The importance of nest cleaning in egg rejection behaviour of great reed warblers Acrocephalus arandinaceas

We tested the importance of nest cleaning in egg rejection behaviour of the great reed warbler Acrocephalus arundinaceus in a highly parasitised population in which about 64% of nests are parasitised by the common cuckoo Cuculus canorus . Three types of objects of the same weight, texture and colour but with different shapes (dummy cuckoo eggs, sticks and disks) were placed into great reed warbler nests. We investigated the response of hosts in two stages of breeding: pre-incubation when the risk of brood parasitism is high, and during incubation when the risk of parasitism is low. The dummy cuckoo eggs were rejected less often than the other objects in both breeding stages, although we did not find any difference in the frequency of rejection between pre-incubation and incubation. We integrate these results into current views on the evolution of host–parasite interactions, and propose a hierarchical concept to understand egg rejection behaviour: (1) hosts reject all non-egg shaped objects as a general cleaning mechanism; (2) adaptations for the hosts' ability to recognise their own eggs allows them to distinguish these eggs from similar objects and parasitic eggs.     

Are blackcaps current winners in the evolutionary struggle against the common cuckoo?

Blackcaps Sylvia atricapilla reject artificial cuckoo eggs, and their eggs vary little in appearance within clutches, whereas among clutches eggs vary considerably. Low variation within clutches facilitates discrimination of parasitic eggs, whereas high variation among clutches makes it harder for the cuckoo to mimic the eggs of a certain host species. These traits have most probably evolved as counteradaptations against brood parasitism by the common cuckoo Cuculus canorus, even though blackcaps are not regularly parasitised today. In this study, we investigated how fine-tuned the rejection of parasitic eggs is in this species by introducing three types of eggs into their nests: a real non-mimetic egg the approximate size of a cuckoo egg, an artificial mimetic egg the size of a cuckoo egg and a real conspecific egg. As the rejection frequency of both mimetic and non-mimetic artificial cuckoo eggs has been shown to be high in previous studies, the variation in rejection behaviour between individuals is low, indicating that most individuals within the population are able to reject parasitic eggs. Thus, we predict that (1) the intraclutch variation in egg appearance should be generally low in all individuals, and that (2) regarding conspecific eggs, rejection decisions should be highly dependent on the degree of mimicry between parasitic and host eggs. We found support for these predictions, which indicates that due to their highly sophisticated countermeasures against brood parasitism, blackcaps can probably be regarded as current winners of the arms race with the common cuckoo. Furthermore, the high and consistent rejection frequency of cuckoo eggs found throughout Europe for this species supports the spatial habitat structure hypothesis, which claims that woodland-nesting species breeding near trees, like blackcaps, presumably experienced a high level of parasitism throughout their range in the past and, therefore, their rejection behaviour, once evolved, spread rapidly to all populations.     

Egg rejection behaviour in the great reed warbler (<Emphasis Type="Italic">Acrocephalus arundinaceus</Emphasis>): the effect of egg type

Egg discrimination in hosts of the common cuckoo Cuculus canorus is frequently studied by experimental parasitism, using model cuckoo eggs. We compared egg rejection behaviour of the great reed warbler Acrocephalus arundinaceus to either model cuckoo eggs made of plastic or painted real host eggs. We simultaneously parasitised host nests by two different egg types to simulate cuckoo parasitism. A previous study revealed very similar, ca. 70%, rejection rates against both of these egg types (beige or bluish background colour maculated with dark brown) when they were used for single parasitism. In the present study we showed 96% average rejection rates against these egg types when they were applied in multiple experimental parasitism, causing a more predictable output for rejection behaviour. Hard plastic eggs and painted real eggs were rejected at similar frequencies, and videotaping revealed that model egg rejection caused extra work for great reed warblers. We revealed a new type of rejection behaviour, when hosts tried to eject hard-shelled model cuckoo eggs: Hosts made little holes in the middle part of these plastic eggs by pecking them several times before ejection, as if seeking the possibility to pierce and hold these eggs in their bills. Painted real eggs were rejected by actually puncturing the eggshell and holding them in the bill during ejection. No instances of grasp ejection were recorded during filming. Most experimental eggs of either type were ejected within 1 day after the introduction of the eggs, indicating that hosts made their rejection decisions quickly. Our observations suggest the lack of plasticity in the mode and timing of ejection behaviour towards experimental cuckoo eggs of different types in great reed warblers.     

No evidence for recognition errors in Acrocephalus warblers

Failure to recognise own eggs (recognition errors) may be an important selective force behind acceptance of parasitic eggs, leading to a balance between rejecters and acceptors in a host population (the equilibrium hypothesis). We predicted that recognition errors should occur frequently among host species with intermediate rejection rates, whose rejection behaviour shows many conditional responses. The reed warbler Acrocephalus scirpaceus and great reed warbler A. arundinaceus fulfil these requirements. These two species were therefore used in an experiment where host birds were exposed to a common cuckoo Cuculus canorus dummy, either <2 m or 5–10 m from the nest, at fishponds in southern Moravia (Czech Republic). The hosts responded to the cuckoo dummy, great reed warblers being much more aggressive than reed warblers, and both species being more aggressive towards the dummy when it was close to the nest than when it was farther away. We furthermore predicted that there should be more eggs rejected (ejected or nest abandoned) due to recognition errors among hosts exposed to a dummy close to the nest than among both those exposed to a dummy farther away from the nest and towards controls not exposed to cuckoo dummies. When comparing egg loss between groups of birds that were exposed to a cuckoo dummy with those that were not, we found no significant difference. However, partial egg loss was frequent among hosts in the studied population, most probably due to cuckoo depredation. We discuss why there were no detectable recognition errors in the studied population, when other researchers have claimed to have found such errors in host populations elsewhere.     

Rejection of artificial cuckoo (Cuculus canorus) eggs in relation to variation in egg appearance among reed warblers (Acrocephalus scirpaceus)

Passerines that are exposed to brood parasitism can evolve reduced intraclutch variation in egg appearance to facilitate recognition and rejection of the parasitic egg. This has been shown to be true for European passerine species that are assumed to have participated in an evolutionary arms race with the cuckoo (Cuculus canorus). However, few investigations have been carried out with the aim of finding out whether there is a relationship between these two traits within a species. In this study, we compare the level of intraclutch variation in egg appearance and the rejection of an unlike parasitic egg within a population of reed warblers (Acrocephalus scirpaceus) in the south-eastern part of the Czech Republic. We parasitized reed warbler nests with an artificial non-mimetic cuckoo egg, and then monitored the reaction of the hosts. In 27 out of 48 nests (56.3%) the parasitic egg was rejected. The rejecter pairs had a statistically significantly lower intraclutch variation in egg appearance than the acceptor pairs. We discuss possible explanations for the observed relationship between rejection of unlike eggs and intraclutch variation in egg appearance within this population of reed warblers. The results are consistent with the evolutionary arms race hypothesis, but the intermediate rejection rate found in this population could also be maintained by an equilibrium between acceptors and rejecters due to rejection costs.     

Do Nest Light Conditions Affect Rejection of Parasitic Eggs? A Test of the Light Environment Hypothesis

Discrimination of foreign eggs is one of the most studied aspects of host defences against avian brood parasites. Although many factors affecting host egg‐recognition processes have already been evaluated, only a few attempts have been made to test the importance of light conditions in microhabitats of host nests. Here, we examined whether the objectively measured nest light environment affects great reed warbler (Acrocephalus arundinaceus) responses towards real common cuckoo (Cuculus canorus) eggs. More specifically, we predicted that parasitic eggs will be rejected with a lower frequency from nests placed in darker conditions than those in lighter conditions. However, we found no effect of the ambient light on egg‐rejection behaviour alone, but the photosynthetically active radiation exhibited a positive interactive effect with chromatic contrast between cuckoo and host eggs. Most rejection events were accomplished when cuckoo eggs of poor mimicry were laid in well‐lit nests. Our study suggests that this phenomenon may have important implications for the evolution of egg mimicry and host egg discrimination. We encourage further testing of the light environment hypothesis in other host species breeding in variable nest microhabitats and light conditions.     

The common cuckoo Cuculus canorus is not locally adapted to its reed warbler Acrocephalus scirpaceus host

The obligate avian brood parasitic common cuckoo Cuculus canorus comprises different strains of females that specialize on particular host species by laying eggs of a constant type that often mimics those of the host. Whether cuckoos are locally adapted for mimicking populations of the hosts on which they are specialized has never been investigated. In this study, we first explored the possibility of local adaptation in cuckoo egg mimicry over a geographical mosaic of selection exerted by one of its main European hosts, the reed warbler Acrocephalus scirpaceus. Secondly, we investigated whether cuckoos inhabiting reed warbler populations with a broad number of alternative suitable hosts at hand were less locally adapted. Cuckoo eggs showed different degrees of mimicry to different reed warbler populations. However, cuckoo eggs did not match the egg phenotypes of their local host population better than eggs of other host populations, indicating that cuckoos were not locally adapted for mimicry on reed warblers. Interestingly, cuckoos exploiting reed warblers in populations with a relatively larger number of co-occurring cuckoo gentes showed lower than average levels of local adaptation in egg volume. Our results suggest that cuckoo local adaptation might be prevented when different cuckoo populations exploit more or fewer different host species, with gene flow or frequent host switches breaking down local adaptation where many host races co-occur.     

Breeding success of common cuckoos Cuculus canorus parasitising four sympatric species of Acrocephalus warblers

We investigated the level of parasitism, egg mimicry and breeding success of cuckoos parasitising four sympatric species of Acrocephalus warblers in southern Moravia, Czech Republic. The parasitism rate was highest in the marsh warbler Acrocephalus palustris (44.8%) followed by great reed warbler A. arundinaceus (33.8%), sedge warbler A. schoenobaenus (26.5%) and reed warbler A. scirpaceus (11.6%). Although the cuckoo eggs showed a high level of mimicry the eggs of the marsh warbler this host species rejected 72% of the cuckoo eggs, resulting in a cuckoo breeding success of only 4.3%. Cuckoo eggs laid in great reed warbler and reed warbler nests showed a similar hatching success, but the cuckoo chicks survived better in great reed warbler nests, resulting in a breeding success of 30.4%, as compared to 16.4% in nests of the reed warbler. The relationship between the level of parasitism, host rejection of cuckoo eggs, cuckoo chick survival and breeding success is discussed for the four host species.     

Explaining variation in brood parasitism rates between potential host species with similar habitat requirements

Host specialization evolved in many parasite-host systems. Evolution and maintenance of host specificity may be influenced by host life-history traits, active host selection by the parasite, and host anti-parasite strategies. The relative importance of these factors is poorly understood in situations that offer parasites a choice between hosts with similar habitat requirements. The common cuckoo Cuculus canorus is a generalist parasite on the species level, but individual females prefer particular host species. In reed beds of the Yellow River Delta, China, two potential hosts with similar nest characteristics, Oriental reed warblers Acrocephalus orientalis and reed parrotbills Paradoxornis heudei, breed in sympatry. We found that warblers were parasitized at much higher rates than parrotbills. Both hosts recognized and rejected non-mimetic model eggs well, indicating that they have been involved in an arms-race with cuckoos. Cuckoo eggs closely resembled warbler eggs, and such eggs were mostly accepted by warblers but rejected by parrotbills. Only warblers recognized adult cuckoos as a specific threat. Both hosts were equally good at raising cuckoo chicks. Low nest density, partial isolation by breeding time, small scale differences in nest and nest site characteristics, and high rejection rates of natural cuckoo eggs are likely cumulatively responsible for the low current parasitism rate in parrotbills. This study emphasizes the importance of integrating the study of general host life-history characteristics and specific anti-parasitism strategies of hosts across all breeding stages to understand the evolution of host specificity.     

Egg phenotype differentiation in sympatric cuckoo Cuculus canorus gentes

The brood parasitic common cuckoo Cuculus canorus consists of gentes, which typically parasitize only a single host species whose eggs they often mimic. Where multiple cuckoo gentes co‐exist in sympatry, we may expect variable but generally poorer mimicry because of host switches or inter‐gens gene flow via males if these also contribute to egg phenotypes. Here, we investigated egg trait differentiation and mimicry in three cuckoo gentes parasitizing great reed warblers Acrocephalus arundinaceus, marsh warblers Acrocephalus palustris and corn buntings Miliaria calandra breeding in close sympatry in partially overlapping habitat types. The three cuckoo gentes showed a remarkable degree of mimicry to their three host species in some but not all egg features, including egg size, a hitherto largely ignored feature of egg mimicry. Egg phenotype matching for both background and spot colours as well as for egg size has been maintained in close sympatry despite the possibility for gene flow.     

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.     

European Cuckoo Cuculus canorus parasitism and host's rejection behaviour in a heavily parasitized Great Reed Warbler Acrocephalus arundinaceus population

An unusually high frequency (64%) of European Cuckoo Cuculus canorus parasitism was found in Great Reed Warbler Acrocephalus arundinaceus clutches in central Hungary. Sixty-four per cent of the parasitized clutches contained one Cuckoo egg, 23% contained two, 10% had three and 3% had four. This means that 58% of the Cuckoo eggs were found in multiply parasitized clutches. In multiple parasitism the laying second Cuckoo removed an egg from the clutch randomly, so preferred neither the host eggs, nor the concurrent Cuckoo egg. Host response towards the parasitic eggs showed 66% acceptance, 12% ejection, 20% desertion and 2% egg burial. We found great variation in both the host and the parasitic egg colour and pattern. This reduces the chance that the parasitic egg's appearance matched that of the hosts' but, in spite of this, almost perfect mimesis was found in 28% of the Cuckoo eggs. Poorly mimetic Cuckoo eggs were more frequently rejected by Great Reed Warblers than parasite eggs that were very similar to the host eggs. This high level of mimicry sometimes makes it difficult for the observer to identify the parasitic egg, especially when it is similar in size to the host eggs. It is also difficult for the host, as shown by the relatively high recognition error and ejection cost.     

Do cuckoos choose nests of great reed warblers on the basis of host egg appearance?

Prevailing theory assumes cuckoos lay at random among host nests within a population, although it has been suggested that cuckoos could choose large nests and relatively active pairs within host populations. We tested the hypothesis that egg matching could be improved by cuckoos choosing nests in which host eggs more closely match their own, by assessing matching and monitoring nest fate in great reed warblers naturally or experimentally parasitized by eggs of European cuckoos. A positive correlation between cuckoo and host egg visual features suggests that cuckoos do not lay at random within a population, but choose nests and this improves egg matching: naturally parasitized cuckoo eggs were more similar to host eggs as perceived by humans and as measured by spectrophotometry. Our results suggest a hitherto overlooked step in cuckoo-host evolutionary arms races, and have nontrivial implications for the common experimental practice of artificially parasitizing clutches.     

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.     

Behaviour of African turdid hosts towards experimental parasitism with artificial red-chested cuckoo Cuculus solitarius eggs

The red-chested cuckoo Cuculus solitarius parasitises many passerines in Africa, but some common species sympatric with this brood parasite are rarely used as hosts. We tested the responses of three turdid hosts to parasitism with artificial cuckoo eggs. The kurrichane thrush Turdus libonyana , which is not regularly parasitized by the cuckoo, rejected 60% of mimetic model eggs and 81% of non-mimetic eggs. We observed female thrush behaviour during the first visit after parasitism, and thrushes appeared to be initially fooled by mimetic eggs in completed clutches in all cases, and incubated. By contrast, in half of the experiments with non-mimetic eggs, these were ejected by the thrushes, with the host grasping the egg and flying away with it. The time spent nest checking prior to ejection was only one third of the time spent nest checking when females decided to incubate the clutch, suggesting that females were immediately aware of a foreign egg in the nest. By contrast, southern olive thrushes T. olivaceous rejected all non-mimetic and accepted all mimetic model eggs, whereas cape robins Cossypha caffra accepted all model eggs, irrespective of whether or not they were mimetic. Our results support the hypothesis that rejection behaviour in these two thrush species evolved as a defence against interspecific nest parasitism, with thrushes appearing to be ahead in this particular host-parasite arms race. The cape robin, by contrast, appears not to reject cuckoo eggs, either because it is to unable to recognize them, or because the costs associated with removal may be too high.     

Coevolution of an avian host and its parasitic cuckoo

Abstract We use a quantitative genetic model to examine the coevolution of host and cuckoo egg characters (termed "size" as a proxy for general appearance), host discrimination, and host and cuckoo population dynamics. A host decides whether to discard an egg using a comparison of the sizes of the eggs in her nest, which changes as host and cuckoo eggs evolve. Specifically, we assume that the probability that she discards the largest egg in her nest depends on how much larger it is than the second largest egg. This decision rule (i.e., the acceptable difference in egg sizes) also evolves, changing both the chance of successful rejection of a cuckoo egg in parasitized nests and the chance of mistaken rejection of a host egg in both parasitized and unparasitized nests. We find a stable equilibrium for coexistence of the host and cuckoo where there is cuckoo egg mimicry, evolutionary displacement of the host egg away from the cuckoo egg phenotype, and host discrimination against unusual eggs. Both host discrimination and host egg displacement are fairly weak at the equilibrium. Cuckoo egg mimicry, although imperfect, usually evolves more extensively and quickly than the responses of the host. Our model provides evidence for both the evolutionary equilibrium and evolutionary lag hypotheses of host acceptance of parasitic eggs.     

Brood parasitism selects for no defence in a cuckoo host

In coevolutionary arms races, like between cuckoos and their hosts, it is easy to understand why the host is under selection favouring anti-parasitism behaviour, such as egg rejection, which can lead to parasites evolving remarkable adaptations to ‘trick’ their host, such as mimetic eggs. But what about cases where the cuckoo egg is not mimetic and where the host does not act against it? Classically, such apparently non-adaptive behaviour is put down to evolutionary lag: given enough time, egg mimicry and parasite avoidance strategies will evolve. An alternative is that absence of egg mimicry and of anti-parasite behaviour is stable. Such stability is at first sight highly paradoxical. I show, using both field and experimental data to parametrize a simulation model, that the absence of defence behaviour by Cape bulbuls (Pycnonotus capensis) against parasitic eggs of the Jacobin cuckoo (Clamator jacobinus) is optimal behaviour. The cuckoo has evolved massive eggs (double the size of bulbul eggs) with thick shells, making it very hard or impossible for the host to eject the cuckoo egg. The host could still avoid brood parasitism by nest desertion. However, higher predation and parasitism risks later in the season makes desertion more costly than accepting the cuckoo egg, a strategy aided by the fact that many cuckoo eggs are incorrectly timed, so do not hatch in time and hence do not reduce host fitness to zero. Selection will therefore prevent the continuation of any coevolutionary arms race. Non-mimetic eggs and absence of defence strategies against cuckoo eggs will be the stable, if at first sight paradoxical, result.     

Effect of nest and nest site characteristics on the risk of cuckoo Cuculus canorus parasitism in the great reed warbler Acrocephalus arundinaceus

The risk of cuckoo Cuculus canorus parasitism for great reed warbler Acrocephalus arundinaceus nests was evaluated in respect to nest and nest site structure in the reedbeds along canals in central Hungary, In total 90 nests were analysed, from which 31 remained unparasitised. 37 were single parasitised and 22 multiple parasitised. Multiple discriminant analysis, based on eight structural variables, separated well the unparasitised clutches from the cases of single and multiple parasitism, but the latter two categories also showed a weaker separation. Nests close to cuckoo vantage points are most vulnerable to parasitism. The variable "distance from nest to closest available cuckoo perch site" (tree or electric wire) played the most important role in the risk of parasitism. It showed highly significant differences among the groups: lowest distances were found in the case of multiple parasitism, and the highest distances were measured when nests remained unparasitised. Additionally, "nest visibility" also showed significant differences among the three groups: a more visible nest is more likely to be parasitised. Risk of parasitism affects on the host on two levels: 1) female cuckoos search for nest-building hosts from a perch site, but 2) when they are in the act of parasitism, they can find more visible nests in the reed. Besides the robust effects of the variables "distance to cuckoo perch site" and "nest visibility", the variable "distance from nest to open water" may reduce, but "nest volume" increases the risk of multiple parasitism. Differences between cases of single and multiple parasitism are weak, mainly affected by chance. We explain it by the high abundance of the cuckoo, the parasitism rate was 65.6% (59/90), Cuckoos parasitised almost all of the available nests in the close vicinity of potential perch sites. There was no evidence that great reed warblers nested closer to each other when risk of parasitism was high.     

Experimental Manipulation of Intraclutch Variation in the Great Reed Warbler Shows No Effect on Rejection of Parasitic Eggs

In the continuing arms race between hosts and brood parasites, hosts are expected to reduce variation in the appearance of their own eggs within clutches, as it facilitates recognition of parasitic eggs. At the same time, by increasing interclutch variation, hosts should make it more difficult for parasites to evolve perfectly mimetic eggs. In this study, we experimentally manipulated intraclutch variation in the great reed warbler, Acrocephalus arundinaceus, in Hungary, where this species is heavily (c. 64%) parasitized by the common cuckoo, Cuculus canorus. We placed artificial cuckoo eggs, which appeared moderately mimetic to humans, in two groups of nests; in one group we increased variability of egg appearance within clutches by exchanging host eggs among nests. These clutches showed a significantly higher intraclutch variability than natural clutches, which we used as a control group. Our results indicate that it has no effect on rejection behaviour in this species, neither when variation was increased experimentally, nor within the natural range of variation displayed by our population. We suggest that when parasitism is high, selection for reduced intraclutch variation may be less important than frequency‐dependent selection for increased variation between individuals within a host population.     

Responses of Black-headed Gulls Larus ridibundus to conspecific brood parasitism

Conspecific brood parasitism in birds occurs when a female inserts her egg into the clutch of her own species. If successful, i.e. the parasitic egg is accepted by the host, then the host female or pair rears the offspring of the parasite. In the present study, we studied natural conspecific brood parasitism in Black-headed Gulls (Larus ridibundus), and conducted series of the experiments with mimetic (conspecific) and non-mimetic (conspecific painted light blue) eggs to explore responses of the tested pairs towards these alien eggs. The natural parasitism rate was 10% and the probability of being parasitized significantly increased with nest density. Experimentally parasitized pairs rejected both types of experimental eggs at a similar rate: 14.3 % for mimetic and 25.5% for non-mimetic within 2 days. Non-mimetic eggs were more selectively rejected than mimetic eggs. The relationships between the probability of egg rejection (dependent variable) and predictor (independent) variables were examined by fitting generalized linear models. Contrast and intraclutch variation in ground color and spotting pattern and the volume of the egg had no significant effect on rejection behavior in either non-mimetic or mimetic eggs. However, nest density significantly positively affected rejection behavior of the Black-headed Gulls in both non-mimetic and mimetic treatments.