reported for descriptive statistics. though this division is somewhat subjective, two We also conducted a seriesof eight parametric observers agreed on the visibility of each nest. discriminant function analyses(SAS 1989) of nest The details of placement of each nest in veg- site variables to compare successfulwith failed etation were recorded in order to address the nests.
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