Foreign egg retention by avian hosts in repeated brood parasitism: why do rejecters accept?

Great reed warblers (Acrocephalus arundinaceus) are frequently parasitized by egg-mimetic common cuckoos (Cuculus canorus) in Hungary, and these hosts reject about a third of parasitic eggs. The timing of parasitism is important, in that the probability of rejection decreases with advancing breeding stages in this host. Also, egg rejection is more common when a clutch is parasitized by a single foreign egg, compared to parasitism by multiple eggs. We repeatedly parasitized great reed warbler clutches with moderately mimetic foreign eggs, either with (1) one foreign egg (single parasitism) and, after 3 days, by all foreign eggs (multiple parasitism), or (2) all foreign eggs and, 3 days later, by only one foreign egg. Hosts ejected 26–53 % of the experimental parasitic eggs in the first stage of the repeated parasitism, but almost all eggs were accepted in the second stage, irrespective of whether the clutch was singly or multiply parasitized. Video-taping of the behavioural responses of hosts to experimental parasitism revealed no evidence for sensory constraints on foreign-egg recognition, because hosts recognized and pecked the parasitic eggs as frequently in the second stage of repeated parasitism, as they did in the first stage. We suggest that the relative timing of parasitism (laying vs. incubation stage), rather than learning to accept earlier-laid foreign eggs, results in higher acceptance rates of cuckoo eggs in repeated parasitism, because there is decreasing natural cuckoo parasitism on this host species and, hence, less need for antiparasitic defences, with the advancing stages of breeding.     

Electrophoretic variants of egg white transferrin indicate a low rate of intraspecific brood parasitism in colonial cliff swallows in the Sierra Nevada,California

Summary The frequency of intraspecific brood parasitism in two colonies of cliff swallows (Hirundo pyrrhonota) nesting in the eastern Sierra Nevada in California was assessed through an electrophoretic analysis of egg white (albumin) proteins. Albumin proteins are maternally derived and are presumed to directly reflect maternal genotype. Thus, a comparison of protein banding patterns produced by eggs collected from a single clutch allows any egg laid by a female other than the putative mother to be detected. Eggs were collected from 13 cliff swallow nests in 1984 and 41 nests in 1987, a total of 54 nests. Of the clutches collected in 1984, one showed evidence of intraspecific brood parasitism, giving a nest parasitism rate of 7.6%. In 1987, one of 41 nests contained a parasitic egg, for a parasitism rate of 2.4%. Over both years combined the rate of intraspecific brood parasitism was 2 of 54 nests, or 3.7%. These parasitism rates are much lower than those estimated for Nebraskan cliff swallows (22–43%) and 2nd-year purple martins (36%). Possible explanations for the discrepancy between parasitism rates in Sierran cliff swallows and other groups of swallows are discussed.Correspondence to: A.P. Smyth     

Intraspecific nest parasitism in the swallow Hirundo rustica: the importance of neighbors

Summary Intraspecific nest parasitism in the swallow Hirundo rustica involves several parasite and anti-parasite tactics. Neighboring swallows breed asynchronously, perhaps because neighboring nests with overlapping egg-laying periods have higher frequency of intraspecific nest parasitims than neighboring nests with no such overlap. Since nest guarding prevents nest parasitism, the effect of nest guarding was removed by putting up old swallow nests. Such experimental nests were parasitized by swallows more often when the nests contained eggs, than when empty, and when the nests were far away from, rather than close to, an active neighboring nest. Close experimental nests with eggs were, however, parasitized less frequently if asynchronous (non-overlapping egg-laying periods), rather than synchronous, to active neighboring nests. Aggressive anti-parasite behavior of nest owners was determined from responses towards female intruders. Responses were more often aggressive before and especially during egg laying than during incubation. Nest owners also behaved aggressively more often when the intruder approached close to as compared to at some distance from their nest. Focal pairs behaved aggressively more often towards closely approaching intruders during the incubation period than towards distantly approaching intruders during the egg-laying period. Swallows reduce the frequency of intraspecific nest parasitism by nesting asynchronously with and close to a neighboring nest. Aggression by neighbors may reduce the success of potential parasitic swallows.     

Adaptations of meadow pipits to parasitism by the common cuckoo

Summary The meadow pipit Anthus pratensis is one of the most frequent hosts in Europe parasitized by the common cuckoo Cuculus canorus. The cuckoo normally removes one or more of the host eggs and replaces them with one of its own. The aim of the present study, which was conducted in an upland area of Central Norway, was to test the following question: assuming the cuckoo has laid a mimetic egg (which is slightly larger than and slightly different in color from that of a meadow pipit egg), under what circumstances are the parent meadow pipits able to detect such parasitism? The reaction of the meadow pipits to artificial parasitism was tested. Plastic model cuckoo eggs were used that bore a striking resemblance to real cuckoo eggs found in other meadow pipit nests in the same area. In addition, in some experiments a stuffed cuckoo dummy was used. The meadow pipits tolerated the experimental procedures and remained in their nests when given an artificial cuckoo egg, with or without removal of one of the host's eggs. However, when a host egg was removed and replaced with an artificial cuckoo egg, and at the same time a stuffed cuckoo dummy was presente, 50% of the birds deserted their nests. The difference between this result and the results of the other experiments was statistically significant. When only the stuffed cuckoo was presented, without any egg manipulations, the meadow pipits reacted in the same way as in the egg experiments. We conclude that the meadow pipit is capable of detecting whether or not its nest has been parasitized, provided it has observed the cuckoo near the nest site. Furthermore, because of the results of our experiments, we reject the hypothesis that the cuckoo has evolved egg removal behavior in order to prevent the host from assessing an increase in egg numbers.     

Does the great spotted cuckoo choose magpie hosts according to their parenting ability?

When brood parasites are about to lay an egg, they have to decide which nest to parasitize. The best nest in which to lay will depend on the parenting ability of the host. We have studied selection of magpie (Pica pica) hosts by great spotted cuckoos (Clamator glandarius). Great spotted cuckoos preferentially parasitize large host nests. Nest volume in magpies is a good indicator of territory quality, since there is a negative relationship between magpie nest size and breeding date, and timing of breeding in magpies is known to be positively related to territory quality. Moreover, magpies occupying high-quality territories have high breeding success. Therefore, nest size is positively related to the quality of magpies. Parasitized magpie nests were of greater volume than the nearest neighbouring nest not parasitized by the great spotted cuckoo. In order to test whether the great spotted cuckoos might select high-quality magpie hosts, we manipulated pairs of parasitized and non-parasitized nests with identical laying dates and habitats, introducing into each of the nests the same number of parasitic and non-parasitic eggs. The number of fledglings reared (magpie plus great spotted cuckoo chicks) in naturally parasitized nests was higher than in experimentally parasitized nests. Thus, the probability of survival of the parasite chicks increased if cuckoo eggs were laid in the nests of high-quality hosts originally chosen by the parasite.     

Costs of brood parasitism and the lack of defenses on the yellow-winged blackbird - shiny cowbird system

The shiny cowbird (Molothrus bonariensis) is a generalist brood parasite that lays either white-immaculate or spotted egg morphs in eastern Argentina and Uruguay. Some hosts accept both morphs, others accept spotted eggs and reject the white morph, but no host has been found to accept white eggs and reject spotted ones. It has been suggested that the yellow-winged blackbird (Agelaius thilius) may be that type of host. The finding of a white acceptor-spotted rejector species would help to explain the occurrence and maintenance of the parasite egg polymorphism. We studied the incidence of shiny cowbird parasitism on this host, its costs for their reproductive success and the presence of antiparasitic defenses in the yellow-winged blackbird - shiny cowbird system. The parasite affected the reproductive success of the host in two ways. Cowbirds punctured host eggs causing a reduction in clutch size, and yellow-winged blackbirds deserted their nests whenever they suffered high egg loss. In addition, parasitized nests suffered higher predation during the nestling stage, but not during egg stages, indicating that the difference found was related to the presence of the cowbird chick, and not to higher exposure of parasitized nests to both parasites␣and predators. Despite the costs imposed by the parasite, yellow-winged blackbirds have not evolved antiparasitic defenses. This host did not reject any egg morph of the shiny cowbird nor desert parasitized nests unless it had suffered high egg loss. Current explanations for the host lack of defenses, the “time lag” and the “equilibrium” hypothesis, are discussed. Received: 29 August 1997 / Accepted after revision: 10 January 1998     

Mechanism of egg recognition in defenses against conspecific brood parasitism: American coots (<Emphasis Type="Italic">Fulica americana</Emphasis>) know their own eggs

Hosts of avian brood parasites use a variety of defenses based on egg recognition to reduce the costs of parasitism; the most important of which is rejecting the parasitic eggs. Two basic recognition mechanisms are possible: “true recognition”, whereby hosts recognize their own eggs irrespective of their relative frequency in the clutch, and minority recognition (or “recognition by discordancy”), whereby hosts respond to the minority egg type. The mechanism of recognition has been experimentally studied in a handful of species parasitized by interspecific brood parasites, but the mechanism used in defenses against conspecific brood parasitism is unknown. I experimentally determined the mechanism of egg recognition in American coots (Fulica americana), a species with high levels of conspecific brood parasitism, egg recognition, and rejection. I swapped eggs between pairs of nests to alter frequencies of host and “parasite” eggs and then used two criteria for recognition: egg rejection and nonrandom incubation positions in the clutch. Eight of 12 nests (66%) given equal frequencies of host and parasite eggs showed evidence of true recognition. In contrast, only one of eight (12.5%) nests where host eggs were in the minority showed evidence of recognition by discordancy. The nonrandom incubation positions of parasitic eggs indicates that birds sometimes recognize parasitic eggs without rejecting them and provides a means of assessing recognition on a per nest basis in species with large clutches. Adaptive recognition without rejection may also be an important evolutionary stepping stone to the evolution of egg rejection in some taxa.     

Why do Horsfield’s bronze-cuckoo <Emphasis Type="Italic">Chalcites basalis</Emphasis> eggs mimic those of their hosts?

The Horsfield’s bronze-cuckoo (Chalcites basalis) egg closely matches the appearance of its host fairy-wren (Malurus spp.) eggs. Mimicry of host eggs by cuckoos is usually attributed to coevolution between cuckoos and hosts, with host discrimination against odd-looking eggs selecting for ever better mimicry by cuckoos. However, this process cannot explain Horsfield’s bronze-cuckoo egg mimicry because fairy-wren hosts rarely reject odd-looking eggs from their nest. An alternative hypothesis is that cuckoos have evolved egg mimicry to disguise their eggs from other cuckoos. Female cuckoos remove one egg from the nest during parasitism and would potentially benefit by selectively removing any cuckoo egg that has already been laid in the nest because otherwise, their egg will be evicted by the first nestling cuckoo along with the host clutch. We used painted, non-mimetic eggs to test whether cuckoos selectively remove odd-looking eggs during parasitism. We found that they were no more likely to remove a non-mimetic egg from a superb fairy-wren Malurus cyaneus clutch than would be expected by chance. Thus, our study does not support the cuckoo egg replacement hypothesis to explain mimicry of host eggs by cuckoos.     

Egg removal and intraspecific brood parasitism in the European starling (Sturnus vulgaris)

Summary From 1983 to 1986 we monitored 284 European starling (Sturnus vulgaris) nests in New Jersey for evidence of intraspecific brood parasitism and egg removal during the laying period. Egg removal occurred significantly more often at nests where intraspecific brood parasitism was detected (12 of 35 nests, 34%) than at unparasitized nests (23 of 249 nests, 9%). Brood parasitism (92% of parasitized nests) and egg removal (74% of nests with egg removal) were most common at nests where egg laying began in April of each year (i.e., early nests). Egg removal occurred at 26 (19%) and brood parasitism at 32 (23%) of 138 early nests. Both brood parasitism and egg removal were concentrated during the first four days in the laying period when brood parasitism is most likely to be successful and when host nests are most vulnerable to parasitism (Romagnano 1987). Both parasitism and removal usually involved a single egg at each nest. We detected brood parasitism and egg removal on the same day at five of 12 nests (42%) where both were observed. Because starlings do not remove foreign eggs from their nests once they begin laying (Stouffer et al. 1987) we hypothesize that parasite females sometimes removed host eggs while parasitizing nests.     

Spectral tuning and perceptual differences do not explain the rejection of brood parasitic eggs by American robins (Turdus migratorius)

By laying their eggs in the nests of other birds, avian brood parasites impose the cost of rearing young upon their hosts. The recognition and rejection of foreign eggs are primary host defenses against costly brood parasitism. Hosts of parasitic brown-headed cowbirds (Molothrus ater) challenge coevolutionary theory because most cowbird hosts accept parasitic eggs despite their drastically different appearance from the hosts’ own eggs. American robins (Turdus migratorius) are one of only 10 % of the over 200 potential cowbird host species to robustly reject parasitic eggs, but the mechanisms driving the sensory bases of foreign egg rejection remain elusive. Our research combined avian visual perceptual modeling and behavioral experimentation to investigate chromatic cues eliciting parasitic egg rejection in American robins. We assessed the effects of perceivable background color differences between real host and model parasite eggs, across all four avian photoreceptors, on rates of rejection of model eggs spanning in color across the entire avian spectral sensitivity range, and including immaculate model eggs matching the natural colors of robin and cowbird eggs. The results suggest that egg rejection in robins is driven by the overall perceivable difference in color between own and artificial eggs, and input from all four single-cone avian photoreceptors affects the rejection decision. The results, however, also reveal that when viewed by the avian eye, natural cowbird eggs appear more similar in background color to robin eggs than predicted by the high rejection rate of these parasitic eggs. This suggests that robins respond specifically to parasitism by cowbirds, despite an apparent lack of sensory tuning toward the detection of the background color of cowbird eggs.     

Differential parasitism of least flycatchers and yellow warblers by the brown-headed cowbird

Summary Brown-headed cowbirds (Molothrus ater) parasitized yellow warblers (Dendroica petechia) six times more frequently than least flycatchers (Empidonax minimus) nesting in the same riparian habitat in southern Manitoba. This difference was surprising because least flycatchers were higher quality hosts; flycatchers always accepted cowbird eggs while warblers did so on only about half the occasions. Both hosts were equally good foster parents, at least until fledging; thus, once an egg was accepted there was probably no further advantage in parasitizing one species over the other. The probability that a nest was parasitized decreased with increasing nest height, perhaps as a consequence of the cowbird's habit of searching for nests close to the ground. Since least flycatchers nested higher in the canopy than yellow warblers, a greater proportion of flycatcher nests probably were not detected by cowbirds. Nevertheless, nest height alone could not account fully for the lower incidence of parasitism on flycatchers. Clutch initiation in flycatchers peaked 6 days after warbler clutch initiations and 5 days after that of cowbirds. Despite the later peak in initiations, flycatcher nests were always available as potential hosts over the entire cowbird laying season and it was not until new clutch initiations by warblers declined in late summer that flycatchers were actually used as hosts. Because least flycatchers responded more aggressively than yellow warblers to a model female cowbird at the nest, we concluded that greater nest defense by flycatchers may have also reduced the rate of brood parasitism in this host. Together, our results suggest the large difference in the frequency of parasitism between these two hosts was primarily a product of nest location but that differences in host nest-defense and breeding season asynchrony may have contributed to preferential host selection.Offprint requests to: J.V. Briskie     

Experimental support for the use of egg uniformity in parasite egg discrimination by cuckoo hosts

Common cuckoo (Cuculus canorus) parasitism drastically reduces the reproductive success of their hosts and selects for host discrimination of cuckoo eggs. In a second stage of anti-parasite adaptation, once cuckoos can lay eggs that mimic those of their hosts, a high uniformity of host egg appearance within a clutch may favour cuckoo egg discrimination. Comparative evidence provides indirect support for this hypothesis although experimental support is currently lacking. Here, we studied the effect of experimentally decreased uniformity of host egg appearance on cuckoo egg discrimination by great reed warbler (Acrocephalus arundinaceus) hosts in a population in which long-term cuckoo parasitism has led to high levels of cuckoo–host egg mimesis. We manipulated host clutch uniformity by adding extra spots to fresh host eggs just after they were laid. Rejection of non-mimetic experimental eggs added to these nests was compared with those in control nests in which uniformity was not altered. Previously, by over-painting real spots in a control group of nests, we showed a negligible effect of our paints on hosts’ perception of their eggs. We show that for the great reed warbler, non-mimetic experimental eggs were relatively more tolerated in experimental nests, i.e. with lower uniformity (40%) than in control nests (5%). This is the first experimental study, to our knowledge, which demonstrates a reduced discrimination of foreign eggs as a consequence of an increase of egg phenotypes variation perception in a cuckoo host.     

House sparrows selectively eject parasitic conspecific eggs and incur very low rejection costs

Most host species of obligate interspecific brood parasites are under strong selection because such parasitism, e.g., that involving evictor nestmates, is highly costly. Egg rejection is one of the most efficient host defences against avian brood parasites. Many hosts have thus evolved egg-recognition ability and rejection behaviour. However, this defensive mechanism has not evolved in most species where only intraspecific brood parasitism occurs, probably because (1) the eggs of conspecific females are very similar in appearance, making egg rejection less likely to emerge, and (2) such parasitism is frequently less costly than interspecific parasitism. Using a captive population of house sparrows (Passer domesticus) with a low breeding density, we here provide new evidence showing that this species actually has a fine capacity to discriminate conspecific eggs and to eject them (44.2% of foreign eggs ejected) while incurring very low rejection costs (4.2% of own eggs ejected). This result contradicts those previously found in high-density house sparrow populations in which very high rejection costs and very high clutch desertion rates were reported, probably as a consequence of intraspecific competition and infanticide provoked by the high breeding density. The house sparrow has only rarely been reported as the host of an interspecific brood parasite, which implies that it is a newly described example of an altricial species in which egg ejection has evolved and is maintained in response to intraspecific brood parasitism.     

Shiny cowbirds share foster mothers but not true mothers in multiply parasitized mockingbird nests

Obligate brood parasitic birds, such as cowbirds, evade parental care duties by laying their eggs in the nests of other species. Cowbirds are assumed to avoid laying repeatedly in the same nest so as to prevent intrabrood competition between their offspring. However, because searching for host nests requires time and energy, laying more than one egg per nest might be favoured where hosts are large and can readily rear multiple parasites per brood. Such ‘repeat parasitism’ by females would have important consequences for parasite evolution because young parasites would then incur indirect fitness costs from behaving selfishly. We investigated shiny cowbird (Molothrus bonariensis) parasitism of a large host, the chalk-browed mockingbird (Mimus saturninus), in a population where over 70 % of the parasitized mockingbird nests receive multiple cowbird eggs. We assessed egg maternity directly, using cameras at nests to film the laying of individually-marked females. We also supplemented video data with evidence from egg morphology, after confirming that each female lays eggs of a consistent appearance. From 133 eggs laid, we found that less than 5 % were followed by the same female visiting the nest to lay again or to puncture eggs. Multiple eggs in mockingbird nests were instead the result of different females, with up to eight individuals parasitizing a single brood. Thus, while cowbird chicks regularly share mockingbird nests with conspecifics, these are unlikely to be their maternal siblings. Our results are consistent with shiny cowbird females following a one-egg-per-nest rule, even where hosts can rear multiple parasitic young.     

Tactics of parasitic American coots: host choice and the pattern of egg dispersion among host nests

Summary I examined the tactics adopted by a conspecific brood parasite, the American coot (Fulica americana), and the degree to which these tactics reflect sources of mortality for parasitic eggs. Only 8% of parasitic eggs produced independent offspring, compared to a 35% success rate for non-parasitic eggs, and most mortality was due to egg-rejection by hosts or the consequences of laying eggs too late in the host's nesting cycle. Parasites usually laid parasitically before initiating their own nests and usually parasitized immediate neighbours. Parasites did not remove host eggs before laying their own egg, and egg disappearance in general was not more common at parasitized nests. I found no evidence for non-random host choice, either on the basis of stage of the host's nesting cycle or the host's brood size. The absence of adaptive host choice is likely a consequence of the fact that, due to host limitation, only a small proportion of parasites had meaningful variation among potential hosts to choose from. The pattern of egg dispersion among host nests by individual parasites appears to be a compromise between constraints imposed by host limitation and the increased success obtained from spreading eggs among nests. Most females laying fewer than five parasitic eggs laid them in a single host nest while females laying five or more eggs normally parasitized two or more hosts. An examination of egg rejection and survival rates showed that parasites would maximize success by laying a single egg per host nest, and the pattern of laying several eggs per host nest is likely a consequence of host limitation. However, no egg that was the fifth laid, or later, parasitic egg in a host nest was ever successful and this probably explains why most females laying five or more eggs parasitized more than one host.     

Should the redstart Phoenicurus phoenicurus accept or reject cuckoo Cuculus canorus eggs?

Hole-nesting habits of redstarts Phoenicurus phoenicurus make laying difficult for parasitic cuckoo Cuculus canorus females and eviction of host eggs difficult for the cuckoo hatchling, causing fitness costs of cuckoo parasitism to be lower than those reported for open nesting hosts. Redstarts have recognition problems when confronted with real cuckoo eggs showing a perfect mimicry with their own eggs since they never eject when parasitized with perfect mimetic cuckoo eggs but instead desert the nest. Here we use a cost-benefit model to assess the effects of parasitism costs and the probability of being parasitized to estimate the reproductive success of redstarts when accepting or rejecting in the presence or absence of parasitism. Baseline data for model calculations come from this and a previous study on a cuckoo parasitized redstart population in Finland. When desertion implies a loss of 50%, we found that below a threshold value of 20% parasitism redstarts should accept cuckoo eggs since the costs of rejection exceed the benefits, whereas above this threshold they should reject. Interestingly, as the cost of desertion increases the threshold value, it should pay the redstart to reject increasingly at an exponential rate. Our field observations on natural parasitism and experiments with artificial cuckoo eggs confirmed the predictions from the model when hatching failures of the cuckoo were taken into account. Therefore, the low cost imposed by cuckoo parasitism in the system, and the presumably high cost of desertion as a response to parasitism favours acceptance over rejection for a wide range of parasitism pressures. This finding could explain the low rejection rate of real cuckoo eggs found in the redstart despite the presumably long history of a coevolutionary relationship with the cuckoo in Finland.     

Mechanisms of avian egg recognition: Which egg parameters elicit responses by rejecter species?

Summary Some species of North American passerines nearly always reject nonmimetic eggs placed in their nests and have apparently evolved this behavior in response to brood parasitism. Experiments presented here examined the specific egg parameters to which rejecter species respond, the relative tolerances rejecters show towards nonmimetic eggs and the degree to which rejection is limited to eggs of the brown-headed cowbird (Molothrus ater), the only parasitic bird widespread in North America.Relative to cowbird eggs, American robin (Turdus migratorius) eggs are larger, blue rather than white and immaculate rather than spotted. Experiments using 10 egg models at 137 nests showed that robins respond to each of these differences (Figs. 2 and 3) but do not usually reject an egg that deviates from their own by only one difference. Eggs that differ in ant two of the three parameters are usually rejected. This built-in tolerance reduces the likelihood that robins will reject their own eggs if these are atypical in size or coloration.Small egg size was the most important parameter eliciting rapid rejections (i.e. within 1 day), probably because differnces in size can be detected by both visual and tactile perception. By contrast, small egg size was the least important parameter determining whether eggs were eventually rejected (i.e. within 5 days, Tables 2 and 3). In terms of their eventual response, robins may be more sensitive to egg coloration than to size because the latter parameter is less reliable in distinguishing between robin and cowbird eggs.Experiments were also carried out at 37 nests of the gray catbird (Dumetella carolinensis), whose immaculate blue-green egg is only slightly larger than a cowbird egg. Catbirds are much more responsive to white ground color than to maculation (Table 4), perhaps because color is more reliable in distinguishing between catbird and cowbird eggs.Rejecter species exhibit degrees of tolerance towards foreign eggs that are proportional to the divergence between their eggs and those of the cowbird. Birds with eggs strongly divergent from cowbird eggs benefit from being relatively tolerant because they avoid rejecting their own eggs but still act against cowbird eggs. Species with cowbird-like eggs must be relatively intolerant to maximize the chances that cowbird eggs are rejected.Experiments show that rejection is not specific to cowbird eggs. Thus, birds have apparently responded evolutionarily to brood parasitism by developing recognition of their own eggs, rather than by developing recognition and rejection specific to parasitic eggs.     

Brood parasitism disproportionately increases nest provisioning and helper recruitment in a cooperatively breeding bird

Obligate avian brood parasites lay their eggs in nests of other species (hosts), which raise parasitic young. Parasitic nestlings are likely to influence host’s parental behaviours as they typically beg for food more vigorously than young host for a given hunger level. However, few studies have tested this idea, with conflicting results. These prior studies were largely limited to biparental hosts, but little is known about the effect of brood parasitism on parental behaviours in hosts that breed cooperatively. We followed a multimodel approach to examine the effect of brood parasitism on nest provisioning and helper recruitment in the baywing (Agelaioides badius), a cooperative breeder parasitised by screaming (Molothrus rufoaxillaris) and shiny (Molothrus bonariensis) cowbirds. Multimodel inference results indicated that feeding visits increased with nestling age, cooperative group size and number of cowbird nestlings in the brood. Brood size had little influence on feeding visits, which further suggests that baywings adjusted their provisioning effort in response to cowbird parasitism. In addition, nests parasitised artificially with shiny cowbird eggs or hatchlings recruited more helpers than unmanipulated nests having only host or screaming cowbird young. Our results provide novel evidence that brood parasitism and cooperative breeding interact in determining the levels of nest provisioning.     

Intraspecific brood parasitism as a conditional reproductive tactic in the treehopper<Emphasis Type="Italic"> Publilia concava</Emphasis>

In species exhibiting egg guarding as well as communal egg laying, females may adopt the strategy of laying eggs in the nests of conspecifics and leaving without providing care (termed intraspecific brood parasitism). This study is the first to describe such a behavior in the insect Publilia concava (Hemiptera: Membracidae) through field studies that followed 849 marked females across 1,828 brood associations. While brood parasitism increased the total number of eggs in a host brood, it did not reduce the overall hatching success of host broods. Solitary females exhibited a range of guarding durations while parasitic females rarely remained to guard eggs. Females exhibiting the parasitic tactic increased their lifetime number of clutches without decreasing the number of solitary clutches that they were able to initiate. Estimates of egg number for these individual broods revealed that females adopting the parasitic tactic (in addition to solitary breeding) had higher lifetime fecundity relative to females that did not parasitize. In females that exhibited both tactics (solitary and parasitic), the parasitic tactic yielded a higher rate of oviposition. The major component of oviposition rate was the time to find hosts and this time decreased with increasing host availability across 222 replicate groups. Females exhibited a shift toward the parasitic tactic when host broods were more abundant (i.e. in larger groups and later in the season). However, the time to find hosts increased as the frequency of the parasitic tactic increased, suggesting that this tactic may be maintained through negative frequency dependence. The results of this study suggest that brood parasitism may be the preferred tactic, as part of a conditional strategy, when hosts are readily available with solitary breeding being the preferred tactic when hosts are in short supply.     

Post-ejection nest-desertion of common cuckoo hosts: a second defense mechanism or avoiding reduced reproductive success?

Hosts of the common cuckoo (Cuculus canorus), an avian brood parasite, develop antiparasite defense mechanisms to increase their reproductive success. Ejection of the parasite egg and desertion of the parasitized nest are the most typical adaptations in response to brood parasitism, but nest desertion may also occur in response to partial clutch reduction, independently from parasitism. Some great reed warblers (Acrocephalus arundinaceus) showed both mechanisms in the same incidence of cuckoo parasitism: in 18% of successful ejections of the parasite eggs, they deserted their nests. We studied if such cases of post-ejection nest-desertion are caused by brood parasitism or reduced clutch value. We experimentally parasitized clutches consisting of five or three host eggs with two painted conspecific eggs to mimic parasitic eggs, as multiple parasitism is frequent in the area. Although hosts ejected these parasitic eggs in both clutch categories (100% and 67% for the larger and smaller inital clutch sizes, respectively), we found that after manipulation, post-ejection nest-desertion frequently occurred at small (3-egg) clutches (40%), but rarely at large (5-egg) clutches (17%). The same phenomenon also occurred when unparasitized 3-egg clutches were reduced by two eggs, but not when 5-egg clutches were reduced in the same way. A logistic regression model revealed that only initial clutch size affected nest desertion of parasitized nests in our experiments. Therefore, we conclude that post-ejection nest-desertion is not a second antiparasite mechanism, which might serve as a redundant antiparasite defense, but a reaction to typically small and further decreased clutch size.