Nest destruction elicits indiscriminate con‐ versus heterospecific brood parasitism in a captive bird

Following nest destruction, the laying of physiologically committed eggs (eggs that are ovulated, yolked, and making their way through the oviduct) in the nests of other birds is considered a viable pathway for the evolution of obligate interspecific brood parasitism. While intraspecific brood parasitism in response to nest predation has been experimentally demonstrated, this pathway has yet to be evaluated in an interspecific context. We studied patterns of egg laying following experimental nest destruction in captive zebra finches, Taeniopygia guttata, a frequent intraspecific brood parasite. We found that zebra finches laid physiologically committed eggs indiscriminately between nests containing conspecific eggs and nests containing heterospecific eggs (of Bengalese finches, Lonchura striata vars. domestica), despite the con‐ and heterospecific eggs differing in both size and coloration. This is the first experimental evidence that nest destruction may provide a pathway for the evolution of interspecific brood parasitism in birds.     

Coevolution between Himalayan cuckoos and 2 sympatric Pycnonotidae hosts

Selection due to cuckoo parasitism is responsible for the evolution of anti-parasitism defenses in hosts. Different host species breeding sympatrically with a single parasitic cuckoo may evolve different strategies to reduce the risk of counter cuckoo parasitism, resulting in different interactions between cuckoos and hosts in areas of sympatry. Here, we studied the coevolutionary interactions between Himalayan cuckoos Cuculus saturatus and 2 sympatric and closely related potential hosts belonging to the family Pycnonotidae, the brown-breasted bulbul Pycnonotus xanthorrhous and the collared finchbill Spizixos semitorques. We investigated parasitism rates and nest-site selection (nest height, nest cover, human disturbance, perch height, forest distance, and degree of concealment) related to parasitism risk, nest defense against a cuckoo dummy, and egg rejection against cuckoo model eggs. Bulbuls used specific nest sites that were further away from forests than those of finchbills, and they behaved more aggressively toward cuckoos than finchbills. In contrast, bulbuls possessed moderate egg rejection ability, whereas the finchbill rejected 100% of cuckoo model eggs. We suggest that selection of a nest site away from forests by the bulbul explains the absence of parasitism by Himalayan cuckoos. We suggest that these interspecific differences in nest-site selection and nest defense indicate alternative responses to selection due to cuckoos.     

Size and material of model parasitic eggs affect the rejection response of Western Bonelli's Warbler Phylloscopus bonelli

Given the high costs of brood parasitism, avian hosts have adopted different defences to counteract parasites by ejecting the foreign egg or by deserting the parasitized nest. These responses depend mainly on the relative size of the host compared with the parasitic egg. Small hosts must deal with an egg considerably larger than their own, so nest desertion becomes the only possible method of egg rejection in these cases. The use of artificial model eggs made of hard material in egg‐recognition experiments has been criticized because hard eggs underestimate the frequency of egg ejection. However, no available studies have investigated the effect of softer material. Here, we test the potential effect of size of dummy parasitic eggs in relation to egg‐rejection behaviour (egg ejection and nest desertion rates) in Western Bonelli's Warbler Phylloscopus bonelli, a small host, using plasticine non‐mimetic eggs of three different sizes. In addition, we tested the potential effect of material, comparing ejection and desertion responses between real and plasticine eggs. As predicted, small eggs were always ejected, whereas nest desertion occurred more frequently with large eggs, thus suggesting that nest desertion occurs because of the constraints imposed by the large eggs. We found that plasticine may misrepresent the responses to experimental parasitism, at least in small host species, because this material facilitates egg ejection, provoking a decrease in nest desertion rate. Thus, particular caution is needed in the interpretation of the results in egg‐rejection experiments performed using dummy eggs made of soft materials.     

Parasitism of maternal investment selects for increased clutch size and brood reduction in a host

The allocation of resources to young that will ultimately beleft to die appears counterintuitive. Yet obligate brood reductionhas evolved in a number of species, despite the waste of reproductiveinvestment this may incur. Here we test whether brood parasitismcould be one factor leading to the evolution of obligate broodreduction because surplus eggs in the nest during incubationoffer some protection from the costs of parasitism. Surpluseggs could benefit females in two ways. First, additional eggsmay protect against the direct costs of parasitism by facilitatingrecognition and removal of parasitic eggs with greater accuracy.Second, additional eggs may protect against the indirect costsof parasitism as parasites often damage or remove host eggswhen entering the host nest; surplus eggs may be an essentialinsurance strategy against this damage. We test these possibilitiesin the Montezuma Oropendola (Psarocolius Montezuma), a speciesexperiencing high levels of parasitism by Giant Cowbirds (Scaphiduraoryzivora) throughout their range. Overall rejection rates ofcowbird eggs were high (72%), and experimental addition of parasiticeggs to empty, one-, and two-egg nests demonstrated that recognitionsuccess was unaffected by the presence of additional host eggsfor comparison. However, the value of surplus eggs when oneegg was removed or damaged by a parasite was high; 31.6% ofsuccessful two-egg clutches lost a single egg during incubationand would have failed to produce a chick without a second egg.This was directly attributable to parasitism in at least 33%of all cases. Therefore, despite highly developed host defensesagainst direct costs of parasitism (recognition and removalof parasitic eggs), the associated indirect costs (egg damageand removal) could play an important role in selection for aclutch size that results in more chicks than can be raised.     

Nest sanitation behavior does not increase the likelihood of parasitic egg rejection in herring gulls

Birds’ behavioral response to brood parasitism can be influenced not only by evolution but also by context and individual experience. This could include nest sanitation, in which birds remove debris from their nests. Ultimately, nest sanitation behavior might be an evolutionary precursor to the rejection of parasitic eggs. Proximately, the context or experience of performing nest sanitation behavior might increase the detection or prime the removal of parasitic eggs, but evidence to date is limited. We tested incubation-stage nests of herring gulls Larus argentatus to ask whether nest sanitation increased parasitic egg rejection. In an initial set of 160 single-object experiments, small, red, blocky objects were usually rejected (18 of 20 nests), whereas life-sized, 3D-printed herring gull eggs were not rejected whether red (0 of 20) or the olive-tan base color of herring gull eggs (0 of 20). Next, we simultaneously presented a red, 3D-printed gull egg and a small, red block. These nests exhibited frequent nest sanitation (small, red block removed at 40 of 48 nests), but egg rejection remained uncommon (5 of those 40) and not significantly different from control nests (5 of 49) which received the parasitic egg but not the priming object. Thus, performance of nest sanitation did not shape individuals’ responses to parasitism. Interestingly, parents were more likely to reject the parasitic egg when they were present as we approached the nest to add the experimental objects. Depending on the underlying mechanism, this could also be a case of experience creating variation in responses to parasitism.     

Parental care of a cowbird host: caught between the costs of egg-removal and nest predation

Avian brood parasites reduce host fitness through the addition of parasitic eggs and the removal of host eggs. Both parasitic egg-addition and host egg-removal may be important sources of selection on host behaviour, creating fitness trade-offs with selection imposed by nest predation. However, the relative costs hosts suffer from egg-addition and host egg-removal and the responses to these costs are largely unstudied. Through experimental manipulations and observations, we demonstrate that increased nest attentiveness by female yellow warblers (Dendroica petechia) reduces the cost of brood parasitism by reducing egg-removal by brown-headed cowbirds (Molothrus ater). However, female attentiveness does not reduce the addition of parasitic eggs. Experimentally parasitized females respond to the threat of egg-removal by increasing nest attentiveness. Increased attentiveness, however, reduces time for females to gather food and requires males to visit the nest more often to feed incubating females. This increased activity in turn increases the risk of nest predation. Thus, brood parasitism (specifically egg-removal) and nest predation produce conflicting selection on incubation strategies, as parasitized hosts are caught between the costs of egg-removal by brood parasites, and the costs of increased nest predation if the female spends more time on the nest to reduce egg-removal.     

How to Spot a Stranger's Egg? A Mimicry‐Specific Discordancy Effect in the Recognition of Parasitic Eggs

Egg discrimination by hosts is an antiparasitic defence to reject foreign eggs from the nest. Even when mimetic, the presence of brood parasitic egg(s) typically alters the overall similarity of all eggs in a clutch, producing a discordant clutch compared to more homogenous clutches of composed only of hosts’ own eggs. In multiple parasitism, the more foreign eggs are laid in the nest, the more heterogeneous the overall clutch appears. Perceptual filters and recognition templates cannot explain the known pattern of lower rejection rates of foreign eggs in multiple vs. single parasitism. We therefore assessed the role of clutch homogeneity and manipulated the colour of one or more eggs in the clutches of great reed warbler (Acrocephalus arundinaceus) hosts of common cuckoos (Cuculus canorus). Varying the colours of both the majority and the minority eggs caused predictable shifts in the rejection of the focal egg(s), and ejection rates of the minority egg colour consistently increased but only when it belonged to a more mimetic egg colour, relative to the less mimetic colour of majority eggs. The results imply that in addition to sensory filters, and template‐based cognitive decision rules, discordancy‐based rejection is affected by the overall clutch appearance and interacts with specific colours varying in the extent of mimicry, to contribute to the recognition decisions of hosts to reject parasitic eggs.     

Effects of nest-site characteristics and parental activity on cowbird parasitism and nest predation in Brown-and-yellow Marshbirds

ABSTRACT Nest‐site selection and nest defense are strategies for reducing the costs of brood parasitism and nest predation, two selective forces that can influence avian nesting success and fitness. During 2001–2002, we analyzed the effect of nest‐site characteristics, nesting pattern, and parental activity on nest predation and brood parasitism by cowbirds (Molothrus spp.) in a population of Brown‐and‐yellow Marshbirds (Pseudoleistes virescens) in the Buenos Aires province, Argentina. We examined the possible effects of nest detectability, nest accessibility, and nest defense on rates of parasitism and nest predation. We also compared rates of parasitism and nest predation and nest survival time of marshbird nests during the egg stage (active nests) with those of the same nests artificially baited with passerine eggs after young fledged or nests failed (experimental nests). Most nests (45 of 48, or 94%) found during the building or laying stages were parasitized, and 79% suffered at least one egg‐predation event. Cowbirds were responsible for most egg predation, with 82 of 107 (77%) egg‐predation events corresponding to eggs punctured by cowbirds. Nests built in thistles had higher rates of parasitism and egg predation than nests in other plant, probably because cowbirds were most active in the area where thistles were almost the only available nesting substrate. Parasitism rates also tended to increase as the distance to conspecific nests increased, possibly due to cooperative mobbing and parental defense by marshbirds. The proportion of nests discovered by cowbirds was higher for active (95%) than for experimental (29%) nests, suggesting that cowbirds used host parental activity to locate nests. Despite active nest defense, parental activity did not affect either predation rates or nest‐survival time. Thus, although nest defense by Brown‐and‐yellow Marshbirds appears to be based on cooperative group defense, such behavior did not reduce the impact of brood parasites and predators.     

On the origin and rarity of interspecific nest parasitism in birds

Interspecific nest parasitism is surprisingly rare in birds given the potential advantages for the parasite of exploiting the parental care of other species. One possibility is that chicks will not thrive with the parental care and food of heterospecifics. I simulated parasitism in nonparasitic congeners by switching eggs between nests of three species of titmice (great tit Parus major, blue tit Parus caeruleus, and coal tit Parus ater). The experiment showed that compatibility of parental care was not a constraint preventing parasitism. I also used the model system to compare fitness consequences of inter- and intraspecific nest parasitism, addressing the problem of which form is ancestral. Fledging success (body mass, survival) was higher when an egg was added to the nest of a smaller species than to the nest of a conspecific and also higher when the parasitic chick hatched early rather than late relative to host chicks. This suggests that interspecific nest parasitism may not require a stage of intraspecific nest parasitism before evolving but may start from a larger species directly exploiting the parental care of a smaller species or a species with shorter incubation period directly exploiting a species with longer incubation period.     

Detecting conspecific brood parasitism using egg morphology in black brant Branta bernicla nigricans

Numerous methods have been proposed to indirectly detect conspecific brood parasitism (CBP) in birds. Egg morphology has been suggested as a predictor of parasitism, assuming that variation in egg size is greater among females than within females. Here we use microsatellite data to assess the use of egg morphology to detect CBP in a sample of black brant Branta bernicla nigricans nests. We attempted to repeat a previously demonstrated technique using cluster analysis and maximum Euclidean distance (MED) to detect parasitized nests within black brant. Additionally we attempted a new technique based on a discriminant function analysis of egg morphology in an attempt to detect brood parasitic eggs. When detecting parasitized nests using egg morphology, the cluster analysis revealed that the MED between the two most dissimilar eggs in each nest was significantly greater for parasitized nests than for non‐parasitized nests (1.62±0.06 and 1.43±0.08, respectively). The extent of overlap in sizes of eggs between parasitized and non‐parasitized nests, however, was such that we were unable to effectively identify parasitized nests. In most cases for each parasitized nest correctly identified, 3 non‐parasitized nests were incorrectly identified as parasitic. When we attempted to detect parasitic eggs we found that parasitic eggs were more different from the expected egg volume than host eggs: mean absolute residual volume of parasitic eggs=2.59±5.79 cm³ while that for host eggs=1.82±2.14 cm³. Overall, we found that the discriminant function analysis was moderately effective in determining whether eggs belonged to the host female using a resubstitution technique (error rate=9.71%) or a jackknife technique (error rate=6.12%). Additionally, we found a higher but moderate error rate when using an independent data set to validate the function (error rate=14.07%). In both cases, however, parasitic eggs accounted for most of the error and were not correctly classified 75%, 70% and 100% of the time respectively. We suggest when developing a predictive function for detecting conspecific brood parasitism based on egg morphology that an appropriate technique be used to validate the function, particularly those techniques that utilize unambiguous identifiers such as molecular and protein fingerprinting techniques.     

Egg ejection cost can limit defence strategies against brood parasitism

A cost associated with the evolution of antiparasite strategies is the failure to recognize parasitic eggs, leading the host to evict its own eggs. However, there is evidence that birds recognize their own eggs through imprinting. This leads to the question of why birds accept parasitic eggs if such eggs can be identified. Here, we tested whether egg ejection per se can be costly due to increased predation risk to the remaining clutch and whether olfactory or visual cues of egg ejection increase predation. We carried out three field experiments to answer the following questions: (a) Does ejecting an egg increase nest predation risk? (b) Does the presence of olfactory cues, such as the smell of a broken egg, increase nest predation risk? And (c) Does the presence of visual cues, such as an egg shell below the nest, increase nest predation risk? We found evidence that egg ejection increases nest predation and that olfactory cues alone also increase nest predation. The presence of visual cues did not change predation rates. These data indicate that egg ejection is costly for both host and parasitic eggs that may remain in the nest. Our results suggest why host and parasite eggs are commonly found within the same nests, despite the possibility that hosts recognize and could possibly eject the parasite’s egg.     

Nest desertion cannot be considered an egg‐rejection mechanism in a medium‐sized host: an experimental study with the common blackbird Turdus merula

Two main mechanisms of egg rejection, the main defence of hosts against brood parasites, have been described: ejection and desertion. Desertion of the parasitized nest is much more costly and is usually exhibited by small‐sized host species unable to remove the parasitic egg. However, nest desertion is frequently assumed to be an anti‐parasite strategy even in medium or large‐sized host species. This assumption should be considered with caution because: 1) large‐sized hosts able to eject the parasitic egg should eject it rather than desert the nest, and 2) breeding birds may desert their nests in response to different disturbances other than brood parasitism. This problem is especially important in the common blackbird Turdus merula because this is a medium‐sized species, potential host of the common cuckoo Cuculus canorus, in which desertion has been frequently reported as a response to cuckoo egg models. Here, we seek to determine whether nest desertion can be considered a response unequivocally directed to the parasitic egg in medium‐sized hosts using the blackbird as the study species. In an experimental study in which we have manipulated levels of mimicry and size of experimental eggs, we have found that both colour (mimetic and non‐mimetic; at least for human vision) and size (small, medium, and large) significantly affected ejection rates but not nest desertion rates. In fact, although large eggs disproportionally provoked nest desertion more frequently than did small or medium‐sized eggs, cuckoo‐sized parasitic eggs were not deserted allowing us to conclude that desertion is unlikely to be an adaptive response to brood parasitism at least for this species.     

Nest desertion by a cowbird host: an antiparasite behavior or a response to egg loss?

Natural selection can favor songbirds that desert nests containingeggs of the parasitic brown-headed cowbird (Molothrus ater).However, the high variability in desertion of parasitized nestswithin species is perplexing in light of the typically highcosts of parasitism. Because nest desertion can also be a responseto partial clutch predation, we first asked if Bell's vireos(Vireo bellii) deserted nests in response to the presence ofcowbird eggs (antiparasite response hypothesis) or to egg removalby predators and female cowbirds (egg predation hypothesis).Second, we asked whether variation in nest desertion was dueto intrinsic differences among individuals or to variation innest contents. We monitored a large number of nests (n = 494)and performed a clutch manipulation experiment to test thesehypotheses. The number of vireo eggs that remained in a nestwas a strong predictor of desertion both within and among pairs.Neither the presence of a single cowbird egg, which leads tonest failure for this host, nor the number of cowbird eggs receivedin a vireo nest influenced nest desertion. Furthermore, vireosdid not desert experimental nests when we immediately exchangedcowbird eggs for vireo eggs but deserted if we removed vireoeggs and replaced them with cowbird eggs the following morning.Desertion of parasitized nests by Bell's vireos can be almostentirely explained as a response to partial or complete clutchloss and does not appear to have been altered by selection frombrood parasitism.     

No change in common cuckoo Cuculus canorus parasitism and great reed warblers’ Acrocephalus arundinaceus egg rejection after seven decades

The coevolutionary process among avian brood parasites and their hosts involves stepwise changes induced by the antagonistic selection pressures of one on the other. As long‐term data on an evolutionary scale is almost impossible to obtain, most studies can only show snapshots of such processes. Information on host behaviour, such as changes in egg rejection rates and the methods of rejection are scarce. In Hungary there is an interesting case between the common cuckoo Cuculus canorus and the great reed warbler Acrocephalus arundinaceus, where the level of parasitism is unusually high (around 50%). We compared host rejection rates and methods of rejection from within our own project to that of an early study carried out and published almost 70 yr ago in the same region. Our comparisons revealed high and stable rates of parasitism (range: 52–64%), and marked fluctuations in the ratio of multiply parasitized nests (range: 24–52%). No difference was revealed in egg rejection rates after 7 decades (34–39%). Linear mixed‐effects modelling revealed no year effect on the type host responses toward the parasitic egg(s) during the years of study (categorized as acceptance, ejection, burial, and nest desertion). Cuckoo egg rejection was primarily affected by the type of parasitism, as more cuckoo eggs were rejected during single parasitism than from multiply parasitized nests. Our comparison did not reveal any directional changes in this cuckoo–host relationship, except a slight decrease in the frequency of multiple parasitism, which is likely to be independent from coevolutionary processes.     

Does the house sparrow Passer domesticus represent a global model species for egg rejection behavior?

Conspecific brood parasitism (CP) is a facultative breeding tactic whereby females lay their eggs in the nests of conspecifics. In some species, potential hosts have evolved the ability to identify and reject foreign eggs from their nest. Previous studies suggest that the ubiquitous house sparrow Passer domesticus in Spain and South Africa employs both CP and egg rejection, while a population in China does not. Given the species’ invasive range expansions, the house sparrow represents a potentially excellent global model system for parasitic egg rejection across variable ecological conditions. We examined the responses of house sparrows to experimental parasitism at three geographically distinct locations (in Israel, North America, and New Zealand) to provide a robust test of how general the findings of the previous studies are. In all three geographic regions egg rejection rates were negligible and not statistically different from background rates of disappearance of control eggs, suggesting that the house sparrow is not a suitable model species for egg rejection experiments on a global scale.     

Conspecific brood parasitism and population dynamics

Conspecific brood parasitism (CBP), defined as parasitic laying of eggs in a conspecific nest without providing parental care, occurs in insects, fishes, amphibians, and many birds. Numerous factors have been proposed to influence the evolution of CBP, including nest site limitation; effects of brood size, laying order, or parasitic status on offspring survival; randomness of parasitic egg distribution; adult life-history trade-offs; and variation in parental female quality or risk of nest predation. However, few theoretical studies consider multiple possible types of parasitism or the interplay between evolution of parasitism and population dynamics. We review existing theory of CBP and develop a synthetic modeling approach to ask how best-of-a-bad job parasitism, separate-strategies parasitism (in which females either nest or parasitize), and joint-strategies parasitism (in which females can both nest and parasitize) differ in their evolutionary conditions and impacts on population dynamics using an adaptive dynamics framework including multivariate traits. CBP can either stabilize or destabilize population dynamics in different scenarios, and the role of comparable parameters on evolutionarily stable strategy parasitism rate, equilibrium population size, and population stability can differ for the different modes of parasitism.     

Molecular identification of brood‐parasitic females reveals an opportunistic reproductive tactic in ruddy ducks

In many taxa, females lay eggs in the nests of other conspecifics. To determine the conditions under which conspecific brood parasitism develops, it is necessary to identify parasitic offspring and the females who produce them; however, for most systems parasitism can be difficult to observe and most genetic approaches have relatively low resolving power. In this study, we used protein fingerprinting from egg albumen and 10 microsatellite loci to genetically match parasitic ducklings to their mothers in a population of ruddy ducks (Oxyura jamaicensis). We found that 67% of nests contained parasitic offspring, and we successfully identified their mothers in 61% of the cases. Of the parasitic females identified, 77% also had nests of their own (i.e. a dual tactic, where females both nest and lay parasitically), and we found no evidence that parasitic females pursued a specialist (parasitism only) tactic. We also found that parasitic egg laying was not influenced by nest loss, predation or female condition. Thus, in contrast to most waterfowl studied to date, female ruddy ducks appear to lay parasitic eggs whenever the opportunity arises.     

Egg discrimination in the Australian reed warbler (Acrocephalus australis): rejection response toward model and conspecific eggs depending on timing and mode of artificial parasitism

In a coevolutionary arms race between an interspecific broodparasite and its host species, both are expected to evolveadaptations and counteradaptations. We studied egg discriminationin the Australian warbler (Acrocephalus australis). This speciesis currently not significantly parasitized by the seven speciesof cuckoo for which it is a suitable host. However, experimentalbrood parasitism in the warbler revealed a fine tuned egg discriminationresponse towards non-mimetic and conspecific eggs, the firstsuch evidence in an Australian passerine: (1) non-mimetic eggswere significantly more often rejected than conspecific eggs;(2) only non-mimetic dummy eggs were rejected selectively,whereas rejection of conspecific eggs entailed a rejectioncost; (3) replacement of a host's egg with a conspecific eggduring egg laying resulted in a significantly higher rejectionrate than after the day of clutch completion; (4) by contrast,rejection rate after addition of a conspecific egg was independentof nest stage; (5) conspecific eggs introduced into a clutchduring the egg laying period led to a significantly highernest desertion rate and a lower egg ejection rate than afterthe day of clutch completion; and (6) addition of a conspecificegg led to egg ejection while egg replacement with a conspecificegg led to nest desertion. The fact that this species respondsdifferentially toward different modes of artificial parasitismsuggests that its egg discrimination has evolved to minimizethe costs of rejection and parasitism. The ability to rejecthighly mimetic conspecific eggs may explain the current paucityof brood parasitism in this species. The significance of thisfor brood parasite-host coevolution is discussed.     

Experimental support for the role of nest predation in the evolution of brood parasitism

In 1965, Hamilton and Orians (HO) hypothesized that the starting point for the evolution of obligate interspecific brood parasitism in birds was the facultative laying of physiologically committed eggs in neighbouring active nests of con‐ and heterospecifics, following predation of a bird’s own nest during the laying stage. We tested this prediction of the HO hypothesis by using captive pairs of zebra finches (Taeniopygia guttata), a species with evidence for intraspecific parasitism both in the wild and in captivity. As predicted, in response to experimental nest removal, subjects laid eggs parasitically in simulated active conspecific nests above chance levels. Across subsequent trials, we detected both repeatability and directional change in laying patterns, with some subjects switching from parasitism to depositing eggs in the empty nest. Taken together, these results support the assumptions and predictions of the HO hypothesis, and indicate that the zebra finch is a potential model species for future behavioural and genetic studies in captive brood parasite research.     

Nest Sanitation as the Evolutionary Background for Egg Ejection Behaviour and the Role of Motivation for Object Removal

Higher interclutch colour variation can evolve under the pressure of brood parasitism to increase the detection of parasitic eggs. Nest sanitation could be a prerequisite for the evolution of anti-parasite defence in terms of egg ejection. In this respect, we used nest sanitation behaviour as a tool to identify: i) motivation and its underlying function and, ii) which features provoke ejection behaviour. Therefore, we experimentally tested whether size, colour or shape may influence ejection behaviour using artificial flat objects. We found a high interclutch variation in egg colouration and egg size in our tree sparrow (Passer montanus) population. Using colour and size we were in fact able to predict clutch affiliation for each egg. Our experiments further revealed the existence of direct anti-parasite behaviours and birds are able to recognise conspecific eggs, since only experimentally-deposited eggs have been removed. Moreover, experiments with different objects revealed that the motivation of tree sparrows to remove experimental objects from their nests was highest during egg laying for objects of varying size, most likely because of parasitism risk at this breeding stage. In contrary, motivation to remove white objects and objects with edges was higher during incubation stage as behavioural patterns connected to hatching started to emerge. The fact that rejection rate of our flat objects was higher than real egg ejection, suggests that egg ejection in tree sparrows and probably more general in small passerines, to be limited by elevated costs to eject eggs with their beaks. The presence of anti-parasite behaviour supports our suggestion that brood parasitism causes variation in egg features, as we have found that tree sparrows can recognise and reject conspecific eggs in their clutch. In conclusion, in tree sparrows it seems that nest sanitation plays a key role in the evolution of the removal of parasitic eggs.