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

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

Experiments and observations of prothonotary warblers indicate a lack of adaptive responses to brood parasitism

It has been suggested that prothonotary warblers, Protonotaria citrea, respond adaptively to brood parasitism by brown-headed cowbirds, Molothrus ater, even though they lack historical habitat and range overlap with cowbirds. I studied behaviours functioning as potential defences against brood parasitism in the prothonotary warbler, a cavity-nesting host species. Opening sizes preferred by prothonotary warblers were not small enough to exclude cowbirds, and warblers were parasitized heavily in nests with larger openings. Male and female prothonotary warblers were always away from their nests before sunrise when cowbirds laid eggs in their nests. Prothonotary warblers infrequently (∼6% of 560 nests) deserted nests that were parasitized during the egg-laying period, but frequently (56% of 151 nests) deserted nests that were parasitized before a female warbler laid her first egg. Prothonotary warblers also deserted 60-70% of nests where a cowbird egg, warbler egg or die were experimentally added before egg laying. However, the experimental addition of one of these three objects during the egg-laying period did not elicit desertion. The desertion of parasitized nests was not affected by nest site availability as has been reported elsewhere in the literature. This lack of a response to brood parasitism by prothonotary warblers may be an example of evolutionary lag, because it is likely that they have only recently been exposed to widespread parasitism, and they accept parasitism at a high cost to their own reproductive success. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Community-wide patterns of parasitism of a host “generalist” brood-parasitic cowbird

The brown-headed cowbird (Molothrusater) is a generalist obligate brood parasite. Despite intensive study and growing concern over the negative impact of cowbird parasitism on populations of many hosts, very little is known about the factors influencing community-wide patterns of cowbird parasitism. Using systematic nest searches, nest parasitism was studied over two breeding seasons at a study site in northeastern Illinois encompassing grassland, forest-edge, and forest habitat, supporting a diverse avian community. Parasitism was observed for 18 out of 34 altricial bird species found nesting at the study site. A total of 299 cowbird eggs and nestlings were found in 191 of a total of 593 nests. Analyses revealed several ecological and behavioral factors associated with frequency of parasitism and the resulting distribution of cowbird eggs. Much higher frequencies of parasitism were found in edge and forest habitats than in grassland. Within the edge habitat, open nests were parasitized significantly more often than cavity nests. Among open nests in the edge habitat, the two largest species were never parasitized. Host behavior, particularly egg-ejection behavior, was associated with a reduced observed frequency of parasitism, but at least three species known to eject cowbird eggs were sometimes parasitized. For six common hosts capable of rearing cowbirds, we found no correlation between level of parasitism and host nest-survivorship, suggesting that fine-grained assessments of host quality by female cowbirds do not influence patterns of parasitism among acceptable host species, or that differences in host quality are not great and/or predictable enough for such fine-grained assessments. Our results suggest that when a variety of possible nests are available, the level of parasitism on a particular species is a balance between a␣cowbird's preference for a particular species and the effectiveness of host species' defenses. A conceptual model was developed that incorporates the observed correlation of cowbird eggs or nestlings with habitat, nest-type, host species' body mass, and host behavioral defenses. Additional community-wide studies of cowbird parasitism will test if this model is applicable to other avian communities. Received: 20 December 1996 / Accepted: 17 May 1997     

Brood parasitism of Black‐capped Vireos: frontline and post‐laying behavioral responses and effects on productivity

The cost of brood parasitism favors the evolution of host behaviors that reduce the risk or expense of being parasitized. Endangered Black‐capped Vireos (Vireo atricapilla) have likely coexisted with brood‐parasitic Brown‐headed Cowbirds (Molothrus ater) for more than 10,000 yr, so it is likely that they have evolved anti‐parasitic behaviors. We monitored naturally parasitized and non‐parasitized vireo nests to evaluate factors that might explain parasitism risk and nest desertion behavior and also assessed whether behaviors that occurred after being parasitized improved reproductive output. Vireos reduced the risk of parasitism by initiating breeding early and nesting farther from open grasslands and edges of woody thickets. Post‐laying, nest desertion was common (70% of parasitized nests) and increased with both the presence of at least one cowbird egg in nests and clutch reduction by cowbirds. After accounting for these cues, desertion was also more likely at nests located closer to cowbird foraging habitat and below potential cowbird vantage points. Despite its regularity, desertion did not appear to provide reproductive benefits to vireos. Instead, accepting cowbird eggs was a more effective strategy because 42% of cowbird eggs did not hatch. Furthermore, cowbird eggs were somehow ejected from at least three vireo nests. Our results suggest that Black‐capped Vireos can behave in a variety of ways that reduce the impact of brood parasitism, with frontline behaviors appearing to provide the greatest benefit. Our results also suggest that habitat management should focus on providing Black‐capped Vireos with adequate breeding habitat that provides access to safe nesting sites, and with high‐quality wintering habitat that allows vireos to migrate and initiate nesting early.     

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     

UV reflectance of eggs of brown-headed cowbirds (Molothrus ater) and accepter and rejecter hosts

Many hosts of obligate brood parasites accept parasitic eggs despite the high costs of parasitism. Acceptance is particularly perplexing in brown-headed cowbird (Molothrus ater) (hereafter “cowbird”) hosts because the eggs of cowbirds and most hosts do not appear to match closely in visual characteristics detectable by humans. However, recent evidence suggests that parasite and host eggs may match in their ultraviolet (UV) reflectance, undetectable by humans, and that birds may use UV signals for egg discrimination. We determined whether egg colour matching in UV reflectance separates accepters and rejecters of cowbird parasitism by comparing the total UV (300–400 nm) reflectance of the eggs of 11 host species to cowbird eggs. Eggs of three of five accepter species and five of five rejecter species differed significantly from cowbird eggs in UV reflectance. We found no significant difference in the UV reflectance of the eggs of three closely related pairs of accepter and rejecter species. There also was no significant difference in the UV reflectance of cowbird eggs laid in nests of five host species, and the UV reflectance of cowbird eggs was not significantly correlated with that of host clutches. Thus, we found no support for the UV-matching hypothesis in brown-headed cowbirds and UV reflectance does not appear to separate accepters and rejecters of parasitism. Differences in UV reflectance between cowbird and host eggs, however, provide potential cues for use in egg discrimination. Experimental testing is needed to determine the relative importance of UV reflectance compared to other visual cues.     

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

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

Density-dependent habitat selection by brown-headed cowbirds (<Emphasis Type="Italic">Molothrus ater</Emphasis>) in tallgrass prairie

Local distributions of avian brood parasites among their host habitats may depend upon conspecific parasite density. We used isodar analysis to test for density-dependent habitat selection in brown-headed cowbirds (Molothrus ater) among tallgrass prairie adjacent to wooded edges, and prairie interior habitat (>100 m from wooded edges) with and without experimental perches. Eight study sites containing these three habitat treatments were established along a geographical gradient in cowbird abundance within the Flint Hills region of Eastern Kansas and Oklahoma, USA. The focal host species of our study, the dickcissel (Spiza americana), is the most abundant and preferred cowbird host in the prairie of this region. Cowbird relative abundance and cowbird:host abundance ratios were used as estimates of female cowbird density, whereas cowbird egg density was measured as parasitism frequency (percent of dickcissel nests parasitized), and parasitism intensity (number of cowbird eggs per parasitized nest). Geographical variation in cowbird abundance was independent of host abundance. Within study sites, host abundance was highest in wooded edge plots, intermediate in the experimental perch plots, and lowest in prairie interior. Cowbirds exhibited a pattern of density-dependent selection of prairie edge versus experimental perch and interior habitats. On sites where measures of cowbird density were lowest, all cowbird density estimates (female cowbirds and their eggs) were highest near (100 m) wooded edges, where host and perch availability are highest. However, as overall cowbird density increased geographically, these density estimates increased more rapidly in experimental perch plots and prairie interiors. Variation in cowbird abundance and cowbird:host ratios suggested density-dependent cowbird selection of experimental perch over prairie interior habitat, but parasitism levels on dickcissel nests were similar among these two habitats at all levels of local cowbird parasitism. The density-dependent pattern of cowbird distribution among prairie edge and interior suggested that density effects on perceived cowbird fitness are greatest at wooded edges. A positive relationship between daily nest mortality rates of parasitized nests during the nestling period with parasitism intensity levels per nest suggested a density-dependent effect on cowbird reproductive success. However, this relationship was similar among habitats, such that all habitats should have been perceived as being equally suitable to cowbirds at all densities. Other unmeasured effects on cowbird habitat suitability (e.g., reduced cowbird success in edge-dwelling host nests, cowbird despotism at edges) might have affected cowbird habitat selection. Managers attempting to minimize cowbird parasitism on sensitive cowbird hosts should consider that hosts in otherwise less-preferred cowbird habitats (e.g., habitat interiors) are at greater risk of being parasitized where cowbirds become particularly abundant.     

Community-level patterns of population recruitment in a generalist avian brood parasite, the brown-headed cowbird

The brown-headed cowbird (Molothrus ater) is a generalist brood parasite that typically parasitizes many host species in a single bird community. Population recruitment in a generalist parasite should be diverse with respect to host species; however, host-specific rates of cowbird recruitment have not been reported in any host community, and the determinants of host quality are poorly known. We studied the combined influence of parasitism level, nest abundance, and host quality on community-level patterns of cowbird recruitment in New Mexico, USA. Our objectives were to: (1) evaluate patterns of host use and quality; (2) compare cowbird egg investment and recruitment among host species; (3) identify host species of most importance to cowbird recruitment. Cowbirds parasitized 11 host species, with five “major” hosts experiencing high parasitism levels (≥1 cowbird egg/nest) and six minor hosts experiencing low parasitism levels (<0.3 cowbird eggs/nest). Parasitism level was not correlated with host species abundance, host mass, host nestling period length, or host success at fledging cowbirds. However, tree-nesting hosts were parasitized more than ground-nesters, and foliage-gleaners more than sally-foragers and ground-foragers. Average estimated survival to fledging of cowbird eggs laid in active host nests was 0.19. Cowbird recruitment was diverse with respect to hosts but was less evenly distributed across the host community than was cowbird egg investment because western tanagers (Piranga ludovicianus) fledged cowbirds more successfully than other hosts. This success in western tanagers was due to high cowbird survivorship in tanager nests and may be associated with the larger body size of tanagers relative to other hosts.     

Brood parasite eggs enhance egg survivorship in a multiply parasitized host

Despite the costs to avian parents of rearing brood parasitic offspring, many species do not reject foreign eggs from their nests. We show that where multiple parasitism occurs, rejection itself can be costly, by increasing the risk of host egg loss during subsequent parasite attacks. Chalk-browed mockingbirds (Mimus saturninus) are heavily parasitized by shiny cowbirds (Molothrus bonariensis), which also puncture eggs in host nests. Mockingbirds struggle to prevent cowbirds puncturing and laying, but seldom remove cowbird eggs once laid. We filmed cowbird visits to nests with manipulated clutch compositions and found that mockingbird eggs were more likely to escape puncture the more cowbird eggs accompanied them in the clutch. A Monte Carlo simulation of this 'dilution effect', comparing virtual hosts that systematically either reject or accept parasite eggs, shows that acceptors enjoy higher egg survivorship than rejecters in host populations where multiple parasitism occurs. For mockingbirds or other hosts in which host nestlings fare well in parasitized broods, this benefit might be sufficient to offset the fitness cost of rearing parasite chicks, making egg acceptance evolutionarily stable. Thus, counterintuitively, high intensities of parasitism might decrease or even reverse selection pressure for host defence via egg rejection.     

Rates of parasitism,but not allocation of egg resources,vary among and within hosts of a generalist avian brood parasite

Brown-headed cowbirds (Molothrus ater) deposit their eggs into the nests of other birds, which then raise the cowbird chick. Female cowbirds thus have limited options for impacting their offspring’s development via maternal effects compared to most other passerines. Cowbirds can impact their offspring’s phenotype by choosing among potential host nests, and by adjusting egg resources based on host characteristics. To examine whether cowbirds exhibit either or both of these strategies, we investigated rates of cowbird parasitism and egg investment (egg size, yolk-to-albumen ratio, and yolk testosterone and androstenedione) among and within host species in a shrubland bird community. We found that the probability of being parasitized by cowbirds, controlling for host status as a cowbird egg accepter or rejecter and ordinal date, varied significantly among host species, indicating an apparent preference for some hosts. Parasitism rates did not differ with host size, however, and despite variation in cowbird egg size among host species, this variation was not related to host size or cowbird preference. Among host species with eggs that are larger than those of the cowbird, cowbirds were significantly more likely to parasitize nests with relatively smaller eggs, whereas parasitism rates did not vary with relative egg size in host species with smaller eggs. There was no evidence for variation in cowbird egg components among or within host species. Our data indicate that cowbirds discriminate among host nests, but do not appear to adjust the composition of their eggs based on inter- or intraspecific host variation.     

Hatching asynchrony, nestling competition, and the cost of interspecific brood parasitism

All parental hosts of heterospecific brood parasites must paythe cost of rearing non-kin. Previous research on nest parasitismby brown-headed cowbirds (Molothrus ater) concluded that competitivesuperiority of the typically more intensively begging and largercowbird chick leads to preferential feeding by foster parentsand causes a reduction in the hosts' own brood. The larger sizeof cowbird nestlings can be the result of at least two causes:(1) cowbirds preferentially parasitize species with smallernestlings and lower growth rates; and/or (2) cowbirds hatchearlier than hosts. I estimated the cost of cowbird parasitismfor each of 29 species by calculating the difference betweenhosts' published brood sizes in nonparasitized and parasitizednests and using clutch size to standardize values. In this analysis,greater incubation length and lower adult mass, surrogate measuresof the hatching asynchrony and size difference between parasiteand hosts, were both related to greater costs of cowbird parasitismwithout bias owing to phylogeny. To establish causality, I manipulatedclutch contents of eastern phoebes (Sayornis phoebe) and examinedwhether earlier hatching by a single cowbird or phoebe egg reducesthe size of the rest of the original host brood. As predicted,greater hatching asynchrony increased the proportion of theoriginal phoebe brood that was lost. This measure of the costof parasitism was partially owing to increased hatching failureof the original eggs in asynchronous broods but was not at allrelated to the size differences of older and younger conspecificnestmates. However, proportional brood loss owing to an earlierhatching conspecific was consistently smaller than brood lossowing to asynchronous cowbirds in both naturally and experimentallyparasitized phoebe nests. These results imply that althoughhatching asynchrony is an important cause of the reduction ofhost broods in parasitized clutches, competitive features ofcowbird nestlings remain necessary to explain the full extentof hosts' reproductive costs caused by interspecific brood parasitism.     

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.     

BEHAVIORAL DEFENSES AGAINST AVIAN BROOD PARASITISM IN SYMPATRIC AND ALLOPATRIC HOST POPULATIONS

The brown-headed cowbird (Molothrus ater) is a widespread, obligate brood parasite of North American passerine birds. In southern Manitoba, where hosts are sympatric with cowbirds, American robins (Turdus migratorius) ejected parasitic eggs from all experimentally parasitized clutches (N = 25) and no eggs were accepted for more than four days. In contrast, robins in northern Manitoba, an area where cowbirds do not breed, accepted parasitic eggs in 33% of nests (N = 18) for at least five days. Acceptance of experimental cowbird eggs by a second host, the yellow warbler (Dendroica petechia), was similar in allopatric (100% of 20 nests) and sympatric (88.6% of 35 nests) populations, but models of a female cowbird elicited greater nest defense by warblers in the area of sympatry. Neither host rejected eggs of conspecifics, thus, rejection of cowbird eggs was not an epiphenomenon of conspecific brood parasitism. These results support the hypothesis that recognition of cowbirds and their eggs evolved as adaptations to counter cowbird parasitism and not some other selection pressure. The expression of anti-parasite defenses by some individuals within allopatric populations further suggests these traits may be controlled genetically but persist in such areas either through the continued introgression of rejecter genes from sympatric populations or because of the low cost of rejection behavior when parasitism is absent or rare.     

Evolutionary lag versus bill-size constraints: a comparative study of the acceptance of cowbird eggs by old hosts

Summary Evolutionary lag and resistance to puncture-ejection are two hypotheses explaining why hosts long exposed to brood parasitism by molothrine cowbirds accept their eggs. The former hypothesis assumes no physical constraints while the latter requires them. We show (1) that most acceptors of Brown-headed Cowbird eggs have small bills, (2) that the single species known to puncture cowbird eggs for ejection also has a small bill, and (3) that all grasp ejectors of Brown-headed Cowbird eggs have large bills. We propose that most acceptors cannot remove cowbird eggs by grasp ejection because their bills are too small. Small hosts probably cannot puncture cowbird eggs for ejection because their unusually thick shells make the costs of puncturing them higher than the benefits of ejection. This hypothesis is capable of explaining the success of cowbirds with scores of host species in the puzzling absence of egg mimicry.     

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.     

To eject or to abandon? Life history traits of hosts and parasites interact to influence the fitness payoffs of alternative anti‐parasite strategies

Hosts either tolerate avian brood parasitism or reject it by ejecting parasitic eggs, as seen in most rejecter hosts of common cuckoos, Cuculus canorus, or by abandoning parasitized clutches, as seen in most rejecter hosts of brown‐headed cowbirds, Molothrus ater. What explains consistent variation between alternative rejection behaviours of hosts within the same species and across species when exposed to different types of parasites? Life history theory predicts that when parasites decrease the fitness of host offspring, but not the future reproductive success of host adults, optimal clutch size should decrease. Consistent with this prediction, evolutionarily old cowbird hosts, but not cuckoo hosts, have lower clutch sizes than related rarely‐ or newly parasitized species. We constructed a mathematical model to calculate the fitness payoffs of egg ejector vs. nest abandoner hosts to determine if various aspects of host life history traits and brood parasites’ virulence on adult and young host fitness differentially influence the payoffs of alternative host defences. These calculations showed that in general egg ejection was a superior anti‐parasite strategy to nest abandonment. Yet, increasing parasitism rates and increasing fitness values of hosts’ eggs in both currently parasitized and future replacement nests led to switch points in fitness payoffs in favour of nest abandonment. Nonetheless, nest abandonment became selectively more favourable only at lower clutch sizes and only when hosts faced parasitism by a cowbird‐ rather than a cuckoo‐type brood parasite. We suggest that, in addition to evolutionary lag and gape‐size limitation, our estimated fitness differences based on life history trait variation provide new insights for the consistent differences observed in the anti‐parasite rejection strategies between many cuckoo‐ and cowbird‐hosts.     

Individual patterns of habitat and nest-site use by hosts promote transgenerational transmission of avian brood parasitism status

Brood parasitic birds impose variable fitness costs upon their hosts by causing the partial or complete loss of the hosts' own brood. Growing evidence from multiple avian host-parasite taxa indicates that exposure of individual hosts to parasitism is not necessarily random and varies with habitat use, nest-site selection, age or other phenotypic attributes. For instance, nonrandom patterns of brood parasitism had similar evolutionary consequences to those of limited horizontal transmission of parasites and pathogens across space and time and altered the dynamics of both population productivity and co-evolutionary interactions of hosts and parasites. We report that brood parasitism status of hosts of brown-headed cowbirds Molothrus ater is also transmitted across generations in individually colour-banded female prothonotary warblers Protonotaria citrea. Warbler daughters were more likely to share their mothers' parasitism status when showing natal philopatry at the scale of habitat patch. Females never bred in their natal nestboxes but daughters of parasitized mothers had shorter natal dispersal distances than daughters of nonparasitized mothers. Daughters of parasitized mothers were more likely to use nestboxes that had been parasitized by cowbirds in both the previous and current years. Although difficult to document in avian systems, different propensities of vertical transmission of parasitism status within host lineages will have critical implications both for the evolution of parasite tolerance in hosts and, if found to be mediated by lineages of parasites themselves, for the difference in virulence between such extremes as the nestmate-tolerant and nestmate-eliminator strategies of different avian brood parasite species.     

Multiple effects of brood parasitism reduce the reproductive success of prothonotary warblers, Protonotaria citrea

Avian brood parasitism often has multiple negative effects on the reproductive success of the host. Most studies have focused on one or two of these effects, but rarely have they all been studied simultaneously for one species. I studied prothonotary warblers to quantify the effects of different intensities of (i.e. multiple) brood parasitism by brown-headed cowbirds, Molothrus ater, on the production of host and cowbird young and on the between-year returns of adult warblers. Host clutch size decreased with an increase in the number of cowbird eggs laid in nests. The hatching success of warbler and cowbird eggs decreased with increased cowbird eggs in nests, but was always higher for cowbird eggs than warbler eggs. The survival of warbler nestlings, but not cowbird nestlings, decreased with increased cowbird nestlings in the brood. An increase in the number of cowbird nestlings in broods resulted in a reduction in the average mass of warbler nestlings but not cowbird nestlings. The number of cowbird eggs or nestlings present did not affect nest predation, and the fledging of cowbirds did not influence the renesting interval of female warblers. In addition, the between-year returns of adult warblers were not negatively affected by brood parasitism. Decreased hatching success and nestling survival reduced the reproductive output of the warblers the most. These effects were substantial and appear to favour the evolution of behavioural responses that reduce the effects of brood parasitism on prothonotary warblers. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour      

Parameters of brown-headed cowbird Molothrus ater egg discrimination in warbling vireos Vireo gilvus

We examined which egg parameters warbling vireos Vireo gilvus use to discriminate brown-headed cowbird Molothrus ater eggs and, by comparing our results to other studies, tested the prediction that ejecter species with eggs more similar in appearance to cowbird eggs will be less tolerant of foreign eggs. In addition, we tested the hypothesis that egg characteristics influence the cost of ejection and probability of committing ejection errors. Warbling vireos ejected 100% of eggs with a cowbird spot pattern and only spot pattern significantly influenced the probability of ejecting a foreign egg, whereas size and nest stage did not. Foreign eggs that differed in two parameters were not ejected significantly more than those that differed in one parameter. Thus, warbling vireos appear to be less tolerant of foreign eggs than species with eggs more divergent from cowbird eggs. There was no significant difference in the number of vireo eggs that were damaged when foreign eggs of different sizes and spot patterns were ejected, which is counter to the assumptions of the evolutionary equilibrium hypothesis. Similarly, foreign egg characteristics did not significantly influence the probability of ejection errors. Finally, egg discrimination in warbling vireos appears to have evolved directly to counter cowbird parasitism because all conspecific eggs switched into their nests were accepted.