Habitat and Landscape Effects on Brood Parasitism,Nest Survival,and Fledgling Production in Swainson's Warblers

ABSTRACT Numerous factors, including nest predation and brood parasitism, may limit populations of neotropical migratory birds. However, nest predation and brood parasitism are not constant, and temporal, biological, habitat, and landscape factors can affect the likelihood of these events. Understanding these patterns is important for species of conservation concern for which managers seek to provide quality habitat. One such species, the Swainson's warbler (Limnothlypis swainsonii), is a neotropical migrant that breeds primarily in bottomland hardwood forests of the southeastern United States. Little is known of factors influencing reproductive success of this rare, yet locally abundant, species. From 2004 through 2007, we examined factors influencing reproductive success of Swainson's warblers at 2 sites in eastern Arkansas, USA, St. Francis National Forest and White River National Wildlife Refuge. We used 2-stage modeling to assess the relationship between 1) temporal and biological, and 2) habitat and landscape factors and brood parasitism, nest survival, and fledgling production. Brood parasitism was greater in this population (36%) than reported elsewhere (≤ 10%), but decreased throughout the breeding season. Nest survival was comparable to or lower than in other populations of this species and increased throughout the breeding season. The probability of brood parasitism was greater near forest edges. Although nests of Swainson's warblers were often associated with giant cane (Arundinaria gigantea), nest survival had a weak negative association with cane density. For nests that were successful, the best predictor of number of Swainson's warblers fledged was brood-parasitism status: nonparasitized nests fledged 2.75 young, whereas parasitized nests fledged 0.60 Swainson's warblers. Our findings suggest that managing and restoring relatively high-elevation bottomland forests that are located far from agricultural edges should increase Swainson's warbler productivity.     

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      

Brood parasitism increases provisioning rate, and reduces offspring recruitment and adult return rates, in a cowbird host

Interspecific brood parasitism in birds presents a special problem for the host because the parasitic offspring exploit their foster parents, causing them to invest more energy in their current reproductive effort. Nestling brown-headed cowbirds (Molothrus ater) are a burden to relatively small hosts and may reduce fledgling quality and adult survival. We documented food-provisioning rates of one small host, the prothonotary warbler (Protonotaria citrea), at broods that were similar in age (containing nestlings 8–9 days old), but that varied in composition (number of warbler and cowbird nestlings) and mass, and measured the effect of brood parasitism on offspring recruitment and adult returns in the host. The rate of food provisioning increased with brood mass, and males and females contributed equally to feeding nestlings. Controlling for brood mass, the provisioning rate was higher for nests with cowbirds than those without. Recruitment of warbler fledglings from unparasitized nests was 1.6 and 3.7 times higher than that of fledglings from nests containing one or two cowbirds, respectively. Returns of double-brooded adult male and female warblers decreased with an increase in the number of cowbirds raised, but the decrease was more pronounced in males. Reduced returns of warbler adults and recruitment of warbler fledglings with increased cowbird parasitism was likely a result of reduced survival. Cowbird parasitism increased the warblers’ investment in current reproductive effort, while exerting additional costs to current reproduction and residual reproductive value. Our study provides the strongest evidence to date for negative effects of cowbird parasitism on recruitment of host fledglings and survival of host adults.     

A generalist brood parasite modifies use of a host in response to reproductive success

Avian obligate brood parasites, which rely solely on hosts to raise their young, should choose the highest quality hosts to maximize reproductive output. Brown-headed cowbirds (Molothrus ater) are extreme host generalists, yet female cowbirds could use information based on past reproductive outcomes to make egg-laying decisions thus minimizing fitness costs associated with parasitizing low-quality hosts. We use a long-term (21 years) nest-box study of a single host, the prothonotary warbler (Protonotaria citrea), to show that local cowbird reproductive success, but not host reproductive success, was positively correlated with the probability of parasitism the following year. Experimental manipulations of cowbird success corroborated that female cowbirds make future decisions about which hosts to use based on information pertaining to past cowbird success, both within and between years. The within-year pattern, in particular, points to local cowbird females selecting hosts based on past reproductive outcomes. This, coupled with high site fidelity of female cowbirds between years, points to information use, rather than cowbird natal returns alone, increasing parasitism rates on highly productive sites between years.     

Comparative detection,density, and reproductive performance of Kirtland's warbler in jack and red pine

The formerly endangered Kirtland's warbler (Setophaga kirtlandii) is among a growing number of conservation-reliant species that depend on active management to avoid reverting to endangered status. Because the Kirtland's warbler is a habitat specialist of young, even-aged jack pine (Pinus banksiana), managers of the recovery effort stressed creating new jack pine stands and monitoring numbers of singing males through an annual census using single visits to individual stands. Kirtland's warbler will occupy and breed in red pine (P. resinosa), but red pine has not been surveyed for Kirtland's warblers in the annual population census. Furthermore, the current monitoring approach cannot determine their species detection probability or individual detection probability, which is essential to evaluate both red pine use and the accuracy of the census. From 2016–2018 we estimated density and detection probabilities in jack pine and red pine stands through repeated visits to a limited number of stands rather than single visits to many stands. Estimates of species detection probability indicated that ≥1 male Kirtland's warbler would be detected on most sites when any were present, but individual detection probabilities were less and varied by stand type, indicating that single visits to sites would underestimate numbers and that accurate estimation of detection probability was important for estimation of density in different stand types. We offer quantitative estimates of detection probabilities for determination of Kirtland's warbler population size in jack pine versus red pine stands in the same areas and breeding seasons. Managers of Kirtland's warblers should incorporate detection probabilities into population surveys to achieve more accurate estimates of population size.     

Cattle grazing in CRP grasslands during the nesting season: effects on avian reproduction

Bird populations in grasslands have experienced declines coinciding with loss and fragmentation of prairies. The United States Department of Agriculture (USDA)-administered Conservation Reserve Program (CRP) is the most extensive grassland restoration program in North America and it has especially benefitted grassland birds. Grazing by domestic cattle has been restricted in CRP during avian nesting seasons despite the potential improvements in structuring habitat for a greater diversity of grassland bird species. Potential negative consequences of grazing in CRP grasslands include trampling of nests by cattle, reductions in nest concealment from predators, and attraction of brood-parasitic brown-headed cowbirds (Molothrus ater). We designed an experiment to test for effects of cattle grazing in CRP fields during the nesting season on nest survival and brood parasitism of 5 bird species that commonly nest in CRP grasslands: mourning dove (Zenaida macroura), grasshopper sparrow (Ammodramus savannarum), dickcissel (Spiza americana), and eastern (Sturnella magna) and western (S. neglecta) meadowlarks. Grazing was implemented during summers 2017 and 2018 on 17 of 36 fields followed by a year of rest on all fields in 2019. Of the 879 nests on grazed fields, only 4 were likely trampled by cattle (vs. 54% of all nests estimated as failing because of depredation). Experimental grazing (grazed vs. ungrazed fields) had variable effects on nest survival and cowbird parasitism among the bird species analyzed. Negative effects of grazing on daily nest survival of dickcissel and meadowlarks were apparent, at least in some years. We found no direct effects of grazing on nest survival of mourning dove or grasshopper sparrow. Probability and intensity (cowbird offspring/nest) of cowbird parasitism in dickcissel nests was higher on grazed versus ungrazed sites but only in conservation practice (CP) CP2 (vs. CP25 fields). Parasitism probability of grasshopper sparrow nests by cowbirds was higher on grazed fields in the 2 years after introduction of cattle in 2017. Greater vegetative concealment around nest sites was associated with reduced cowbird parasitism of meadowlark and grasshopper sparrow nests and higher nest survival for grasshopper sparrows. Reductions in vegetative height caused by longer-term or high-intensity grazing might therefore have negative consequences for some grassland birds by increasing nest site visibility and exposure to cowbird parasitism. Our results indicate that cattle grazing in CRP fields during the nesting season might have some negative effects on reproductive success of some grassland bird species, at least in the short term; however, the potential improvements of structuring habitat to accommodate more grassland bird species and increasing landowner participation in the CRP are considerable.     

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     

Evidence of a Minimum Patch Size Threshold of Reproductive Success in an Endangered Songbird

ABSTRACT That area-sensitive songbirds breed only in relatively large patches suggests that there may be a minimum patch size threshold in which they will breed, even when controlling for the total amount of habitat in the landscape. We searched for minimum patch size thresholds of presence, territory establishment by males, pairing success, and reproductive success for 2 migratory songbirds that differed in sensitivity to patch size: golden-cheeked warblers (Dendroica chrysoparia) and white-eyed vireos (Vireo griseus). We assessed 2 potential limiting factors: brown-headed cowbird (Molothrus ater) parasitism and arthropod biomass (food resource). We determined whether either factor was related to patch size and compared measurements of each above and below the observed thresholds. We monitored 24 golden-cheeked warbler and 47 white-eyed vireo territories in 12 patches. We found evidence of a minimum patch size threshold (between 15.0 ha and 20.1 ha) of reproductive success for golden-cheeked warblers, but not for white-eyed vireos. We found no minimum patch size thresholds for presence, territory establishment by males, or pair formation for either species. Conservation practices based on thresholds of presence, territory establishment, or pair formation might not address issues of reproduction for golden-cheeked warblers. We failed to find evidence that cowbird parasitism or arthropod biomass were limiting factors. The ability to identify patch size thresholds of reproductive success for target species could be useful in conservation and management for setting goals for retention and restoration of target species’ habitat patch size.     

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.     

Host switching in cowbird brood parasites: how often does it occur?

Avian obligate brood parasites lay their eggs in nests of host species, which provide all parental care. Brood parasites may be host specialists, if they use one or a few host species, or host generalists, if they parasitize many hosts. Within the latter, strains of host‐specific females might coexist. Although females preferentially parasitize one host, they may occasionally successfully parasitize the nest of another species. These host switching events allow the colonization of new hosts and the expansion of brood parasites into new areas. In this study, we analyse host switching in two parasitic cowbirds, the specialist screaming cowbird (Molothrus rufoaxillaris) and the generalist shiny cowbird (M. bonariensis), and compare the frequency of host switches between these species with different parasitism strategies. Contrary to expected, host switches did not occur more frequently in the generalist than in the specialist brood parasite. We also found that migration between hosts was asymmetrical in most cases and host switches towards one host were more recurrent than backwards, thus differing among hosts within the same species. This might depend on a combination of factors including the rate at which females lay eggs in nests of alternative hosts, fledging success of the chicks in this new host and their subsequent success in parasitizing it.     

Distance from riparian edge reduces brood parasitism of southwestern willow flycatchers,whereas parasitism increases nest predation risk

The southwestern willow flycatcher (Empidonax traillii extimus) is a federally endangered subspecies that breeds in increasingly fragmented and threatened habitat. We examined whether temporal and habitat characteristics were associated with risk of predation and probability of brood parasitism by brown-headed cowbirds (Molothrus ater) on flycatcher nests at 6 sites in southern Nevada and northwestern Arizona, USA. For nest predation, we found the most support for a model that included date and an interaction between parasitism status and nesting stage. Daily nest survival decreased from 0.87 (95% CI = 0.81–0.93) to 0.78 (95% CI = 0.72–0.84) through the season for parasitized nests but remained relatively constant for unparasitized nests (0.93, 95% CI = 0.91–0.95 to 0.92, 95% CI = 0.91–93). Parasitized nests had lower survival than non-parasitized nests during the incubation (0.85, 95% CI = 0.84–0.86 vs. 0.92, CI = 0.91–0.93) and nestling (0.79, 95% CI = 0.77–0.81 vs. 0.91, 95% CI = 0.90–0.92) stages. Of the variables included in our parasitism candidate models, model-averaged coefficients and odds ratios supported only distance to habitat edge; odds of parasitism decreased 1% for every 1 m from the habitat edge. Nests greater than 100 m from an edge were 50% less likely to be parasitized as those on an edge, however, only 52 of 233 nests (22%) were found at this distance. Where management and conservation goals include reducing nest losses due to parasitism, we recommend restoration of habitat patches that minimize edge and maximize breeding habitat further from edges. At sites where cowbirds have been documented as important nest predators, controlling cowbirds may be one option, but further study of the link between parasitism and nest predation and the identification of major nest predators at specific sites is warranted. © 2011 The Wildlife Society.     

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.     

REED WARBLER HOSTS FINE‐TUNE THEIR DEFENSES TO TRACK THREE DECADES OF CUCKOO DECLINE

Interactions between avian hosts and brood parasites can provide a model for how animals adapt to a changing world. Reed warbler (Acrocephalus scirpaceus) hosts employ costly defenses to combat parasitism by common cuckoos (Cuculus canorus). During the past three decades cuckoos have declined markedly across England, reducing parasitism at our study site (Wicken Fen) from 24% of reed warbler nests in 1985 to 1% in 2012. Here we show with experiments that host mobbing and egg rejection defenses have tracked this decline in local parasitism risk: the proportion of reed warbler pairs mobbing adult cuckoos (assessed by responses to cuckoo mounts and models) has declined from 90% to 38%, and the proportion rejecting nonmimetic cuckoo eggs (assessed by responses to model eggs) has declined from 61% to 11%. This is despite no change in response to other nest enemies or mimetic model eggs. Individual variation in both defenses is predicted by parasitism risk during the host's egg‐laying period. Furthermore, the response of our study population to temporal variation in parasitism risk can also explain spatial variation in egg rejection behavior in other populations across Europe. We suggest that spatial and temporal variation in parasitism risk has led to the evolution of plasticity in reed warbler defenses.     

Effects of Jack Pine Plantation Management on Barrens Flora and Potential Kirtland's Warbler Nest Habitat

Jack pine barrens, once common in northern lower Michigan, mostly have been converted to managed jack pine plantations. Management of the disturbances associated with logging provides the opportunity to maintain the unique plant assemblages of jack pine barrens and nest habitat of the federally endangered Kirtland's warbler. Studies indicate that Carex pensylvanica can develop into dense mats and strongly compete with other barrens species such as Vaccinium angustifolium, which seem to be important species for Kirtland's warbler nest locations. According to forest managers, the most important factors facilitating high cover of V. angustifolium and reducing cover of C. pensylvanica are the amount of shade produced by tree crowns before harvest (pre‐harvest shade), the length of time between harvest and planting (planting delay), and fire. We found that high or low levels of pre‐harvest shade had no effect on cover of either V. angustifolium or C. pensylvanica. Planting delays of at least three years following prescribed burns generally increased cover of V. angustifolium in forest plots, which are important for warbler nesting. Analysis of community composition in openings indicated that burning enhanced the growth of barrens species. We found only weak evidence for a negative correlation between the cover of V. angustifolium and C. pensylvanica on our study sites. The openings created in the jack pine plantation are important refugia for barrens flora that would likely be lost under forests managed strictly for jack pine. Maintenance of jack pine barrens flora and Kirtland's warbler nest habitat is possible within the context of a heavily managed forest plantation system.     

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.     

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.      

Brood‐parasite‐induced female‐biased mortality affects songbird demography: negative implications for conservation

Parasites, of all sorts, can profoundly affect host population dynamics. Parasites commonly cause sex‐biased mortality and this can add to their impact. Female‐biased mortality in particular can destabilize dynamics and promote population collapse. We previously reported in a correlative study that brown‐headed cowbird Molothrus ater brood parasitism of song sparrows Melospiza melodia appears to cause female‐biased host nestling mortality. Here, we report results from ‘infestation’ and ‘de‐infestation’ experiments designed to test whether brood parasitism causes female‐biased mortality, and we document the resulting demographic impact using a simulation model. Experimental cowbird infestation of song sparrow nests halved the proportion of female host nestlings (0.31±0.07 vs 0.59±0.06; infested vs unparasitized nests at day 6) replicating the halving reported in naturally cowbird‐parasitized nests (0.28±0.01 vs 0.57±0.05; parasitized vs unparasitized). De‐infestation of naturally cowbird‐parasitized nests in turn wholly eliminated any effect on the proportion of female host nestlings (0.53±0.13 vs 0.54±0.06; de‐infested vs unparasitized) confirming that brood parasitism is the cause. This halving of the proportion of females fledging is likely to be as significant as nest predation in affecting population dynamics, based on the elasticities derived from our demographic model (–0.50 vs –0.59). Experimental infestation reduced the testosterone levels, begging behaviour, and body mass of six day old female host nestlings, whereas males were largely unaffected, suggesting that it is the exacerbation of intra‐brood competition that may be primarily responsible for the resulting female‐biased mortality. The brown‐headed cowbird is invasive in most of North America and has been implicated in regional population declines of many native species. We suggest that female‐biased host offspring mortality is likely to be commonplace among the 144 host species the cowbird successfully parasitizes, and we discuss the negative implications for songbird conservation, given the projected demographic impact.     

Could diffuse coevolution explain the generic eggshell color of the brown-headed cowbird?

The brown-headed cowbird (hereafter cowbird) is an avian brood parasite that produces an egg dissimilar to those produced by the majority of its diverse host community. The cowbird’s generic egg may result from a Jack-of-all-trades strategy; however, the evolutionary mechanisms that select for their generic eggs are unclear. Here we propose that the cowbird’s eggshell phenotypes have evolved via diffuse coevolution, which results from community-level selective pressures, rather than via pairwise coevolution that occurs between a particular host species and its brood parasite. Under diffuse coevolution the cowbird’s host community, with varying eggshell phenotypes and recognition abilities, would select for a cowbird eggshell phenotype intermediate to those of its host community. This selection is exerted by hosts that reject cowbird eggs, rather than those that accept them; therefore, we expect cowbird eggshell colors can be approximated by both the phenotypes and rejection abilities of their host community. Here we use eggshell reflectance data from 43 host species to demonstrate that the cowbird eggshell phenotypes are reasonably predicted (within 2 just noticeable differences) by the eggshell phenotypes and rejection rates of their hosts. These findings suggest that cowbird eggshell phenotypes, and potentially those of other some generalist parasites, may evolve via diffuse coevolution. Importantly, this research provides insight into the underlying evolutionary processes that explain observed phenotypic variation and provides a framework for studying selection on both specialist and generalist parasites’ traits.     

Repeatability of Host Female and Male Aggression Towards a Brood Parasite

Current research on behavioural consistency showed that various types of animal behaviour are highly repeatable in the context of mate choice, exploration and parental care, including nest protection. However, the repeatability of aggressive nest defence has not yet been studied in hosts of brood parasites, although host aggression against adult parasites represents a crucial line of antiparasitic defences. Here, we investigated the between‐season repeatability of the great reed warbler (Acrocephalus arundinaceus) aggression towards a stuffed dummy of the brood parasitic common cuckoo (Cuculus canorus). We found that under the relatively stable risk of brood parasitism across breeding seasons, female responses to the cuckoo were highly repeatable, whereas male responses were variable. We suggest that the potential explanation for the observed patterns of female and male behaviours may lie in female's prominent roles in offspring care and nest protection, and in her lower renesting potential in comparison with that of males. However, further studies on the relationship between host aggression and other types of behaviours (host personality) and their fitness consequences under the fluctuating parasitism pressures are required to clarify the adaptive significance of aggressive behaviour observed in hosts of brood parasites.     

The ghosts of parasitism past: lingering frontline anti-brood parasite defenses in a former host

Coevolutionary arms races between brood parasites and hosts provide tractable systems for understanding antagonistic coevolution in nature; however, little is known about the fate of frontline antiparasite defenses when the host “wins” the coevolutionary arms race. By recreating bygone species interactions, using artificial parasitism experiments, lingering defensive behaviors that evolved in the context of parasitism can be understood and may even be used to identify the unknown agent of parasitism past. Here we present the first study of this type by evaluating lingering “frontline” nest defenses that have evolved to prevent egg laying in a former brood parasite host. The Australian reed warbler Acrocephalus australis is currently not parasitized but is known to exhibit fine-tuned egg discrimination—a defensive behavior indicative of a past brood parasite–host arms race and common in closely related parasitized species. Here, using 3D-printed models of adult brood parasites, we examined whether the Australian reed warbler also exhibits frontline defenses to adult brood parasites, and whether we could use these defenses to identify the warbler’s “ghost of parasitism past.” Our findings provide evidence that the Australian reed warbler readily engages in frontline defenses that are considered adaptive specifically in the context of brood parasitism. However, individuals were unable to discriminate between adults of different brood parasite species at their nest. Overall, our results demonstrate that despite a relaxation in selection, defenses against brood parasitism can be maintained across multiple stages of the host’s nesting cycle, and further suggest that, in accordance with previous findings, that learning may be important for fine-tuning frontline defense.