Free‐moving artificial eggs containing temperature loggers reveal remarkable within‐clutch variance in incubation temperature

Incubation is a crucial aspect of avian parental care and measuring incubation temperature in the wild can improve our understanding of life history tradeoffs and inform conservation efforts. However, there are challenges associated with measuring the temperature of eggs in natural nests. Most studies to date have measured incubation temperature by using a single, stationary logger in each nest. However, real eggs are rotated and moved throughout the nest by the parent during the incubation period, and thus, a stationary logger may not accurately represent the temperature experienced by individual eggs within the entire clutch. We recorded incubation temperature in nests by using multiple, mobile artificial egg temperature loggers. We installed six mobile loggers and one stationary logger in wood duck Aix sponsa nests to compare the two logger types in the field. We found that at a given ambient temperature, mobile loggers recorded lower average and more variable temperatures than stationary loggers. Further, temperatures recorded by stationary loggers showed no relationship with clutch size, while mobile loggers captured temperatures that were lower and more variable as clutch size increased. Also, the multiple mobile loggers revealed that eggs within a nest experienced a substantial range of temperatures throughout the incubation period. We discuss potential limitations of this method, but believe that it is a promising way to collect biologically‐relevant incubation temperature data and provides an opportunity to advance our understanding of incubation temperature as a parental effect.     

Plasticity of brood patch development and its influence on incubation periods in the yellow-eyed penguin Megadyptes antipodes: an experimental approach

In many bird species, eggs laid late in the laying period hatch after a shorter incubation period than early-laid eggs. However, the mechanisms that explain these seasonal declines in incubation periods among clutches remain poorly understood. In this study we investigated the plasticity of brood patch development during incubation in yellow-eyed penguins Megadyptes antipodes and established whether differences exist in brood patch formation among early, mean and late-breeding penguins. We also examined whether brood patch development was influenced by sex and age of birds. We then placed an artificial egg in nests a few days prior to egg laying to investigate whether the presence of an egg influences brood patch development and whether an advanced brood patch development at the time of egg laying causes declines in incubation periods. Initial brood patch width on the day the first egg was laid was dependent on sex and age, while the development of brood patch width after first egg laying was slower in early-laying birds than in mean- and late-laying birds. Initial brood patch temperature as well as temperature throughout incubation was largely dependent upon sex, whereby males had higher brood patch surface temperatures than females. Placement of an artificial egg in nests stimulated successfully brood patch development in manipulated birds, so that by the time they laid their own first egg, their brood patches were wider and had higher temperatures than those of control birds. Moreover, incubation periods of first eggs from manipulated nests were significantly shorter (43.5 days) than were those from control nests (47.3 days). Thus, variation in brood patch development and related differences in incubation temperature during early incubation could contribute to seasonal declines in incubation periods.     

Incubation prior to clutch completion accelerates embryonic development and so hatch date for eggs laid earlier in a clutch in the great tit Parus major

The ability of parents to respond to changes in food supply within a season will have a large effect on fitness through the number and quality of chicks fledged. Great tits, Parus major, attempt to synchronise their production of chicks with a seasonal food peak, but when food supply fails, hatching asynchrony of chicks provides a mechanism by which some young can be fledged because more developed chicks outcompete their less developed siblings for the reduced parental food supply. We tested whether female great tits can potentially control the degree of hatching asynchrony by using incubation before clutch completion, so that early laid eggs develop faster and hatch sooner. The temperature of an artificial egg placed in 29 nests during the laying period was measured with data loggers, and nocturnal incubation of eggs similar to incubation post clutch completion was recorded in all nests. We then demonstrated that eggs removed from the nest for 72 hour periods prior to clutch completion hatched later than eggs remaining in the nest for the entirety of the laying period. Our results show that variable pre clutch completion incubation (which was mostly nocturnal) can lead to faster embryo development and earlier hatching, so potentially providing a mechanism for adaptive female control of degree of hatching asynchrony.     

Egg temperature and initial brood patch area determine hatching asynchrony in Magellanic penguin Spheniscus magellanicus

In birds, the adaptive significance of hatching asynchrony has been under debate for many years and the parental effects on hatching asynchrony have been largely assumed but not often tested. Some authors suggest that hatching asynchrony depends on the incubation onset and many factors have been shown to influence hatching asynchrony in different species. Our objective was to analyze the exact timing of the onset of incubation and if this affects hatching asynchrony; and, in addition, which other factors (brood patch development, incubation position, adult body condition, intra‐clutch egg dimorphism, laying date and year) affect hatching asynchrony in Magellanic penguins Spheniscus magellanicus. We first estimated the eggshell temperature at which embryo development starts, with a non‐destructive and novel method. We then recorded individual egg temperatures in 61 nests during incubation, and related them, and other breeding parameters, to hatching asynchrony. We also observed incubation positions in 307 nests. We found a significant positive relationship between hatching asynchrony and the temperature that the first‐laid egg experienced during egg laying and between hatching asynchrony and the initial brood patch area. We also found a negative relationship between hatching asynchrony and the difference in temperature between second and first‐laid eggs within a clutch, measured after the egg‐laying period was finished. We ruled out position of the eggs during incubation, adult body condition, egg volume, laying date, and study year as factors influencing hatching asynchrony. The egg temperature during laying and the difference in temperature between eggs of a clutch are determinants of hatching asynchrony in Magellanic penguins.     

The role of partial incubation and egg repositioning within the clutch in hatching asynchrony and subsequent effects on breeding success

The main mechanism to achieve hatching asynchrony (HA) for incubating birds is to start heating the eggs before clutch completion. This might be achieved through partial incubation and/or early incubation. Even in the absence of incubation behaviour during the laying phase, clutches still experience a certain degree of asynchrony. Recent studies have shown that eggs located in the centre of the nest receive more heat than peripheral ones during incubation. As eggs receiving more heat would develop faster, we hypothesized that HA should be shorter in nests where eggs were moved homogeneously along the centre–periphery space during incubation than in those nests where eggs repeatedly remained in the same locations, either centrally or peripherally. We explored the relative roles of egg repositioning and partial incubation in determining HA in wild birds by (1) removing eggs from 20 Great Tit Parus major nests on the day of laying and replacing them with fake eggs to avoid partial incubation, and returning them when full incubation began; (2) monitoring twice a day the position of each individually marked egg relative to the clutch centre during incubation, and estimating the coefficient of variation of the distances; and (3) determining HA in each nest. Preventing partial incubation reduced HA by 51% days in experimental nests. It also caused negative effects for the incubating females (lengthening the full incubation period) and positive effects for the brood (increasing fledging success). However, our hypothesis about the role of egg repositioning on HA was not supported: all the females moved the eggs with remarkable consistency, generally attaining a coefficient of variation of the distances around 33%, and it was not related to the HA experienced. We therefore conclude that partial incubation is an important factor regulating HA, and females compensate for the potential effects of differential heating by moving the eggs homogeneously within the clutch.     

Incubation temperature in the Australasian Gannet Moms senator

Gannets (Sulidae) and some other pelecaniforms incubate their eggs under the webs of their totipalmate feet. These species have a wide latitudinal distribution from tropics to subarctic, but little is known of the incubation temperatures attained. I measured egg temperatures of the Australasian Gannet Morns serrator at Cape Kidnappers, New Zealand, employing a data logger to obtain records every 15 min for day-long sample periods at undisturbed nests. Egg surface temperatures were relatively stable and little affected by ambient temperature. Mean surface temperature of natural eggs was 34.9^oC for samples taken during the first 4 days of incubation, but this then increased and stabilized at 36.5^oC. Internal temperature of pipped eggs was about 1^oC higher, attributable to embryonic heat production. Upper surface temperature of eggs kept in a fixed position was about 2^oC below deep adult body temperature (40.3^oC), suggesting heat is transferred directly from the body through the feet. Clutch size does not appear to be limited by an inability to warm two eggs. These results are in general agreement with measurements from other web incubators and are well within the range for species with conventional brood patch incubation.     

Avian Incubation Patterns Reflect Temporal Changes in Developing Clutches

Incubation conditions for eggs influence offspring quality and reproductive success. One way in which parents regulate brooding conditions is by balancing the thermal requirements of embryos with time spent away from the nest for self-maintenance. Age related changes in embryo thermal tolerance would thus be expected to shape parental incubation behavior. We use data from unmanipulated Black-capped Chickadee (Poecile atricapillus) nests to examine the temporal dynamics of incubation, testing the prediction that increased heat flux from eggs as embryos age influences female incubation behavior and/or physiology to minimize temperature fluctuations. We found that the rate of heat loss from eggs increased with embryo age. Females responded to increased egg cooling rates by altering incubation rhythms (more frequent, shorter on- and off- bouts), but not brood patch temperature. Consequently, as embryos aged, females were able to increase mean egg temperature and decrease variation in temperature. Our findings highlight the need to view full incubation as more than a static rhythm; rather, it is a temporally dynamic and finely adjustable parental behavior. Furthermore, from a methodological perspective, intra- and inter-specific comparisons of incubation rhythms and average egg temperatures should control for the stage of incubation.     

Male boobies expel eggs when paternity is in doubt

We analyzed the effect of increased risk of cuckoldry on maleparental investment in eggs in the colonial blue-footed booby(Sula nebouxii). Seventeen experimental males were removedfrom the nesting territory for 10-12 h on a single day, 1-5days before laying (females' supposed fertile period = SFP),and 17 control males were removed for the same amount of timeon a single day, 7-29 days before laying (before the SFP).These removals were intended to simulate extended absence fromthe nest on a foraging excursion. Female extrapair courtshipand copulation rates did not increase during the removal ofthe social mate, and there was no evidence that experimentaland control males differed quantitatively in incubation ordefense of the clutch. However, 43% of experimental males expelledthe first-laid egg from the nest, whereas no control male didso. Apparently, male boobies drastically reduce parental investmentin eggs with a presumed elevated probability of extrapair fertilizationby destroying them.     

Costs and response to conspecific brood parasitism by colonial red-breasted mergansers

Costs of conspecific brood parasitism (CBP) are expected to be influenced by a species’ life history traits. Precocial birds lay large clutches, and clutches that have been enlarged by CBP can affect host fitness through a longer incubation period, displaced eggs, and lower hatching success. We examined costs and response to CBP by hosts in a population of colonial red-breasted mergansers (Mergus serrator; n?=?400 nests over 8 years) within which 29% of parasitized clutches were enlarged considerably (≥?15 eggs). Length of the incubation period did not increase with clutch size. The mean number of eggs displaced from a parasitized nest during incubation (2.8) was 2×?greater than at an unparasitized nest (1.4). Hatching success declined by 2% for each additional egg in the nest. Thus, for a nest with?≥?15 eggs, one or more fewer host eggs hatch relative to an unparasitized nest with the same number of host eggs, assuming equal probability of success for all eggs. Hosts were 40% more likely to desert nests receiving 2 or 6 experimental eggs relative to unparasitized control nests, although it is unknown whether hens deserting a nest renested elsewhere. Our study indicates that costs of CBP to hosts during nesting may be limited to those red-breasted mergansers incubating the largest clutches (≥?15 eggs), and it raises questions about the adaptive significance of deserting a parasitized clutch.     

Temperature and life history: experimental heating leads female tree swallows to modulate egg temperature and incubation behaviour

1. Life-history decisions are strongly affected by environmental conditions. In birds, incubation is energetically expensive and affected significantly by ambient temperature. We reduced energetic constraints for female tree swallows (Tachycineta bicolor) by experimentally heating nests during incubation by an average of 6.9 degrees C to test for changes in incubation behaviour. 2. Females in heated boxes (hereafter 'heated females') increased time spent incubating and maintained higher on-bout and off-bout egg temperatures. This indicates that female energetic constraints, not maximizing developmental conditions of offspring, determine incubation investment. Furthermore, this result suggests that embryonic developmental conditions in unmanipulated nests are suboptimal. 3. We found individual variation in how females responded to experimental heating. Early-laying (i.e. higher phenotypic quality) females with heated nests increased egg temperatures and maintained incubation constancy, while later-laying (lower quality) heated females increased incubation constancy. Changes in egg temperature were due to changes in female behaviour and not due directly to increases in internal nest-box temperatures. 4. Behaviour during the incubation period affected hatching asynchrony. Decreased variation in egg temperature led to lower levels of hatching asynchrony, which was also generally lower in heated nests. 5. Our study finds strong support for the prediction that intermittent incubators set their incubation investment at levels dictated by energetic constraints. Furthermore, females incubating in heated boxes allocated conserved energy primarily to increased egg temperature and increased incubation attentiveness. These results indicate that studies investigating the role of energetics in driving reproductive investment in intermittent incubators should consider egg temperature and individual variation more explicitly.     

Egg coloration and selection for crypsis in open-nesting blackbirds

High variation in egg coloration among birds has traditionally been explained as adaptation for camouflage. We tested this hypothesis by conducting reciprocal clutch exchanges (n=301) among Brewer's blackbirds Euphagous cyanocephalus , red-winged blackbirds Agelaius phoeniceus , and yellow-headed blackbirds Xanthocephalus xanthocephalus . We predicted that clutches placed against their natural nest backgrounds would have higher survival rates than heterospecific clutches. Intraspecific clutch exchanges were used as a control. Clutch survival was monitored for a 9-d period at all nests, during which time incubation rhythm and nest defense were quantified. Intraspecific clutch exchanges did not influence incubation or nest defense. For two of the species, intraspecific clutch exchanges did not influence clutch survival; in red-winged blackbirds, however, intraspecifically exchanged clutches had somewhat depressed survival curves relative to control clutches (P=0.08). The effect of interspecific clutch exchanges differed by host species. In Brewer's nests, eggs of the yellow-headed blackbird had lower survival than Brewer's eggs (P=0.02), but survival of red-winged blackbird eggs did not differ from Brewer's eggs (P=0.50). In nests of red-winged blackbirds, all three clutch types had approximately equal survival. In yellow-headed blackbird nests, eggs of the red-winged blackbird had lower survival than yellow-headed blackbird eggs (P=0.06), and survival of Brewer's eggs did not differ from yellow-headed blackbird eggs (P=0.31). These findings support a role for egg coloration as camouflage in two of the three species studied.     

Nest construction by a ground-nesting bird represents a potential trade-off between egg crypticity and thermoregulation

Predation selects against conspicuous colors in bird eggs and nests, while thermoregulatory constraints select for nest-building behavior that regulates incubation temperatures. We present results that suggest a trade-off between nest crypticity and thermoregulation of eggs based on selection of nest materials by piping plovers (Charadrius melodus), a ground-nesting bird that constructs simple, pebble-lined nests highly vulnerable to predators and exposed to temperature extremes. Piping plovers selected pebbles that were whiter and appeared closer in color to eggs than randomly available pebbles, suggesting a crypsis function. However, nests that were more contrasting in color to surrounding substrates were at greater risk of predation, suggesting an alternate strategy driving selection of white rocks. Near-infrared reflectance of nest pebbles was higher than randomly available pebbles, indicating a direct physical mechanism for heat control through pebble selection. Artificial nests constructed of randomly available pebbles heated more quickly and conferred heat to model eggs, causing eggs to heat more rapidly than in nests constructed from piping plover nest pebbles. Thermal models and field data indicated that temperatures inside nests may remain up to 2–6°C cooler than surrounding substrates. Thermal models indicated that nests heat especially rapidly if not incubated, suggesting that nest construction behavior may serve to keep eggs cooler during the unattended laying period. Thus, pebble selection suggests a potential trade-off between maximizing heat reflectance to improve egg microclimate and minimizing conspicuous contrast of nests with the surrounding substrate to conceal eggs from predators. Nest construction behavior that employs light-colored, thermally reflective materials may represent an evolutionary response by birds and other egg-laying organisms to egg predation and heat stress. An erratum to this article can be found at     

Intraspecific nest parasitism in the grey starling (Sturnus cineraceus)

We studied intraspecific nest parasitism in the grey starling (Sturnus cineraceus) in 1992 and 1993. We used three criteria to detect nest parasitism: (i) the appearance of more than one egg per day while the host was laying; (ii) the appearance of extra eggs after the host completed its clutch; and (iii) the appearance of eggs which were of a different shape, size and color to other eggs in the clutch. There were 290 nests (157 nests in 1992; 133 nests in 1993) in which the clutch was completed early (clutches initiated before May 10). Twenty-nine (1992) and 32 (1993) nests contained at least one parasitic egg. Parasitic eggs hatched if they were laid during the laying period and early in the incubation period of their host, and a few of them fledged. Fledging success of parasitic eggs was not different from that of eggs in non-parasitized nests if parasitic eggs were laid during the host's laying period. However, fledging success of all parasitic eggs was fewer than that of eggs in non-parasitized nests. By comparison, fledging success of parasitized nests was not a great as that of non-parasitized nests.     

Conspecific brood parasitism and egg rejection in Great-tailed Grackles

In this study, we tested whether conspecific brood parasitism (CBP) has selected for egg rejection behavior in the colonial Great-tailed Grackle Quiscalus mexicanus . No evidence of CBP was recorded at 797 Great-tailed Grackle nests, and we did not induce CBP by experimentally removing nests while grackles were laying. We determined experimentally that Great-tailed Grackles are determinate layers, an attribute opposite to that sometimes associated with CBP. Despite the absence of CBP, Great-tailed Grackles rejected 8% of experimentally introduced conspecific eggs, rarely rejecting or damaging their own eggs. Conspecific eggs added to nests during incubation tended to be rejected more frequently than eggs switched between nests, and eggs that differed the most from the host's eggs tended to be rejected sooner. There was no relationship between rejection and the stage of the nest cycle when experimental parasitism occurred; however, eggs were rejected faster when added during the prelaying and incubation stages than during laying. Evidence suggests, therefore, that egg rejection behavior in Great-tailed Grackles has not evolved in response to CBP.     

The effects of exposure and microbes on hatchability of eggs in open-cup and cavity nests

Many avian species initiate incubation before clutch completion, resulting in an asynchronous hatch of their eggs. Several studies suggest that early laid eggs in birds that exhibit synchronous hatching may be more vulnerable to the negative impacts of ambient temperatures and/or trans-shell infection by microbes. However, nearly all of these studies have exposed fertile eggs to environmental conditions in artificial cavity nests, and thus, the effects of exposure of eggs to environmental conditions in open-cup nests remains largely unknown. Therefore, we directly compared hatchability and rates of trans-shell infection in fertile domestic chicken eggs that were exposed for 1–5 days in either open-cup or cavity nests. Eggs in open-cup nests had significantly higher rates of trans-shell infection and lower hatchability than those in cavity nests. These differences may result from different environmental conditions in open-cup nests, as well as in rates of microbial colonization of eggs. Cavity nests maintained slightly higher temperatures than did open-cup nests in the same location; thus, eggs in cavity nests were exposed for a longer period of time to temperatures ≥27°C, the temperature at which antibacterial enzymatic activity is initiated in the albumen. Moreover, microbial growth on eggs was much higher in open-cup nests even when eggs in these nests were cleaned daily with alcohol. It may be that the increased exposure to rain events in open-cup nests may facilitate microbial growth and egg infection. Thus, our data suggest that open-cup nesters may face constraints on reproduction different from those that cavity nesters face, and therefore may make choices regarding incubation that reflect these challenges.     

Low frequency of observed cowbird parasitism on eastern kingbirds: host rejection, effective nest defense, or parasite avoidance?

Eastern kingbird (Tyrannus tyrannus) nests rarely are parasitizedby brown-headed cowbirds (Molotrus ater). Kingbirds are oneof a dozen or so species known to eject cowbird eggs from theirnests. We hypothesized that either kingbirds eject cowbird eggsso quickly that researchers normally do not detect the eggsduring daily nest inspections, or that cowbirds avoid parasitizingkingbirds. We tested these alternative hypotheses by experimentallyintroducing real cowbird eggs into eastern kingbird nests duringthe pre-egg, early laying, late laying, and incubation stages.We recorded the interval between "parasitism" and ejection ofthe cowbird eggs. Although kingbirds ejected 87 of 88 cowbirdeggs placed in their nests, about 40% of the eggs remained innests for more than 24 h. Thus, during daily nest inspectionswe should have observed cowbird eggs if nests were parasitizedat all. In fact, we detected only one parasitized nest amongthe 402 inspected daily. The time for ejection was longest atnests parasitized early in laying, and shorter at nests parasitizedbefore and after. This variation in ejection times may reflectthe time kingbirds require to learn to recognize their own eggs.Although kingbirds defend their nests aggressively, they donot respond to female cowbirds as unique threats and do notguard their nests before sunrise when cowbirds lay. We concludethat cowbirds avoid parasitizing eastern kingbirds because theireggs most likely will be wasted. The rejection behavior persistspossibly because it is almost cost-free (a maximum of 0.07 kingbirdegg lost or damaged per cowbird egg ejected), or it evolvedin response to conspecific rather than cowbird parasitism. Foreignkingbird eggs introduced into nests at different nest stageswere ejected only during the pre-egg stage. This result supportsthe hypothesis that rejection behavior in eastern kingbirdsevolved in response to cowbird parasitism.     

Egg concealment is an antipredatory strategy in a cavity‐nesting bird

Birds have developed different behavioural strategies to reduce the risk of predation during the breeding period. Bird species that nest in the open often cover their eggs to decrease the risk of predators detecting the clutches. However, in cavity nesters, the potential functions of egg covering have not been explored despite some bird species that nest in cavities also covering their eggs as open nesters do. We analysed whether egg covering is an antipredatory behaviour in the blue tit (Cyanistes caeruleus). We simulated an increase in the perceived risk of predation at experimental nests by adding predator scent inside the nest boxes during the egg‐laying period, whilst adding lemon essence or water to control nest boxes. Birds exposed to predator chemical cues in the nest of experimental pairs more frequently covered their eggs than birds exposed to an odorous control. These results suggest that egg covering may have evolved as an antipredatory behaviour also in cavity nesters to reduce the risk of egg predation and thus increase reproductive success in birds.     

The moult of British thrushes and chats

Incubated clutches of the Mandarin Duck were larger earlier in the breeding season. At all nests there were days on which a single egg, and days on which no egg was added to a clutch; days on which more than one egg was added to a clutch were less frequent. Larger clutches had proportionately more days on which two or more eggs were added; the proportion of days on which no eggs were added was unrelated to clutch size. Larger clutches had a relatively shorter laying period than expected, but a longer incubation period than smaller clutches. Unhatched eggs resulted mainly from embryonic mortality, which was concentrated in the period just prior to pipping. Similar proportions of unhatched eggs resulted from eggs laid into clutches before and after the start of incubation. Larger clutches did not have proportionately more unhatched eggs. As all clutches had at least one day on which no eggs were laid, it is proposed that the proportion of nests parasitized was greater than that estimated from nests with more than one egg laid on any one day.     

Plasticity in incubation behavior and shading by king rails Rallus elegans in response to temperature

King rails experience a wide range of temperatures during the course of the breeding season throughout their rapidly contracting geographic range. Incubating parent birds are adapted to keep their eggs within a temperature range appropriate for embryo development, but king rail clutches are at risk of exceeding lethal temperatures in the latter half of the nesting season. We investigated whether behavioral plasticity during incubation enables parents to maintain clutch temperature within tolerable limits for embryo development. Video revealed that king rail parents interrupted incubation to stand above and shade their eggs. We tested the hypothesis that the onset of shading was a direct response to ambient temperature (adaptive plasticity). We monitored clutch temperature directly by experimentally adding into clutches a model egg embedded with a programmable iButton. We measured ambient temperature at the nest site simultaneously. Parents spent proportionately more time shading and less time incubating their eggs at higher ambient temperatures. Shading may primarily function in cooling the parent. The frequency and duration of shading bouts were significantly greater at higher ambient temperatures. Parents also took more frequent but shorter recesses in hotter conditions. Diurnal recesses exposed eggs to direct sunlight, and the highest clutch temperatures were recorded under these conditions. Complete hatching failure in at least one nest was attributable to high clutch temperature for an extended period. Because mean ambient temperature increases throughout the breeding season, we investigated seasonal patterns in onset of incubation and its effect on hatching rate. Later in the season, parents tended to initiate incubation earlier, and hatching asynchrony increased significantly. Together these results suggest that breeding king rails may be constrained in their ability to cope with sustained high temperatures should seasonal averages continue to rise as predicted.     

Nest Sanitation Plays a Role in Egg Burial by Yellow Warblers

Some hosts of the brown-headed cowbird ( Molothrus ater ) possess defences that eliminate all or most parasitism costs. Yellow warblers ( Dendroica petechia ) bury cowbird eggs, possibly to clean nests rather than serving strictly as an anti-parasite defence, as non-egg-shaped objects have been ejected, buried, or deserted by other hosts. With two experiments, we tested the 'nest sanitation' hypothesis by recording warblers' responses to objects (1) similar in volume, mass, and colour to cowbird eggs, and (2) half the mass and volume (more easily ejected), placed into nests before and during incubation. We compared outcomes at control nests with responses to objects that were dissimilar (stars) and moderately similar (dumbbells) to eggs, and to real cowbird and warbler eggs. We tested whether rejection (1) declines from stars through dumbbells and real eggs, (2) is similar between stages, and (3) non-egg-shaped objects are ejected because this is the least costly rejection method. Large stars were rejected (most buried) significantly more frequently (43.8%) than cowbird eggs (16.3%) in pre-incubation, suggesting that warblers reject objects shaped unlike their own eggs to rid nests of debris. Objects spent less time in nests the more they diverged from eggs. Warblers rarely rejected large stars and dumbbells, and cowbird eggs during incubation, possibly because burial and desertion are too costly by this time. Responses to small stars and dumbbells, and to foreign yellow warbler eggs did not differ between stages; also warblers rejected stars, mostly by ejection and selective burial, more frequently (28.8%) than dumbbells (1.3%) and warbler eggs (0%). Rejection by yellow warblers, especially burial, may keep nests clean, but also functions in rejecting cowbird eggs.