SEARCH

SEARCH BY CITATION

Keywords:

  • anti-predator strategy;
  • nest defence;
  • nest predation;
  • nest site;
  • parental activity

Summary

  1. Top of page
  2. Summary
  3. Introduction
  4. Methods
  5. Results
  6. Discussion
  7. Conclusions
  8. Acknowledgements
  9. References
  • 1
    A simple model of relative effects of parental behaviour (parents present vs. absent) and nest concealment on probability of nest predation was evaluated by measuring survival of paired natural/artificial nests of four open nesting passerines over 3 years.
  • 2
    The ratio of rodent to corvid predation (i) decreased from yellowhammer (small eggs, ground/near ground nests) through blackcap (small, shrub) to song thrush and blackbird (medium sized, shrub/subcanopy); (ii) was highest in years when rodent abundance peaked – this effect was clear in yellowhammer, detectable in blackcap, but not found in thrushes. An inverse relationship was found between mean annual nest survival and abundance of the major nest predator for each species. Predators differed between poorly (corvids) and well-concealed (rodents) nests in blackcap.
  • 3
    The effects on nest survival differed among species, including: positive effect of parental behaviour combined with neutral effect of concealment (thrushes); independent positive effects of behaviour and concealment (yellowhammer); neutral effect of behaviour combined with positive effect of concealment (blackcap). These patterns are consistent with hypothesis that relatively larger species with conspicuous nests (thrushes) should either engage in more vigorous nest defence or their defence is more efficient, compared with smaller species with less conspicuous nests (blackcap).
  • 4
    The positive relationship between nest concealment and survival resulted either from an effect of nest site per se (yellowhammer) or from an effect of parental behaviour (blackcap) that changed from negative (nest disclosure) to positive (nest defence) between poorly and well-concealed nests, respectively. The view that lack of a within-species relationship between nest concealment and survival (thrushes) results from parental behaviour compensating for predation risk associated with poor nest concealment was not supported.
  • 5
    This study demonstrated (i) multiple interactions among factors influencing the rate of nest predation, both between and within species; (ii) potential bias associated with drawing general conclusions from small-scale experiments.

Introduction

  1. Top of page
  2. Summary
  3. Introduction
  4. Methods
  5. Results
  6. Discussion
  7. Conclusions
  8. Acknowledgements
  9. References

Nest predation is one of the major selective agents shaping avian life histories. Anti-predator strategies evolved by birds include direct effects of parental behaviour (nest defence, sensuMontgomerie & Weatherhead 1988) as well as indirect ones, such as decision where (nest site selection) and when (timing) to breed. Nest defence may be active, including direct attacks, mobbing, distraction display, nest guarding and vocalization, or passive through sitting on the nest (nest attentiveness), thereby hiding the eggs from view and serving to camouflage the nest and to discourage some of the potential predators (Montgomerie & Weatherhead 1988; Martin 1992a). The effectiveness of different kinds of nest defence depends on the bird’s capacity to perceive risk associated with the particular predator (McLean & Rhodes 1991; Burhans 2000; Veen et al. 2000) and capacity to deter or distract it (Blancher & Robertson 1982; Komdeur & Kats 1999; Schmidt et al. 2001a). Apart from cost through allocation of time between nest defence and competing activities (Martin 1992a; Komdeur & Kats 1999), parental activity associated with nesting can be costly because it can attract predator’s attention to the nest (Martin et al. 2000a; Martin, Scott & Menge 2000b). The relative strength of the positive (‘nest defence’) and negative (‘nest disclosure’) effects of parental activity on nest survival in particular bird species is likely to depend on the type of nest predators involved and their foraging behaviour.

Selection of nest sites by birds is supposed to be nonrandom and adaptive with regard to the risk of predation (Martin 1998; Clark & Shutler 1999; but see Misenhelter & Rotenberry 2000 and Willson & Gende 2000). By selecting safe nest sites, birds can either decrease nest encounter rate for incidental nest predators (Vickery, Hunter & Wells 1992; Schmidt, Goheen & Naumann 2001b) or decrease nest detectability for actively searching predators. Among the various nest site characteristics, the concealment by vegetation is probably the most important factor for investigators’ attention. Yet, previous studies yielded conflicting evidence for the expected positive relationship between nest concealment and survival (Martin 1992b; Götmark et al. 1995; Howlett & Stutchbury 1996; Burhans & Thompson 1998).

The lack of consistent patterns in nest predation studies has traditionally been attributed to spatial/temporal variation in abundance of nest predators, or to interactions among the effects of nest site, parental and predator behaviour (Götmark et al. 1995; Hogstad 1995; Murphy, Cummings & Palmer 1997; Söderstrom, Pärt & Rydén 1998; Rangen, Clark & Hobson 1999; Schmidt 1999; Dion, Hobson & Larivière 2000; Martin et al. 2000b; Schmidt et al. 2001b). For example, Cresswell (1997) ascribed the lack of uniform (across species) effect of nest concealment to a compensating effect of parental nest defence operating on the within-species level. This hypothesis predicts that species capable of efficient nest defence will show a weaker relationship between nest concealment and survival than species that cannot deter predators and must rely on passive protection through nest concealment. Assumption implicit in this explanation is that on the within-species level, parent birds can respond to nest site associated risk of predation by altering their behaviour (McLean, Smith & Stewart 1986; Hobson, Bouchart & Sealy 1988; Burhans & Thompson 2001). Ricklefs (1977) found some support for an interspecific relationship between mean nest concealment and intensity of nest defence. Yet, sound interpretation of the observed patterns of predation is often limited by unknown identity of principal nest predators in the particular study system (Söderstrom et al. 1998; Pietz & Granfors 2000; Willson & Gende 2000).

Relative effects of parental behaviour and nest site have often been inferred from comparisons between artificial (without parents) and natural (with parents) nests (Cresswell 1997; Wilson, Brittingham & Goodrich 1998; King et al. 1999; Komdeur & Kats 1999; Davison & Bollinger 2000; Martin et al. 2000b). Considering the diversity of patterns documented in such studies, there has been little effort to interpret them within a conceptual framework permitting formulation of testable hypotheses about the underlying mechanisms (Schmidt 1999; Schmidt et al. 2001a,b). Next I present a simple model of the possible mechanisms behind the patterns in survival of natural and artificial nests (Fig. 1).

image

Figure 1. Model relating nest survival to possible effects of parental behaviour and nest concealment, as reflected in survival of natural (with parents; solid lines and filled points) and experimental (without parents; dashed lines and open points) nests. Points indicate the mean survival rate across all levels of nest concealment. The net effect of parental behaviour can be: (A) neutral – either no effect or antagonistic effects that offset each other; (B) positive –‘nest defence’; (C) negative –‘nest disclosure’. Effect of nest concealment unconfounded by parental behaviour can be: (a, d) neutral; (b, c) positive. Effects of parental behaviour and nest concealment can be: (a, c) independent; (b, d) interacting. Each subplot is identified by column (A–C) and row (a–d) coordinates.

Download figure to PowerPoint

It is assumed that survival of experimental nests reflects effect of nest site (e.g. concealment) regardless of parental behaviour, whereas survival of natural nests reflects combined effects of nest site and behaviour. The positive (nest defence) or negative (nest disclosure) effects of parental behaviour are broadly defined as any effects due to presence of parents at the nest that have positive or negative effect on nest survival. These positive and negative effects may combine, thus yielding the net effect of behaviour that can be neutral (positive and negative effects offset each other), positive or negative. This net effect can be inferred from comparison of the mean survival rates between natural and experimental nests: no difference (neutral effect; Fig. 1A); higher survival of natural nests (positive effect; Fig. 1B); lower survival of natural nests (negative effect; Fig. 1C). Effect of nest concealment unaffected by parental behaviour can be inferred from the relationship shown by experimental nests: neutral effect (Fig. 1a,d); positive effect (Fig. 1b,c). The null model is that neither behaviour nor concealment effect is present (Fig. 1Aa). Any of the behaviour effects may combine with any of the concealment effects, both effects being either independent (Fig. 1a,c) or interacting (Fig. 1b,d). Interaction means that the effect of behaviour is not constant across different levels of nest concealment and can be of two opposite directions. First, parental behaviour can compensate for the effect of nest concealment, yielding no relationship between concealment and survival in natural nests (Fig. 1b). Alternatively, parental behaviour can generate relationship between concealment and survival of natural nests in cases, where effect of nest concealment per se is neutral (Fig. 1d).

The model shows that each of the two basic patterns exhibited by survival of natural nests (Fig. 1a,b vs. c,d) can be explained by several alternative mechanisms. First, lack of relationship between concealment and survival of natural nests (Fig. 1a,b) can result either from no effect of concealment combined with independent effect of parental behaviour (Fig. 1a), or from positive effect of concealment masked by an interacting effect of behaviour (Fig. 1b). Second, the positive relationship between concealment and survival of natural nests (Fig. 1c,d) can result either from positive effect of concealment combined with independent effect of parental behaviour (Fig. 1c), or may be due to effect of behaviour interacting with no effect of concealment (Fig. 1d). The above alternatives generate different predictions for survival of artificial nests relative to that of natural nests, and thus can be evaluated experimentally.

This study was designed to test these alternative hypotheses using data on four open nesting passerines characterized by different body size (presumed correlate of defence potential) and different nest size and placement (correlates of overall nest conspicuousness, accessibility to predators and risk of disclosure through parental activity). The survival of natural and experimental nests of each species was concurrently monitored to separate the relative effects of parental behaviour and nest site. Cresswell’s (1997) hypothesis generates the following predictions for the species under study. Song thrush (Turdus philomelos, Brehm) and blackbird (T. merula, Linnaeus) – relatively high nest defence potential and conspicuous nests: natural nests should survive on average better than experimental nests; there should be no relationship between concealment and survival for natural nests but a positive one for experimental nests (Fig. 1Bb). Blackcap (Sylvia atricapilla, Linnaeus) and yellowhammer (Emberiza citrinella, Linnaeus) – relatively low nest defence potential and concealed nests: there should be no difference in mean survival rates between natural and experimental nests, survival should increase with concealment in both natural and experimental nests (Fig. 1Ac).

Methods

  1. Top of page
  2. Summary
  3. Introduction
  4. Methods
  5. Results
  6. Discussion
  7. Conclusions
  8. Acknowledgements
  9. References

STUDY AREA

The study was conducted in a farmland area of c 83 km2 north of the village Luže (49°54′N, 16°02′E, 240–350 m above sea level) in the Eastern Bohemia, Czech Republic, in 1998–2000. The local landscape is a mosaic of arable land with settlements and remnants (≤25 ha) of deciduous woodland. The tree vegetation is dominated by Fraxinus excelsior, Populus nigra, Alnus glutinosa, Quercus petraea and Salix spp., and the dense shrub layer is formed by Sambucus nigra, Padus avium and Prunus spinosa. The study took place on 22–24 plots (2–10 ha) representative of local woodland habitats and located in distinct habitat patches surrounded by arable land or grassland. The median distance to the nearest neighbouring plot was 750 m, minimum was 300 m.

MODEL SPECIES

Three (1998, 1999) or four (2000; see Data analysis) open-cup nesting passerine species that differ in nest placement (Fig. 2), nest size and egg size and crypsis were studied. Yellowhammer: medium sized nest placed on or close to ground and usually well hidden in herbaceous vegetation; cryptic eggs (21·6 mm × 16·2 mm); only female incubates. Blackcap: small nest placed typically on thin branches of shrubs, 0·5–1·5 m above ground, moderately concealed; cryptic eggs (19·6 mm × 14·7 mm); male and female take about equal share of incubation. Song thrush: conspicuous bulky nest, placed in shrubs or trees, 1·5–4 m above ground, often little concealed; blue-green eggs (27·3 mm × 20·6 mm) contrast with yellowish and unlined nest cup; only female incubates. Blackbird: similar nest as in song thrush but with cup lining; cryptic eggs (29·4 mm × 21·6 mm); only female incubates.

image

Figure 2. Relationship between concealment category and height above ground for active (•) and experimental (▴) nests. Median and interquartile range are shown. Relative frequency of concealment categories for active (left bars) and experimental (right bars) nests shown by histogram. Note that there is a partial overlap between the samples of active and experimental nests (see Methods for samples definition); the total number of physical nests was 201 in yellowhammer, 885 in blackcap and 532 in thrushes.

Download figure to PowerPoint

NATURAL NESTS

Nests were systematically searched for in the shrub and herbaceous vegetation with approximately constant effort from about middle of April until the end of July (Weidinger 2001a). Nest content was inspected every 5th day to determine fate. All successive visits to each individual nest were approximately (±2 h) at the same time of day, thus keeping the exposure time comparable among nests. All nests that failed from reasons other than predation (inclement weather, disturbance by forestry or agriculture) were excluded from the analysis. The following nest site characteristics were recorded on the last visit to a nest: nest supporting plant, height above ground (m), concealment category (subjective score ranked from 0 = low to 4 = high). The latter was obtained as a sum of horizontal and overhead concealment (each scored as: 0 = nest well visible from most directions; 1 = intermediate; 2 = nest not visible in any direction from a distance of c. 2 m). A total of 146 (yellowhammer), 654 (blackcap), 225 (song thrush) and 78 (blackbird; only in 2000) nests were analysed.

EXPERIMENT 1A

Natural nests with known fate (successful or predated) were left in their original position and baited with artificial clutches about 15 days after the natural nesting attempt had terminated. The artificial clutches consisted of four plastic eggs attached to the nest and mimicking the size and coloration of real eggs (for details see Weidinger 2001b). The levels of disturbance to natural and experimental nests were maintained at a similar level. Because natural nests were inspected from close proximity and eggs were handled to determine the stage of incubation, human scent was present on both experimental and natural nests. Nest markers were not used. Experimental nests were exposed for a 15-day period, which was close to the mean duration of the egg stage (laying and incubation) of the natural nests (16 days in yellowhammer and blackcap, 17 days in thrushes), and were checked in 5-day intervals in the same way as the natural nests. A nest was considered depredated when at least one egg was damaged, pulled out of the nests, missing or buried in nest lining. Because of development of vegetation throughout the breeding season, I recorded nest concealment also after termination of each experimental nest; the other nest characteristics remained the same as for the natural nest. Data were collected in 1998 and 1999 from a total of 105 (yellowhammer), 418 (blackcap) and 183 (song thrush) experimental nests.

EXPERIMENT 1B

All procedures were the same as in Experiment 1a, except that the artificial clutches consisted of one Japanese quail (Coturnix japonica) egg not attached to the nest. Data were collected in 2000 from 52 (yellowhammer), 325 (blackcap), 125 (song thrush) and 129 (blackbird) experimental nests.

EXPERIMENT 2

Active nests of blackcap found from the stage of laying (≥2 eggs laid) to the 2nd day after clutch completion (i.e. ≥10 days before hatching) were paired with an experimental nest – undamaged blackcap nests collected on the study plots in the previous breeding season. These nests were baited with a clutch of four tree sparrow (Passer montanus) eggs, the size (mean: 19·1 mm × 14·2 mm) and coloration (degree of crypsis) of which resembled those of blackcap. Experimental nests were placed within 10–15 m of the paired active nest. Nest sites were kept similar within each nest pair: the same species of nest supporting plant or at least similar type of vegetation, height above ground, concealment and microhabitat (e.g. isolated shrub vs. closed shrub layer). Before placing an experimental nest, I tried to make sure that no other nest was active within a radius c. 10 m except the focal blackcap nest. Given the difficulties with satisfying all the above conditions, not all blackcap nests found could be included in this experiment. Paired nests were checked simultaneously in 5-day intervals to determine fate (active nest) or survival over the 15 day exposure period (experimental nests). When either of the paired nests was depredated, I continued visiting the remaining nest. Nest site characteristics were recorded in the same way for both active and experimental nests. In all, 46 (1999) and 74 (2000) nest pairs provided useful data.

PREDATORS

The occurrence of potential avian predators (corvids) was recorded visually throughout the breeding season. A frequency of species occurrence in the total number of plot-visits (528 per year; each lasted about 2 hours) was used as a rough measure of relative abundance. Small mammals were snap-trapped on the study (plus two additional) plots in the second half of August (24 plots × 2 transects × 30 traps × 1 night exposure = 1440 trap-nights per year). One transect was placed along the habitat edge and the other inside the patch within each study plot. No attempt was made to determine predators of natural nests from egg remains or nest appearance (Pietz & Granfors 2000). Predators (classified as bird vs. mammal) of experimental nests could be inferred from tooth or beak marks left in the soft layer of paint on the plastic eggs (Experiment 1a) or from remains of quail eggs (Experiment 1b). In the latter case it was only possible to distinguish eggs attacked by small mammals (unbroken eggs or egg shells with tooth scratches; Marini & Melo 1998) from those destroyed by a large predator. Many quail eggs disappeared from the nests, which prevented any inferences about predators involved.

A study aimed at identification of predators of artificial nests was initiated in 2000 on the same plots where the experiments were running. Automatic cameras (Olympus® AF-10 mini) equipped with a photo-cell trigger were placed at a distance of c. 1·5 m from inactive nests of the model species, baited with plastic and quail eggs in the same way as the experimental nests. The first year of the camera study provided only qualitative results. The frequently documented nest predators included jay (Garrulus glandarius), pine marten (Martes martes), wood/yellow-necked mouse (Apodemus sylvaticus/A. flavicolis) and bank vole (Clethrionomys glareolus). Occasional predators included red squirrel (Sciurus vulgaris), weasel/stoat (Mustela nivalis/M. erminea) and great-spotted woodpecker (Dendrocopos major).

DATA ANALYSIS

The unit of analysis in Experiment 1 was a nest, on which two repeated measurements of survival were made: one for natural nesting attempt (i.e. active nest; A) and the second for the experimental nest (E). I defined two data sets of different extent according to whether only ‘paired nests’ (nests for which both repeated measurements were available) or ‘all nests’ (including those nests with either A or E measurement missing) were included. Paired nests (total n = 882) represent a subset of all nests (total n = 1618). The unit of analysis in Experiment 2 was a pair of simultaneously exposed nests (A and E; total n = 120).

Two response variables were used to measure nest survival. (i) Daily survival rate (‘DSR’= number of successful days divided by the number of exposure days; Aebischer 1999); applicable to all experimental nests and to those natural nests that have been observed active during the incubation stage (from laying of the penultimate egg to hatch day). Number of exposure days was estimated by the ‘Early termination’ variant of the Mayfield method (Manolis, Andersen & Cuthbert 2000) with an exception that the exposure period of successful (i.e. hatched) nests was truncated by the hatching day and not by the last visit before the potential hatching interval. For the purposes of graphical presentation, I extrapolated DSR to a 15-day period (Hensler 1985). (ii) Survival over 15-day exposure period (‘FATE’= binary response coded 0/1); applicable to all experimental nests and to those natural nests found active more than 15 days before the earliest possible fledging date (defined as the day when nestlings were 8 days old). Nests analysed using the FATE response represent a subset of nests analysed using the DSR response.

Simultaneous effects of several predictors on either type of response variable were analysed by fitting generalized linear models, with logit link and binomial error distribution (proc genmod; SAS Institute Inc. 2000), under general guidelines by Crawley (1993) and Aebischer (1999). The original models containing all two- and three-way interactions were reduced by sequentially deleting nonsignificant (P > 0·05) interaction terms. Significance was assessed by the Type III tests based on the generalized score statistic, which is asymptotically χ2 distributed. When a significant interaction of species/year effect with a categorical predictor was detected, the analysis was partitioned for species/years. When a significant interaction with a covariate (date, nest height) was detected, a model with separate slopes was fitted. Significance of terms included in the models derived from the all nests sample was further evaluated by fitting these models to a restricted sample of paired nests.

Data collected in 2000 did not reveal significant differences between song thrush and blackbird in nest site characteristics, mean nest survival and the examined patterns (direction and magnitude of differences or trends). Hence, data on these two species were merged in one category referred to as ‘thrushes’. Conclusions did not change in any way when blackbird was excluded from the analyses, but it was retained to increase sample size and power of the tests. The ‘year’ was included as a predictor to account for an effect of the type of artificial eggs used (Experiment 1) and effects of uncontrolled factors that may have changed among years (see Discussion). The ‘study plot’ was considered as a blocking variable to account for the site-specific effects (e.g. individual predators). It follows from spacing and size of the study plots that each plot in this study probably represented only one pair of corvids, but many individual small mammals, home range of which is much less than plot area. The effect of study plot was not included in the analyses of sparse data sets (yellowhammer and Experiment 2) because of many empty cells in the data table. To simplify analysis, and because an overall ‘concealment’ differed markedly among species (Fig. 2), I dichotomized nest concealment within each species as relatively good (score 4 in yellowhammer, scores 3–4 in blackcap, scores 2–4 in thrushes) or poor (the remaining scores). ‘Height’ of nests above ground was partially redundant with factor ‘species’ (Fig. 2), it was therefore included only in the separate-species models.

The effect size observed in this study was evaluated by estimating the 95% confidence interval on differences between the estimated DSR for A and E nests. To get some impression about statistical power of the tests performed, power analysis (Steiger 1999) was conducted for the simplest case encountered in this study, the McNemar’s test for paired binary responses. Given the number of nests available per species (years pooled), type I error rate = 0·05, power = 0·80, and η (proportion of nests with different outcomes on the two measurement occasions) = 0·4, the minimum detectable difference between A and E nests was estimated at 19% (yellowhammer), 9% (blackcap) and 13% (thrushes) in Experiment 1 and 16% in Experiment 2.

Results

  1. Top of page
  2. Summary
  3. Introduction
  4. Methods
  5. Results
  6. Discussion
  7. Conclusions
  8. Acknowledgements
  9. References

NEST PREDATORS

Jay, probably the only important avian nest predator of the studied species, increased numbers steadily, while abundance of rodents showed more than two-fold fluctuations (Table 1). The relative contribution of rodents to overall predation on experimental nests (Experiment 1) decreased, on average, from yellowhammer (81%) through blackcap (51%) to thrushes (44%). A logistic model relating the type of predator (mammal vs. bird) to species, year and nest concealment (Fig. 3), revealed significant species × year (χ24= 10·1, P = 0·040) and species × concealment (χ22= 9·8, P = 0·007) interactions. Because of relatively sparse data sets, the partitioned analyses were based on exact tests of independence in contingency tables (shown are exact P-values). Mammalian share of total nest predation differed: (i) among species in years of high rodent abundance (1998: P < 0·001; 2000: P = 0·012), but not in the year of low rodent abundance (1999: P = 0·708); (ii) among years in yellowhammer (P = 0·003) and blackcap (P = 0·010), but not in thrushes (P = 0·222); (iii) between nest concealment categories in blackcap (P < 0·001), but not in yellowhammer (P = 0·294) and thrushes (P = 0·326).

Table 1.  Relative abundance of potential nest predators on the study plots. Corvids: frequency of occurrence per 100 plot-visits (528 plot-visits per year); mammals: numbers captured per 100 trap-nights (1440 trap-nights per year)
 199819992000
Jay 6·6 9·111·4
Hooded crow 6·1 4·9 4·2
Magpie 2·1 1·1 1·1
Total corvids14·815·216·7
Wood/yellow-necked mouse21·316·938·3
Bank vole39·710·324·6
Total small mammals67·829·964·1
image

Figure 3. Proportion of predation events on experimental nests (Experiment 1) ascribed to small mammals, according to year and nest concealment (poor = left bars, good = right bars). Annual means (concealment categories combined) shown by points connected by a dashed line. Sample size refer to the number of predation events (= 100%) that allowed predator discrimination. Relative abundance of rodents indicated as high/low (see Table 1).

Download figure to PowerPoint

EXPERIMENT 1

A simple comparisons of mean survival rates (not adjusted for effects of nest site and date) between active and experimental nests showed differences of both directions and generally less than c. 15%. A marked exception was yellowhammer in 1998, where active nests have survived by about 30% better than experimental nests (Fig. 4). Significance tests on these raw data were precluded because of the time lag between active and experimental nests and seasonal trends in nest survival rate. Nest survival measured by DSR and FATE responses was similar within all species-year samples in both active and experimental nests (Fig. 4).

image

Figure 4. Observed (unadjusted) survival rates (means with 95% CI) of active (hatched bars) and experimental (open bars) nest. Comparison based on: DSR – daily survival rates extrapolated to 15-day period; FATE – binary proportion of nests that have survived over 15 days since discovery (active nests) or initial exposure date (experimental nests). The nests used for FATE comparison represent a subsample of those used for DSR comparison (see Methods for sample definition). Only paired nests are included – number of nests shown above bars applies to both active and experimental nests.

Download figure to PowerPoint

The most complex multispecies model (including effects of nest activity, concealment, year, species, date and study plot) was examined for presence of interactions and for sensitivity to the type of response variable and the extent of data included. Using DSR response and all nests sample (n = 1618) revealed significant three–way interactions: activity × species × year (χ24= 10·1, P = 0·038) and activity × concealment × year (χ22= 7·8, P = 0·021). Using DSR response and paired nests sample (n = 822) revealed similar patterns, but only two-way interactions among the above effects were significant: activity × species (χ22= 8·1, P = 0·018), species × year (χ24= 10·9, P = 0·027) and concealment × year (χ22= 15·7, P = 0·001). Using FATE response led to essentially the same conclusions as using DSR, hence the results are not reported. Because of the significant interactions with species effect, I next performed separate species analyses.

The model for thrushes (Table 2; Figs 5–7) showed that DSR differed among years and that all other effects were consistent across years. The only significant interaction was detected between effects of activity and date: DSR increased with date, which effect was stronger in active (2·37 ± 0·49; regression slope ± SE, × 0·01; logit scale) than in experimental nests (0·45 ± 0·27). Nevertheless, for any particular date of the breeding season, DSR was on average higher for active than for experimental nests. The positive effect of concealment on DSR, though strong when considered separately, was only marginal and not significant after controlling for date.

Table 2.  Experiment 1 – thrushes. Generalized linear models (binomial error, logit link) relating the daily survival rate to nest activity (active vs. experimental: repeated factor within individual nests), concealment (poor vs. good), year (1998–2000), date and height above ground (continuous covariates), and effect of study plot (blocking variable). Given for each effect or interaction of effects is P-value associated with the generalized score statistic based on Type III test. A model including only main effects (M) and a model with interactions (I) is shown. The latter was obtained by sequential deleting nonsignificant (P > 0·05) interaction terms from an original model containing all two- and three-way interactions. Significance of terms included in the models derived from the all nests sample was further evaluated by fitting these models to a restricted sample of paired nests (see Methods for sample definition)
Effectd.f.All nests (n = 532)Paired nests (n = 208)
MIMI
  1. × Main effects involved in the interaction terms retained in the model.

  2. P < 0·05 shown in bold.

Activity (within nest) 10·002×0·009×
Concealment 10·2310·2260·7550·747
Year 20·0040·0010·0580·039
Date 10·001×0·002×
Height 10·9630·9000·1020·097
Study plot210·0540·0400·5430·481
Activity × Date 1 0·001 0·071

The model for blackcap revealed significant three-way interactions: activity × height × year (χ22= 9·4, P = 0·009) and activity × concealment × year (χ22= 7·5, P = 0·024). Next separate year analyses were performed (Table 3; Figs 5–7). DSR tended to decrease with date in 1998 (–0·78 ± 0·45) but increased in 1999 (1·85 ± 0·44) and 2000 (0·78 ± 0·31). In two of three years a marginally significant interaction was found between effects of nest activity and height. The only significant effect of height revealed by the separate slopes models was that for active nests in 1998 (106·3 ± 43·3). Accordingly, the main effect models did not reveal an overall effect of nest height in any year. A significant interaction between effects of nest activity and concealment was found in two of three years (Table 3; Fig. 7). DSR increased with concealment in active nests (1998: χ21= 14·8; 1999: χ21= 22·6, both P < 0·001) but not in experimental nests (P > 0·1 in both years). As a result, active nests showed significantly higher survival than experimental nests within the good-concealment category (1998: χ21= 12·8, P < 0·001; 1999: χ21= 5·0, P = 0·026), while nonsignificant (P > 0·3 in both years) effects of opposite direction were found in the poor-concealment category (Figs 6 and 7). Only marginally significant decrease of DSR with concealment, but neither significant interactions, nor overall effect of nest activity was found in one of three years (2000).

Table 3.  Experiment 1 – blackcap. Generalized linear models (binomial error, logit link) relating the daily survival rate to nest activity (active vs. experimental: repeated factor within individual nests), concealment (poor vs. good), date and height above ground (continuous covariates), and effect of study plot (blocking variable). The separate year models were obtained by partitioning the total blackcap model, that revealed significant (P < 0·05) activity × concealment × year and activity × height × year interaction. For other details see Table 2
Effectd.f.All nests (n = 885)Paired nests (n = 512)
1998 (n = 236)1999 (n = 289)2000 (n = 360)1998 (n = 112)1999 (n = 168)2000 (n = 232)
MIMIM = IMIMIM = I
  1. × Main effects involved in the interaction terms retained in the model.

  2. P < 0·05 shown in bold.

Activity (within nest) 10·125×0·721×0·6100·432×0·692×0·807
Concealment 10·078×0·001×0·0500·015×0·001×0·098
Date 10·2610·1030·0010·0010·0150·3770·3190·0040·0030·242
Height 10·131×0·959×0·9440·480×0·435×0·858
Study plot210·3200·2920·1430·1660·0030·1990·3310·1840·2050·029
Activity × Concealment 1 0·001 0·013  0·127 0·119 
Activity × Height 1 0·028 0·048  0·671 0·079 
image

Figure 7. Nest survival rates (‘least square’ means with 95% CI) estimated for three-way interaction year × concealment × activity (active = hatched, experimental = open). Based on the separate-species models containing all two- and three-way interactions, fitted to the total nest sample.

Download figure to PowerPoint

The model for yellowhammer (Table 4, Figs 5–7) showed that DSR was higher for active than for experimental nests and increased with nest concealment, which effects were consistent across years. The only significant interaction was detected between effects of date and year: DSR did not change with date in 1998 (–0·48 ± 0·66) and 1999 (1·71 ± 1·07) but increased in 2000 (2·12 ± 0·82). The main effect model suggests marked annual variation in mean DSR.

Table 4.  Experiment 1 – yellowhammer. Generalized linear models (binomial error, logit link) relating the daily survival rate to nest activity (active vs. experimental: repeated factor within individual nests), concealment (poor vs. good), year (1998–2000), date and height above ground (continuous covariates). For other details see Table 2
Effectd.f.All nests (n = 201)Paired nests (n = 102)
MIMI
  1. × Main effects involved in the interaction terms retained in the model.

  2. P < 0·05 shown in bold.

Activity (within nest)10·0010·0010·0030·003
Concealment10·0590·0220·0230·014
Year20·001×0·001×
Date10·285×0·609×
Height10·2630·2130·2970·286
Year × Date1 0·018 0·272

EXPERIMENT 2

The fates (success/predation) over 15-day exposure period of paired active and experimental blackcap nests tended to be positively associated, but this effect was not significant (exact test of independence in a 2 × 2 table) in either year (1999: n = 46, P = 0·376; 2000: n = 70, P = 0·357). In contrast to Experiment 1, a simple paired comparison of survival rates between active and experimental nests (exact McNemar’s test for paired binary responses) was relevant here because of simultaneous exposure of paired nests. No significant effect was found in either year (1999: P > 0·9; 2000: P = 0·110). A generalized linear model (Table 5; Fig. 5) showed consistent results for the two response variables and verified the findings of simple paired comparisons. All main effects were consistent across years and generally matched those found for blackcap in Experiment 1. DSR decreased from 1999 to 2000 but did not differ significantly between active and experimental nests. DSR increased with date in poor-concealment nests (4·75 ± 1·51) but decreased in good-concealment nests (–3·29 ± 1·26).

Table 5.  Experiment 2 – blackcap. Generalized linear models (binomial error, logit link) relating the daily survival rate (DSR) or nest survival over 15-day period (FATE) to nest activity (active vs. experimental: repeated factor within nest-pairs), concealment (poor vs. good), year (1999 vs. 2000), date and height above ground (continuous covariates). For other details see Table 2
EffectDSR (n = 120)FATE (n = 116)
MIMI
  1. × Μain effects involved in interaction terms retained in the model.

  2. P < 0·05 shown in bold.

  3. d.f. = 1 for all effects.

Activity (within nest-pair)0·0680·0880·2290·275
Concealment0·994×0·132×
Year0·0040·0080·0020·004
Date0·807×0·663×
Height0·1390·1900·2830·345
Concealment × Date 0·001 0·001
image

Figure 5. Nest survival rates (‘least square’ means with 95% CI) estimated for the main effects of year, concealment and activity (A = active, E = experimental). Based on the separate-species models containing all two- and three-way interactions, fitted to the total nest sample (Experiment 1) or paired nest sample (Experiment 2). The annual number of nests is shown for each species.

Download figure to PowerPoint

Discussion

  1. Top of page
  2. Summary
  3. Introduction
  4. Methods
  5. Results
  6. Discussion
  7. Conclusions
  8. Acknowledgements
  9. References

METHODOLOGICAL CONSIDERATIONS

Artificial nest experiments suffer from many drawbacks (Major & Kendal 1996). While it was hardly possible to check for all of them simultaneously, an attempt was made to eliminate different biases in turn in the separate experiments. The experimental nests were designed to resemble as closely as possible the real models in all aspects but the target variable – presence of parents. Experiment 2 came close to this goal, but at the expense of limited sample size and coverage of only one species. In spite of this effort, the natural and experimental nests inevitably differed in many uncontrolled or only statistically controlled traits. Interpretation of the experimental results thus relies on the assumption that the difference between survival of natural and experimental nests is mostly, but not necessarily completely, attributable to parental effects. Review of past work on artificial nest methodology made this assumption plausible. It has long been realized that survival of artificial nests does not measure an absolute survival of natural nests, but as relative measure can be compared among experimental treatments (Major & Kendal 1996). The interactions between effect of nest type (i.e. presence of parents) and other factors (nest site, date, year, species) cast some doubt on general validity of this assumption, supporting instead the recent critical view (Butler & Rotella 1998; Wilson et al. 1998; Buler & Hamilton 2000; Davison & Bollinger 2000; Weidinger 2001b).

The two types of artificial eggs used in Experiment 1 were associated with different biases. The plastic eggs were of natural size and colour, whereas the quail (= natural) eggs were of unnatural size (all species) and colour (thrushes). Only the latter provided reward (food) to the predator but their shell was probably difficult to break for mice (Marini & Melo 1998; Maier & DeGraaf 2000; Rangen, Clark & Hobson 2000). The effect on nest survival of fluctuating rodent abundance would probably be easier to detect by using natural eggs of appropriate size, as suggested by the larger year effect revealed in Experiment 2 (sparrow eggs). Coloration of artificial eggs may not represent serious limitation of this study, as it was shown to have little effect on the risk of predation in these species (Weidinger 2001c). The effect of year was partially confounded with that of egg type in Experiment 1, but the differences among years can not be simply explained by the different type of eggs used in the last year of the study. Marked year-to-year variation occurred throughout the study and was not parallel across all species (Figs 5–8). Moreover, the annual variation in nest survival was not the focus of this study. Instead, annual variation (of whatever cause) was statistically controlled for when evaluating the effects of primary interest (behaviour, concealment).

Perhaps the main impression from this study is the high variability in results. None of the patterns observed was consistent across all species and/or years, but all results were consistent for the two response variables. Contrary to main effects, conclusions about significance of interactions were sensitive to the extent of data included. The paired nests sample (about half size) was not sufficient to detect significant interactions revealed by the analysis of the all nests sample (Tables 2–4), although all effects detected were of the same direction and of comparable size. The number of nests observed and their spatial distribution seem sufficient to exclude spurious effects typical of small data sets, but not all species–year samples were sufficient to detect effects of clear biological significance (e.g. 10% difference in nest survival over the incubation period) with an acceptable power (Fig. 8). Also representativeness of the analysed nest samples differed among species. The well-concealed ground (vs. less concealed off-ground) nests in yellowhammer and canopy (vs. shrub) nests in thrushes were under-represented (Weidinger 2001a), whereas blackcap nests, being located in a searchable vegetation, represented the least biased sample.

image

Figure 8. Differences (with 95% CI) in daily survival rates between active and experimental nests estimated by the separate-species models (see Figs 5 and 6). Positive values indicate better survival of active nests. The two ‘pooled’ values shown for yellowhammer refer to combined years 1999 and 2000 (point) or all three years (bar), respectively.

Download figure to PowerPoint

NEST PREDATORS

A review of past work (Söderstrom et al. 1998) showed that birds prevailed over nonavian predators for shrub compared to ground nests, and a similar trend has also been predicted from foraging theory (Schmidt 1999). Mice are known to be important predators of passerine nests (Ketterson et al. 1996; Bureš 1997; Honza et al. 1998; Maier & DeGraaf 2000; Pietz & Granfors 2000; Schmidt et al. 2001a). This study supported these findings; the overall ratio of mammalian to avian predation decreased from yellowhammer (small eggs, ground/near ground nests) through blackcap (small, shrub) to thrushes (medium sized, shrub/subcanopy). Several pieces of evidence suggest a link between annual changes of nest predation and the predators involved. The ratio of mammalian to avian predation was higher in years when rodent abundance peaked; an effect that was clear in yellowhammer, detectable in blackcap, but not found in thrushes. Finally, for each species, the mean annual nest survival was inversely related to abundance of the major nest predator and its share of total predation.

The dominant nest predators differed not only among species or broadly defined nest sites (ground/shrub), but also among nest sites within species (see also Rangen et al. 1999; Dion et al. 2000; Hansson, Bensch & Hasselquist 2000). Visually hunting corvids took mainly the poorly concealed nests, whereas rodents preyed chiefly upon the nests well hidden in vegetation. This could be because the most conspicuous nests were quickly lost to diurnal corvids before nocturnal predation by rodents could take place, while the opposite may happen to well-concealed nests. Alternatively, rodents may avoid foraging in open places to reduce risk of avian predation on themselves (Korpimäki, Koivunen & Hakkarainen 1996; Manson & Stiles 1998; but see Schmidt et al. 2001b), whereas the most concealed nests may not be detectable for corvids. Such a clear pattern was consistently found in blackcap, whose nests showed a high variation in overall concealment and were heavily preyed upon by both rodents and corvids. A similar (though not significant owing to low number of nests) effect was found also in yellowhammer, but only in the year of relatively low rodent abundance, when corvids became important nest predators. In this case the increased predation by corvids did not compensate for reduced predation by rodents and disproportionately affected the poorly concealed nests. No such effects were found in thrushes with their conspicuous nests, where concealment had little effect on nest survival and where apparently all nests were detectable for corvids.

NEST CONCEALMENT

This study revealed generally positive relationship between concealment and survival of active nests (see also Martin 1992b), which effect disappeared in thrushes after controlling for date. Previous studies reported similar relationships in blackcap (positive: Hoi-Leitner, Nechtelberger & Hoi 1995), song thrush (neutral: Götmark et al. 1995) and blackbird (neutral: Cresswell 1997; positive: Hatchwell, Chamberlain & Perrins 1996). The relationships between nest concealment and survival were parallel for active and experimental nests in thrushes and yellowhammer. In blackcap, however, survival of experimental nests was independent of concealment, indicating that the positive relationship found in active nests was largely due to parental behaviour. Hence, this study concurs with others that the effect of concealment may differ between active and artificial nests, but the patterns reported are conflicting: either the effect was found in active nests but not in artificial ones (this study; Dion et al. 2000), or vice versa (Götmark et al. 1995; Cresswell 1997).

PARENTAL BEHAVIOUR

The net effect of parental behaviour on nest survival differed among species, being either positive (thrushes, yellowhammer) or neutral (blackcap). The finding that positive effects (nest defence) prevailed over the negative ones (nest disclosure) (Fig. 8) is further supported by the lack of increase in overall predation rates between the egg (low activity) and nestling (high activity) stages in all four species (K. Weidinger, unpublished data; see Martin et al. 2000b for an opposite result). Two other similarly designed experiments on comparable species also have shown higher survival for active than for artificial nests (Cresswell 1997; Wilson et al. 1998).

On the within-species level, the effect of behaviour was either independent of nest concealment (thrushes, yellowhammer) or interacted with it (2 of 3 years in blackcap). In the latter case, effect of behaviour was positive for well-concealed nests but negative for poorly concealed nests. The interaction was thus of opposite direction than that reported for thrushes (Götmark et al. 1995; Cresswell 1997), and not consistent with expectations derived from optimality theory of nest defence. According to the latter, parents with exposed nests should exhibit more vigorous nest defence than those with concealed nests (McLean et al. 1986; Hobson et al. 1988; Montgomerie & Weatherhead 1988). The above prediction was not supported for thrushes, but the test was inconclusive for blackcap, where the effect of parental behaviour on survival of differently concealed nests (but not necessarily the intensity of nest defence) was influenced by concealment × predator interaction.

Finally, an interaction between effects of behaviour and date was found in thrushes; the positive effect of parental behaviour on nest survival tended to increase as the breeding season progressed. This is consistent with hypothesis predicting seasonal increase in intensity of parental nest defence as re-nesting potential declines (Montgomerie & Weatherhead 1988). All species under study have similar re-nesting capacity, but presumably differ in nest defence potential. Accordingly, the above seasonal effect was detectable only in thrushes. This effect cannot be accounted for by changes in nest defence between nesting stages (Montgomerie & Weatherhead 1988; McLean & Rhodes 1991) because the analysis was restricted to the incubation stage, neither by seasonal changes in nest concealment (Hobson et al. 1988), which effect was controlled statistically.

ANTIPREDATOR STRATEGIES

Thrushes are capable of active nest defence against birds, including direct attacks (own observations; Cresswell 1997; Grim & Honza 2001), and their nests are unlikely to be threatened by mice when attended by parents. On the other hand, the large nests are conspicuous on their own, so the risk of nest disclosure is unlikely to be greatly increased by parental activity. Resulting net effect of parental behaviour on nest survival is thus positive (Table 6) and independent of nest concealment, which itself has little effect (Fig. 1Ba). Hence, nest protection through maximizing nest concealment may not be an efficient strategy in these species, instead, nest placement that facilitates vigilance and active defence (e.g. intermediate concealment, Götmark et al. 1995) may be selected for.

Table 6.  Summary of hypothesized (H) and observed effects in this study. Effect of parental activity on nest survival indicated as positive (nest defence), negative (nest disclosure) or neutral (0)
 ThrushesBlackcapYellowhammer
Nest predatorscorvidsrodentscorvidsrodentscorvidsrodents
Predation on poor-concealment nests (Fig. 3)highmediumhighmediumlow (medium)high
Predation on good-concealment nests (Fig. 3)highmediummediumhighlowhigh
Parental nest defence potential (H)yesyesnoyesnoyes
Parental nest disclosure potential (H)nonoyesnoyesno
Effect of parental behaviour on nest survival:
 by type of predator/nest site (Figs 6 and 7)+++0 (–)+
 overall (Figs 5 and 8)+0+ (0)

Blackcap has limited capacity to deter large predators such as corvids. Nonetheless, passive defence against rodents is possible through nest attentiveness, because mice are supposed to take eggs or nestlings opportunistically from unattended nests. Incubating blackcaps are typically rather tame; their nest attentiveness (proportion of all spot observations with parent present) increases from about 11% on the first egg day to 84% on the last egg day and the rest of incubation (K. Weidinger, unpublished data). Compared to thrushes, blackcap nests and eggs are relatively inconspicuous, so that an overall risk of nest disclosure to visually searching predators may be, at least potentially, increased by parental activity. The nest defence/disclosure effect in blackcap thus depends on the type of principal predator, which in turn differs among nest sites, yielding negative and positive net effects of parental behaviour in poorly and well-concealed nests, respectively (Fig. 1Ad). These opposing effects offset each other across all nest sites, so that neutral net effect of behaviour is observed on the population level (Table 6).

Similar main effects of behaviour and concealment on survival of blackcap nests were found in two independent experiments, while their interaction was detected only in Experiment 1 (larger sample). Moreover, one of three annual replicates of Experiment 1 supported predictions of the null model (Fig. 1Aa). Further research is needed to evaluate these alternatives over a longer time scale and to test whether the positive relationship between concealment and survival of active nests results mainly from behaviour × concealment interaction (Fig. 1Ad; Experiment 1) or represents an effect of nest site per se (Fig. 1Ac; Experiment 2). This study suggests, that although blackcaps selecting nest site face a trade-off between avoidance of corvid and mice predation, maximizing nest concealment should represent an efficient antipredator strategy for this species with limited capacity for active nest defence against large visually oriented predators. Other experiments (S. Bureš & V. Pavel, unpublished data) showed that blackcaps behaved inconspicuously in the presence of predator models, apparently so that the nest site is not disclosed.

The nest defence/disclosure potential in yellowhammer is similar to that in blackcap, but nests are preyed upon mainly by rodents. In contrast to corvids, nest predation behaviour of mice probably results from random encounters with unattended nests (incidental predation; Schmidt et al. 2001b) rather than from active responses to observed parental activity. The risk of nest disclosure is then low, while nest attentiveness by parents probably represents an important mechanism of passive nest defence. The resulting net effect of parental behaviour on nests survival is thus positive (Table 6) and independent of nest concealment, which itself also has a marginally positive effect (Fig. 1Bc). Schmidt et al. (2001a) estimated that about 60–75% of veery (Catharus fuscescens) nests encountered by rodents were not predated because of parental nest defence. By applying their method (1 – (daily predation rate of active nests/daily predation rate of experimental nests)) to yellowhammer I estimated a similar value of 72%. Relative importance of parental behaviour and concealment is hypothesized to vary among years in relation to fluctuating abundance of rodent and their effect on total nest predation.

Conclusions

  1. Top of page
  2. Summary
  3. Introduction
  4. Methods
  5. Results
  6. Discussion
  7. Conclusions
  8. Acknowledgements
  9. References

The null model (Fig. 1Aa) was rejected – apart from multitude of other suspected factors, nest survival was influenced by parental behaviour and/or nest concealment in all species and in most years. The basic prediction was supported; the principal nest predators and the relative importance of behaviour and nest site effects differed among species. The observed patterns included: positive effect of parental behaviour combined with neutral effect of nest concealment (Fig. 1Ba; thrushes); independent positive effects of behaviour and concealment (Fig. 1Bc; yellowhammer); neutral effect of behaviour combined with positive effect of concealment (Fig. 1Ad/Ac; blackcap). These species-specific patterns are consistent with the hypothesis (Ricklefs 1977; see also Murphy et al. 1997; Willson et al. 2001) that relatively larger species with more conspicuous nests should engage in more vigorous nest defence compared to smaller species with less conspicuous nests. The question of whether the observed species-specific effects of behaviour result more from differences among species in the intensity of nest defence or from differences in its efficiency remains open to further study. The interspecific pattern was further complicated by the within-species interactions between nest site and type of predator, which effects may prove to be quite common (Rangen et al. 1999; Dion et al. 2000). Moreover, predation risk associated with particular nest sites may be influenced by interactions among several different predators, both within and between trophic levels (Schmidt et al. 2001b).

Experimental control of parental behaviour enabled the mechanisms behind the observed effects of nest concealment to be distinguished. The positive relationship between concealment and nest survival resulted either from effect of nest site per se (yellowhammer) or from the effect of parental behaviour (blackcap), which changed from negative (nest disclosure) to positive (nest defence) between poorly and well-concealed nests, respectively. The alternative view, that the lack of within-species relationship between nest concealment and survival (thrushes) results from parental behaviour compensating for the predation risk associated with poor nest concealment (Cresswell 1997), was not supported.

The evolutionary patterns in life-history traits associated with the risk of nest predation depend on the proximate mechanisms of predator–prey interactions (Martin et al. 2000b). The accumulating evidence in support of multiple and interactive effects on nest predation is reflected in a growing feeling that further studies should integrate, among others, effects of nest site, parental behaviour and foraging behaviour of predators in a single conceptual framework (Murphy et al. 1997; Schmidt 1999; Martin et al. 2000b; Willson & Gende 2000). Potential explanatory power of mechanistic approach has been demonstrated by Schmidt et al. (2001a,b). Highly variable and even conflicting results of this study demonstrated the importance of scale in nest predation studies. When considered separately, the apparently clear-cut results of partial analyses (single years/species), would lead to different conclusions, which might be the case with small-scale experiments.

Acknowledgements

  1. Top of page
  2. Summary
  3. Introduction
  4. Methods
  5. Results
  6. Discussion
  7. Conclusions
  8. Acknowledgements
  9. References

This work was supported by the Grant Agency of the Czech Republic (GAČR 206/98/P119 and 206/01/P028) and by the Ministry of Education of the Czech Republic (MSM 153100012). I would like to thank V. Remeš, V. Pavel, M. Krist, T. Grim and S. Bureš for insightful comments on the work and F. Krause for kindly supplying tree sparrow eggs. The paper benefited from criticism by two referees.

References

  1. Top of page
  2. Summary
  3. Introduction
  4. Methods
  5. Results
  6. Discussion
  7. Conclusions
  8. Acknowledgements
  9. References
  • Aebischer, N.J. (1999) Multi-way comparisons and generalized linear models of nest success: extensions of the Mayfield method. Bird Study, 71, 2231.
  • Blancher, P.J. & Robertson, R.J. (1982) Kingbird aggression: does it deter predation? Animal Behaviour, 30, 929930.
  • Buler, J.J. & Hamilton, R.B. (2000) Predation of natural and artificial nests in a southern pine forest. Auk, 117, 739747.
  • Bureš, S. (1997) High common vole Microtus arvalis predation on ground-nesting bird eggs and nestlings. Ibis, 139, 173174.
  • Burhans, D.E. (2000) Avoiding the nest: responses of field sparrows to the threat of nest predation. Auk, 117, 803806.
  • Burhans, D.E. & Thompson, F.R. (1998) Effects of time and nest-site characteristics on concealment of songbird nests. Condor, 100, 663672.
  • Burhans, D.E. & Thompson, F.R. (2001) Relationship of songbird nest concealment to nest fate and flushing behavior of adults. Auk, 118, 237242.
  • Butler, M.A. & Rotella, J.J. (1998) Validity of using artificial nests to assess duck-nest success. Journal of Wildlife Management, 62, 163171.
  • Clark, R.G. & Shutler, D. (1999) Avian habitat selection: Pattern from process in nest-site use by ducks? Ecology, 80, 272287.
  • Crawley, M.J. (1993) GLIM for Ecologists. Blackwell, London.
  • Cresswell, W. (1997) Nest predation: the relative effects of nest characteristics, clutch size and parental behaviour. Animal Behaviour, 53, 93103.
  • Davison, W.B. & Bollinger, E. (2000) Predation rates on real and artificial nests of grassland birds. Auk, 117, 147153.
  • Dion, N., Hobson, K.A. & Larivière, S. (2000) Interactive effects of vegetation and predators on the success of natural and simulated nests of grassland songbirds. Condor, 102, 629634.
  • Götmark, F., Blomqvist, D., Johansson, O.C. & Bergkvist, J. (1995) Nest site selection: a trade-off between concealment and view of the surroundings? Journal of Avian Biology, 26, 305312.
  • Grim, T. & Honza, M. (2001) Differences in behaviour of closely related thrushes (Turdus philomelos and T. merula) to experimental parasitism by the common cuckoo Cuculus canorus. Biologia, 56, 549556.
  • Hansson, B., Bensch, S. & Hasselquist, D. (2000) Patterns of nest predation contribute to polygyny in the Great Reed Warbler. Ecology, 81, 319328.
  • Hatchwell, B.J., Chamberlain, D.E. & Perrins, C.M. (1996) The reproductive success of blackbirds Turdus merula in relation to habitat structure and choice of nest site. Ibis, 138, 256262.
  • Hensler, G.L. (1985) Estimation and comparison of functions of daily nest survival probabilities using the Mayfield method. Statistics in Ornithology (eds B. J. T.Morgan & P. M.North), pp. 289301. Springer-Verlag, New York.
  • Hobson, K.A., Bouchart, M.L. & Sealy, S.G. (1988) Responses of naive yellow warblers to a novel nest predator. Animal Behaviour, 36, 18231830.
  • Hogstad, O. (1995) Do avian and mammalian nest predators select for different nest dispersion patterns of fieldfares Turdus pilaris? A 15-year study. Ibis, 137, 484489.
  • Hoi-Leitner, M., Nechtelberger, H. & Hoi, H. (1995) Song rate as a signal for nest site quality in blackcaps (Sylvia atricapilla). Behavioral Ecology and Sociobiology, 37, 399405.
  • Honza, M., Oien, I.J., Moksnes, A. & Røskaft, E. (1998) Survival of reed warbler Acrocephalus scirpaceus clutches in relation to nest position. Bird Study, 45, 104108.
  • Howlett, J.S. & Stutchbury, B.J. (1996) Nest concealment and predation in hooded warblers: experimental removal of nest cover. Auk, 113, 19.
  • Ketterson, E.D., Nolan, V., Cawthorn, M.J., Parker, P.G. & Ziegenfus, C. (1996) Phenotypic engineering: using hormones to explore the mechanistic and functional bases of phenotypic variation in nature. Ibis, 138, 7086.
  • King, D.I., Degraaf, R.M., Griffin, C.R. & Maier, T.J. (1999) Do predation rates on artificial nests accurately reflect predation rates on natural bird nests? Journal of Field Ornithology, 70, 257262.
  • Komdeur, J. & Kats, R.H. (1999) Predation risk affects trade-off between nest guarding and foraging in Seychelles warblers. Behavioral Ecology, 10, 648658.
  • Korpimäki, E., Koivunen, V. & Hakkarainen, H. (1996) Microhabitat use and behavior of voles under weasel and raptor predation risk: predator facilitation? Behavioral Ecology, 7, 3034.
  • Maier, T.J. & DeGraaf, R.M. (2000) Predation on Japanese quail vs. house sparrow eggs in artificial nests: small eggs reveal small predators. Condor, 102, 325332.
  • Major, R.E. & Kendal, C.E. (1996) The contribution of artificial nest experiments to understanding avian reproductive success: a review of methods and conclusions. Ibis, 138, 298307.
  • Manolis, J.C., Andersen, D.E. & Cuthbert, F.J. (2000) Uncertain nest fates in songbird studies and variation in Mayfield estimation. Auk, 117, 615626.
  • Manson, R.H. & Stiles, E.W. (1998) Links between microhabitat preferences and seed predation by small mammals in old fields. Oikos, 82, 3750.
  • Marini, M.Â. & Melo, C. (1998) Predators of quail eggs, and the evidence of the remains: implications for nest predation studies. Condor, 100, 395399.
  • Martin, T.E. (1992a) Interaction of nest predation and food limitation in reproductive strategies. Current Ornithology, Vol. 9 (ed. D. M.Power), pp. 163197. Plenum, New York.
  • Martin, T.E. (1992b) Breeding productivity considerations: What are the appropriate habitat features for management? Ecology and Conservation of Neotropical Migrants (eds J. M.Hagan & D. W.Johnston), pp. 455473. Smithsonian Institution, Washington, DC.
  • Martin, T.E. (1998) Are microhabitat preferences of coexisting species under selection and adaptive? Ecology, 79, 656670.
  • Martin, T.E., Martin, P.R., Olson, C.R., Heidinger, B.J. & Fontaine, J.J. (2000a) Parental care and clutch sizes in North and South American birds. Science, 287, 14821485.
  • Martin, T.E., Scott, J. & Menge, C. (2000b) Nest predation increases with parental activity: separating nest site and parental activity effects. Proceedings of the Royal Society of London Series B – Biological Sciences, 267, 22872293.
  • McLean, I.G. & Rhodes, G. (1991) Enemy recognition and response in birds. Current Ornithology, Vol. 8 (ed. D. M.Power), pp. 173211. Plenum, New York.
  • McLean, I.G., Smith, J.M. & Stewart, K.G. (1986) Mobbing behavior, nest exposure, and breeding success in the American robin. Behaviour, 96, 171185.
  • Misenhelter, M.D. & Rotenberry, J.T. (2000) Choices and consequences of habitat occupancy and nest site selection in sage sparrows. Ecology, 81, 28922901.
  • Montgomerie, R.D. & Weatherhead, P.J. (1988) Risks and rewards of nest defense by parent birds. Quarterly Review of Biology, 63, 167187.
  • Murphy, M.T., Cummings, C.L. & Palmer, M.S. (1997) Comparative analysis of habitat selection, nest site and nest success by cedar waxwings (Bombycilla cedrorum) and eastern kingbirds (Tyrannus tyrannus). American Midland Naturalist, 138, 344356.
  • Pietz, P.J. & Granfors, D.A. (2000) Identifying predators and fates of grassland passerine nests using miniature video cameras. Journal of Wildlife Management, 64, 7187.
  • Rangen, S.A., Clark, R.G. & Hobson, K.A. (1999) Influence of nest-site vegetation and predator community on the success of artificial songbird nests. Canadian Journal of Zoology, 77, 16761681.
  • Rangen, S.A., Clark, R.G. & Hobson, K.A. (2000) Visual and olfactory attributes of artificial nests. Auk, 117, 136146.
  • Ricklefs, R.E. (1977) Reactions of some Panamanian birds to human intrusion at the nest. Condor, 79, 376379.
  • SAS Institute Inc. (2000) SAS OnlineDoc., Version 8. SAS Institute Inc., Cary. NC.
  • Schmidt, K.A. (1999) Foraging theory as a conceptual framework for studying nest predation. Oikos, 85, 151160.
  • Schmidt, K.A., Goheen, J.R. & Naumann, R. (2001a) Incidental nest predation in songbirds: behavioral indicators detect ecological scales and processes. Ecology, 82, 29372947.
  • Schmidt, K.A., Goheen, J.R., Naumann, R., Ostfeld, R.S., Schauber, E.M. & Berkowitz, A. (2001b) Experimental removal of strong and weak predators: mice and chipmunks preying on songbird nests. Ecology, 82, 29272936.
  • Söderstrom, B., Pärt, T. & Rydén, J. (1998) Different nest predator faunas and nest predation risk on ground and shrub nests at forest ecotones: an experiment and a review. Oecologia, 117, 108118.
  • Steiger, J.H. (1999) Statistica Power Analysis. StatSoft Inc., Tulsa, OK.
  • Veen, T., Richardson, D.S., Blaakmeer, K. & Komdeur, J. (2000) Experimental evidence for innate predator recognition in the Seychelles warbler. Proceedings of the Royal Society of London Series B – Biological Sciences, 267, 22532258.
  • Vickery, P.D., Hunter, M.L. & Wells, J.V. (1992) Evidence of incidental nest predation and its effects on nests of threatened grassland birds. Oikos, 63, 281288.
  • Weidinger, K. (2001a) Laying dates and clutch size of open-nesting passerines in the Czech Republic: a comparison of systematically and incidentally collected data. Bird Study, 48, 3847.
  • Weidinger, K. (2001b) How well do predation rates on artificial nests estimate predation on natural passerine nests? Ibis, 143, 632641.
  • Weidinger, K. (2001c) Does egg colour affect predation rate on open passerine nests? Behavioral Ecology and Sociobiology, 49, 456464.
  • Willson, M.F. & Gende, S.M. (2000) Nesting success of forest birds in southeast Alaska and adjacent Canada. Condor, 102, 314324.
  • Willson, M.F., Morrison, J.L., Sieving, K.E., De Santo, T.L., Santisteban, L. & Diaz, I. (2001) Patterns of predation risk and survival of bird nests in a Chilean agricultural landscape. Conservation Biology, 15, 447456.
  • Wilson, G.R., Brittingham, M.C. & Goodrich, L.J. (1998) How well do artificial nests estimate success of real nests? Condor, 100, 357364.