Attractive blue‐green egg coloration and cuckoo−host coevolution

Recent evidence suggests that blue‐green coloration of bird eggshells may be related to female and/or egg phenotypic quality, and that such colour may affect parental effort and therefore the nutritional environment of developing nestlings. Here we suggest that these relationships and the signal function of eggshell coloration would affect the outcome of coevolution between avian brood parasites and their hosts in at least three different non‐exclusive evolutionary pathways. First, by laying blue‐green coloured eggs, cuckoo females may exploit possible sensory biases of their hosts, constraining the evolution of parasitic egg recognition, and thus avoid rejection. Second, because of the relatively high costs of laying blue eggs, cuckoo females may be limited in their ability to mimic costly blue‐green eggs of their hosts because cuckoo females lay many more eggs than their hosts. Furthermore, costs associated with foreign egg recognition errors would be relatively higher for hosts laying blue eggs. Third, cuckoos may use coloration of host eggs for selecting individuals or specific hosts of appropriate phenotypic quality (i.e. parental abilities). We here explored some predictions emerging from the above scenarios and found partial support for two of them by studying egg coloration of European cuckoos (Cuculus canorus) and that of their 25 main hosts, as well as parasitism and rejection rate of hosts. Cuckoo hosts parasitized with more blue, green, and ultraviolet cuckoo eggs, or those laying more blue‐green eggs, were more prone to accept experimental parasitism with artificial cuckoo eggs. In addition, coloration of cuckoo eggs is more variable when parasitizing hosts laying bluer‐greener eggs, even after controlling for the effect of host egg coloration (i.e. degree of egg matching). Globally, our results are consistent with the proposed hypothesis that host egg traits that are related to phenotypic quality of hosts, such as egg coloration, may have important implications for the coevolutionary interaction between hosts and brood parasites. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 154–168.     

Function of egg punctures by Shiny Cowbirds in parasitized and nonparasitized Creamy‐bellied Thrush nests

ABSTRACT Avian brood parasites usually remove or puncture host eggs. Several hypotheses have been proposed to explain the function of these behaviors. Removing or puncturing host eggs may enhance the efficiency of incubation of cowbird eggs (incubation‐efficiency hypothesis) or reduce competition for food between cowbird and host chicks in parasitized nests (competition‐reduction hypothesis) and, in nonparasitized nests, may force hosts to renest and provide cowbirds with new opportunities for parasitism when nests are too advanced to be parasitized (nest‐predation hypothesis). Puncturing eggs may also allow cowbirds to assess the development of host eggs and use this information to decide whether to parasitize a nest (test‐incubation hypothesis). From 1999 to 2002, we tested these hypotheses using a population of Creamy‐bellied Thrushes (Turdus amaurochalinus) in Argentina that was heavily parasitized by Shiny Cowbirds (Molothrus bonariensis). We found that 56 of 94 Creamy‐bellied Thrush nests (60%) found during nest building or egg laying were parasitized by Shiny Cowbirds, and the mean number of cowbird eggs per parasitized nest was 1.6 ± 0.1 (N= 54 nests). At least one thrush egg was punctured in 71% (40/56) of parasitized nests, and 42% (16/38) of nonparasitized nests. We found that cowbird hatching success did not differ among nests where zero, one, or two thrush eggs were punctured and that the proportion of egg punctures associated with parasitism decreased as incubation progressed. Thus, our results do not support the incubation‐efficiency, nest‐predation, or test‐incubation hypotheses. However, the survival of cowbird chicks in our study was negatively associated with the number of thrush chicks. Thus, our results support the competition‐reduction hypothesis, with Shiny Cowbirds reducing competition between their young and host chicks by puncturing host eggs in parasitized nests.     

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.     

Egg morphology fails to identify nests parasitized by conspecifics in common pochard: a test based on protein fingerprinting and including female relatedness

Conspecific brood parasites lay eggs in nests of other females of the same species. A variety of methods have been developed and used to detect conspecific brood parasitism (CBP). Traditional methods may be inaccurate in detecting CBP and in revealing its true frequency. On the other hand more accurate molecular methods are expensive and time consuming. Eadie developed a method for revealing CBP based on differences in egg morphology. That method is based on Euclidean distances calculated for pairs of eggs within a clutch using standardized egg measurements (length, width and weight). We tested the applicability of this method in the common pochard Aythya ferina using nests that were identified as parasitized (39 nests) or non‐parasitized (16 nests) based on protein fingerprinting of eggs. We also analyzed whether we can distinguish between parasitic and host eggs in the nest. We found that variation in MED can be explained by parasitism but there was a huge overlap in MED between parasitized and non‐parasitized nests. MED also increased with clutch size. Using discriminant function analysis (DFA) we found that only 76.4% of nests were correctly assigned as parasitized or non‐parasitized and only 68.3% of eggs as parasitic or host eggs. Moreover we found that MED in parasitized nests increased with relatedness of the females that laid eggs in the nest. This finding was supported by positive correlation between MED and estimated relatedness in female–female pairs. Although variation in egg morphology is associated with CBP, it does not provide a reliable clue for distinguishing parasitized nests from non‐parasitized nests in common pochard.     

Evidence of selection for egg crypsis in conspicuous nests

ABSTRACT The value of egg coloration as crypsis, once accepted as a general principle, has recently been questioned because most experiments have failed to show that egg coloration deters predation. The nest‐crypsis hypothesis postulates that, among species that build conspicuous nests, selection for egg crypsis is relaxed or absent because visually searching predators detect nests prior to eggs. I tested the nest‐crypsis hypothesis using the large, relatively conspicuous nests of American Robins (Turdus migratorius), and eggs that differed markedly in color that were collected from the nests of Red‐winged Blackbirds (Agelaius phoeniceus), Brewer's Blackbirds (Euphagus cyanocephalus), and Yellow‐headed Blackbirds (Xanthocephalus xanthocephalus). Each nest (N= 22) received a clutch of each species during three sequential predation trials that were 16 d in duration. The order of clutch presentation was randomized for each nest. Survival trends for Brewer's and Yellow‐headed Blackbirds were similar, and higher than those for clutches of Red‐winged Blackbirds. By the end of trials, overall survival of the three clutch types was roughly equivalent. However, clutches of Red‐winged Blackbird eggs, the most conspicuous egg type to the human eye, were discovered sooner by predators. Because the experimental design controlled for effects of nest crypsis, nest location, and nest size, this difference in egg survival can be attributed to differences in egg pigmentation. Thus, my results support a role for egg coloration as camouflage in conspicuous nests.     

Common Cuckoos Cuculus canorus change their nest‐searching strategy according to the number of available host nests

In recent decades, numerous studies have examined factors affecting risk of host nest parasitism in well‐known avian host–parasite systems; however, little attention has been paid to the role of host nest availability. In accordance with other studies, we found that nest visibility, reed density and timing of breeding predicted brood parasitism of Great Reed Warblers Acrocephalus arundinaceus by the Common Cuckoo Cuculus canorus. More interestingly, hosts had a greater chance of escaping brood parasitism if nesting was synchronized. Cuckoo nest searching was governed primarily by nest visibility at high host‐nest density. However, even well‐concealed nests were likely to be parasitized during periods when just a few hosts were laying eggs, suggesting that Cuckoos adjust their nest‐searching strategy in relation to the availability of host nests. Our results demonstrate that host vulnerability to brood parasitism varies temporally and that Cuckoo females are able to optimize their nest‐searching strategy. Moreover, our study indicated that Cuckoos always manage to find at least some nests to parasitize. Thus, in this case, the co‐evolutionary arms race should take place mainly in the form of parasitic egg rejection rather than via frontline pre‐parasitism defence.     

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.     

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.     

Temporal patterns of host availability, brown-headed cowbird brood parasitism, and parasite egg mass

I studied relationships between temporal patterns of host availability, brood parasitism, and egg mass for the parasitic brown-headed cowbird (Molothrus ater). At a study site consisting largely of edge habitat in north-eastern Illinois, I found 834 bird nests from 27 species. A total of 407 cowbird eggs and nestlings were found in these nests over three laying seasons. Nearly all (n= 379; 93%) were found in the nests of seven host species. For these species and all taken together, weekly nest availability generally decreased whereas parasitism frequency generally increased throughout the cowbird laying season, but the proportions of nests parasitized and the mean number of cowbird eggs in them did not. Additionally, no correlation was found between the proportions of nests parasitized and nest availability. Cowbird egg mass generally increased throughout the laying season, indicating that foraging conditions improved and that, early in the laying season, egg mass and quality may be less important than quantity. Consistently high weekly levels of parasitism indicate that cowbird reproduction was less limited by resources needed for egg production and more by the availability of suitable host nests. Fluctuating weekly host availabilities suggest that previously established, constant rates of cowbird egg laying would produce an excess of eggs during periods of low host availability. Further, the low frequency of parasitism (1%) of nests in stages too advanced for successful parasitism, and of abandoned nests, is consistent with the hypothesis that cowbirds' consistently high rate of egg production helps assure an egg is available when an appropriate nest is found. Frequently, nests were parasitized multiple times, raising the possibility that cowbirds were interfering with their own reproduction. A diverse host community increases the possibility that a decline of any one host species is unlikely to affect cowbird reproduction significantly. Received 11 July 1997 / Accepted: 31 March 1998     

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.     

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.     

Decoding dumping ducks

Conspecific brood parasitism, where females of the same species lay eggs in each other's nests, is common in waterfowl, and is usually considered costly to host females, which are stuck looking after eggs and chicks that are not their own. However, since female waterfowl often exhibit an unusual propensity to nest near where they were born, there has been some uncertainty over whether, in ducks and geese, laying in nests of conspecifics really is parasitism. Do parasitic and host females tend to be related? And is parasitism actually a form of cooperation in disguise? In a population in Hudson Bay, Andersson & Waldeck (this issue) found that ‘parasitic’ eggs in nests of the common eider, Somateria mollissima sedentaria, are more closely related to host eggs than expected by chance. In fact, host and ‘donor’ eggs are more closely related than are females breeding at neighbouring nests. The Hudson Bay population of common eiders is unusual, because unlike in more benign climates, females do not tend to breed near their natal nest. Spatial proximity alone cannot account for the high relatedness between host eggs and ‘dumped’ or donor eggs. Instead, the high relatedness values are probably the result of active recognition, where females favour kin, either when dumping or accepting eggs. These new data, along with evidence indicating that the donor lays the first egg in the nest nearly half the time, suggest that what appears to be parasitism in common eiders may be a form of kin‐based cooperation.     

Timing of Spawning and Host-nest Choice for Brood Parasitism by the Japanese Minnow,Pungtungia herzi,on the Japanese Aucha Perch,Siniperca kawamebari

A field study of the breeding ecology of the Japanese aucha perch, Siniperca kawamebari, and brood parasitism by the Japanese minnow, Pungtungia herzi, on nests of the perch was carried out from 1989 to 1991. Observations of perch nests under natural conditions in 1990 showed that brood parasitism by the minnow was concentrated on host nests in which nest owners had just begun their nesting cycle. When spawned in a perch nest with recently spawned perch eggs, parasite eggs always hatched earlier than host eggs. An experiment with imitation perch eggs in 1991 confirmed that changing colour of host eggs was the cue for the parasites to distinguish between different developmental stages of host eggs. Parasite eggs rapidly disappeared without guarding by a host male (Baba et al. 1990). This loss was caused by predation by fishes. Parasite fry left the nest immediately after hatching, so parasite eggs spawned in a host nest in an early stage should be well guarded until they hatch. In the field, minnows deposited their eggs in perch nests which had larger numbers of newly spawned perch eggs. Since the perch males always deserted their nests when their own eggs disappeared, the parasite's choice of host nests with larger numbers of host eggs may ensure survival of the parasite eggs. The timing of egg deposition and choice of host nest by the minnow appear to be adaptive in terms of brood parasitism on nests of the perch.     

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.     

Blue–Green Eggs in Pied Flycatchers: An Experimental Demonstration that a Supernormal Stimulus Elicits Improved Nestling Condition

It has been proposed that blue–green egg colouration in certain avian species has evolved as a signal of female and egg quality to males, affecting their investment in the brood according to the differential allocation scenario. A recent experiment has successfully manipulated male investment by placing dummy eggs simulating the extremes in the blue–green natural reflectance. Here we have substituted one egg in certain pied flycatcher clutches with a single deep‐blue (blue) or light‐blue (pale) dummy egg and used other clutches as controls. Blue dummy eggs reflected more in the blue–green part of the spectrum than natural and pale dummy eggs, which were similar in reflectance. Nestlings in nests with one blue dummy egg during laying and incubation attained a significantly higher mass and condition than those in nests with one pale dummy egg or in control broods, when controlling for phenology, brood size and blue–green chroma of the clutch. Only a shift in parental investment can have induced this highly significant effect. Preference for deeply coloured clutches associated with high‐quality females and their maternal effects may explain stimulation by a single supernormal stimulus emanating from multiple objects like clutches.     

Breeding Season Aggression of Female and Male Eastern Bluebirds (Sialia sialis) to Models of Potential Conspecific and Interspecific Egg Dumpers

Naturally occurring aggression between female eastern bluebirds (Sialia sialis) is dramatic, resulting in severe injuries and even death. Furthermore, aggression among bluebirds is usually sex specific: males attack males, females attack females. We hypothesized that the primary function of female-female aggression is to guard against the threat of intraspecific egg dumping and that, in this context, same sex aggression is related to the possibility of advantages for males of parasitism (egg dumping) of their nests. Our hypotheses to explain variation in naturally occurring aggression predict temporal variation in aggressive tendency within nest cycles and between the sexes depending on asymmetries in threats to the residents. We report the results of experimental trials in the field designed to determine temporal variation in the aggressive tendencies of resident females to models of intruder females of two species, eastern bluebirds and brown-headed cowbirds (Molothrus ater). Both species dump eggs in the nests of bluebirds. Female aggression to eastern bluebird models is greatest during early stages of nest cycles; the patterns are most consistent with protection against egg dumping and protection of nest sites from usurpation. Male residents seldom attack female eastern bluebird models, but often attack models of female brown-headed cowbirds, a result inconsistent with the hypothesis that patterns of differential parental care control aggression of female and male residents.     

Experimental Shifts in Intraclutch Egg Color Variation Do Not Affect Egg Rejection in a Host of a Non-Egg-Mimetic Avian Brood Parasite

Avian brood parasites lay their eggs in the nests of other birds, and impose the costs associated with rearing parasitic young onto these hosts. Many hosts of brood parasites defend against parasitism by removing foreign eggs from the nest. In systems where parasitic eggs mimic host eggs in coloration and patterning, extensive intraclutch variation in egg appearances may impair the host’s ability to recognize and reject parasitic eggs, but experimental investigation of this effect has produced conflicting results. The cognitive mechanism by which hosts recognize parasitic eggs may vary across brood parasite hosts, and this may explain variation in experimental outcome across studies investigating egg rejection in hosts of egg-mimicking brood parasites. In contrast, for hosts of non-egg-mimetic parasites, intraclutch egg color variation is not predicted to co-vary with foreign egg rejection, irrespective of cognitive mechanism. Here we tested for effects of intraclutch egg color variation in a host of nonmimetic brood parasite by manipulating egg color in American robins (Turdus migratorius), hosts of brown-headed cowbirds (Molothrus ater). We recorded robins’ behavioral responses to simulated cowbird parasitism in nests where color variation was artificially enhanced or reduced. We also quantified egg color variation within and between unmanipulated robin clutches as perceived by robins themselves using spectrophotometric measures and avian visual modeling. In unmanipulated nests, egg color varied more between than within robin clutches. As predicted, however, manipulation of color variation did not affect rejection rates. Overall, our results best support the scenario wherein egg rejection is the outcome of selective pressure by a nonmimetic brood parasite, because robins are efficient rejecters of foreign eggs, irrespective of the color variation within their own clutch.     

Nest, but Not Egg, Fidelity in a Territorial Salamander

Egg recognition and subsequent egg brooding are costly forms of parental investment in many species of vertebrates. Life history factors, such as coloniality or risk of brood parasitism, may constrain egg recognition in vertebrates. Female red-backed salamanders ( Plethodon cinereus ) from my study site are territorial and do not share nest sites with other females. They are terrestrial and neither they nor their eggs are likely to be displaced by environmental factors such as flooding. I experimentally tested, in the laboratory, the hypothesis that female red-backed salamanders can discriminate between their own eggs and the eggs of unfamiliar females. Each female was allowed to move about a test chamber containing two clutches of eggs, one clutch with which it was found in the forest and one that had been found with a distant female. Most females remained with one clutch of eggs, which they brooded during the entire observation period. However, they did not significantly prefer to brood their own eggs over the eggs of another female. In a corollary field experiment, I tested whether brooding females that were displaced 1 m from their nest sites would return to their territories and commence brooding behaviour within 3 d. All 10 displaced females returned to their own nest within this time period and were found brooding their eggs. Because female red-backed salamanders at my study site do not tend to share nest sites with other females and because their eggs remain in stationary nests, selection may not have favoured egg recognition. However, the results suggest that female salamanders indirectly recognize their own eggs by actively recognizing their territorial nest sites.     

Brood Parasitism and Nest Takeover in Common Eiders

Conspecific brood parasitism (CBP) is an alternative breeding tactic that occurs in many brood‐tending animals and can have important fitness effects for both host and parasite. We use protein fingerprinting of egg albumen to distinguish the eggs from different females and to estimate the frequency, pattern and tactics of CBP and other forms of mixed maternity in a Hudson Bay population of common eiders (Somateria mollissima sedentaria). Mixed clutches, containing eggs from more than one female, occurred in 31% of the 86 nests studied that progressed to clutch completion. Other females than the host laid 8% of the eggs. In 11 (41%) of the mixed clutches another female laid before the host started laying, corroborating the hypothesis that takeover of nests started by other females accounts for many of the mixed clutches in this population. Our results also indicate that traditional non‐molecular methods of identifying foreign eggs may considerably underestimate the frequency of mixed clutches.     

Influence of Shape on Egg Discrimination in American Robins and Gray Catbirds

The eggs of some obligate brood parasites are more spherical than the eggs of their non‐parasitic relatives and hosts, which contributes to the increased strength of their shells. We examined whether egg shape, including the more spherical shape of brown‐headed cowbird eggs (Molothrus ater), influenced egg discrimination in American robins (Turdus migratorius) and gray catbirds (Dumetella carolinensis). We added a series of artificial objects to robin and catbird nests that varied in shape from a control host egg, a rounded, cowbird‐like egg, to odd‐shaped objects. Real cowbird eggs were significantly more spherical than catbird and robin eggs, which confirmed a potential cue for egg recognition. Object shape significantly influenced the probability of rejection and time to rejection in both robins and catbirds. However, rounded eggs and spheres were rejected infrequently and at frequencies similar to control eggs. Therefore, the shape of a brood parasite's egg does not appear to influence egg discrimination in these two rejecters. Robins and catbirds rejected significantly more odd‐shaped objects than egg‐shaped objects and odd‐shaped objects were rejected significantly sooner than egg‐shaped objects. The rejection of odd‐shaped objects likely represents an expression of nest‐sanitation behaviour where debris is removed from the nest. By comparison with other studies of accepters of cowbird eggs, robins and catbirds appear to reject higher proportions of odd‐shaped objects, which suggests they may have more refined abilities to discriminate against foreign objects in their nests.