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.     

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.     

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.     

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     

Reducing the conservation reliance of the endangered Kirtland's warbler through adaptive management

Species are considered conservation-reliant when their continued existence is dependent on human assistance. Conservation reliance challenges the conservation community in terms of their ability to sustain the funding and public-private partnerships needed for indefinite management. Although increased funding for conservation is critical, reducing conservation reliance through adaptive management represents an attractive alternative. We used a large-scale ecological experiment as a case study in the use of adaptive management to reduce conservation reliance. For >40 years, the United States Fish and Wildlife Service has trapped and lethally removed an obligate brood parasite, the brown-headed cowbird (Molothrus ater), to protect the Kirtland's warbler (Setophaga kirtlandii) from the negative effects that brood parasitism has on its reproductive success. To determine if the conservation reliance of the Kirtland's warbler could be reduced through optimization of the cowbird control program, we used an adaptive management approach. In collaboration with stakeholders, we experimentally reduced cowbird trapping effort across nearly all of the Kirtland's warbler breeding range. We monitored the resulting cowbird abundance and rate of parasitism, and then adjusted the scale of trap reductions based on the previous year's results. Despite reducing (2015–2017) and eventually eliminating (2018) cowbird trapping, we detected only 20 cowbirds (2015–2017) and found that just 4 of 514 (<1%) nests were parasitized (2015–2018). Our results indicate that the cowbird control program can at least temporarily be suspended, thereby reducing conservation reliance in the Kirtland's warbler and freeing funds for other management. We urge the conservation community to consider the use of adaptive management to reduce conservation reliance in other threatened and endangered taxa. © 2019 The Wildlife Society.     

Urban-associated habitat alteration promotes brood parasitism of Acadian Flycatchers

ABSTRACT.   High rates of brood parasitism are generally associated with agricultural landscapes, but recent evidence suggests that urbanization may also increase the likelihood of brood parasitism. I evaluated the extent to which brood parasitism by Brown-headed Cowbirds ( Molothrus ater ) was explained by differences in (1) body size of adult hosts, presumably relating to the ability to defend nest from cowbirds, (2) nest placement in substrate and relative to habitat edges, (3) habitat structure surrounding nests, (4) host density, (5) cowbird abundance, both absolute and relative to host numbers, (6) landscape composition, and (7) Julian date. From 2001 to 2006, I monitored nest fate and measured vegetation characteristics surrounding nests of Acadian Flycatcher ( Empidonax virescens ) breeding in mature riparian forests in central Ohio, USA. The likelihood that a nest would be parasitized was best explained by the number of understory stems surrounding the nest and, to a lesser extent, by the amount of urbanization in the surrounding 1-km-radius landscape. Parasitized nests were surrounded by 1.6 times more stems and nearly twice the amount of urbanization than nonparasitized nests. Numbers of understory stems were positively associated with increasing urbanization, primarily due to invasion of urban forests by Amur honeysuckle ( Lonicera maackii ). Thus, urban-associated changes in habitat characteristics around nests may be important contributors to the greater vulnerability of urban nests to brood parasitism than nests in more rural landscapes. This pattern suggests that ecological restoration, such as removing exotic shrubs, may be an effective strategy to ameliorate certain negative consequences of urbanization near wooded reserves.     

Genetic analysis of brown-headed cowbirds Molothrus ater raised by different hosts: data from mtDNA and microsatellite DNA markers

Ecological and behavioural data suggest that female brood parasitic brown-headed cowbirds Molothrus ater are host generalists; this predicts that there should be no genetically differentiated host races within cowbird populations. We tested this hypothesis by comparing differentiation in two rapidly evolving DNA markers (mtDNA control region sequence and nuclear DNA microsatellite loci) among unrelated cowbird chicks raised by two ecologically distinct hosts. No differentiation was observed in either marker supporting the absence of host race hypothesis.     

Eggs yield nuclear DNA from egg-laying female cowbirds, their embryos and offspring

Here we report methods forextracting maternal DNA from avian eggshells oroffspring DNA from eggshells and embryos. Thesemethods offer alternative techniques forobtaining DNA from oviparous organisms. UsingDNA extracted from eggshells, we obtainmicrosatellite genotypes of the brood parasiticbrown-headed cowbird (Molothrus ater)female that laid the eggs and/or her hatchedoffspring. Using DNA extracted from embryos, weobtain microsatellite genotypes of offspring.We demonstrate that separate extractionsperformed on the embryo and shell from a singleegg can provide DNA from the embryo and itsmother, respectively. This single-egg approachfor obtaining both maternal and embryonic DNAsimplifies paternity analyses because allelesunique to the embryo can be considered paternalin origin. Finally we report two newmicrosatellite loci and primer sequences forbrown-headed cowbirds.     

Partial host fidelity in nest selection by the shiny cowbird (Molothrus bonariensis), a highly generalist avian brood parasite

Obligate avian brood parasites can be host specialists or host generalists. In turn, individual females within generalist brood parasites may themselves be host specialists or generalists. The shiny cowbird Molothrus bonariensis is an extreme generalist, but little is known about individual female host fidelity. We examined variation in mitochondrial control region sequences from cowbird chicks found in nests of four common Argentinean hosts. Haplotype frequency distributions differed among cowbird chicks from nests of these hosts, primarily because eggs laid in nests of house wrens Troglodytes aedon differed genetically from those laid in nests of the other three hosts (chalk-browed mockingbird Mimus saturninus, brown-and-yellow marshbird Pseudoleistes virescens, and rufous-collared sparrow Zonotrichia capensis). These differences in a maternally inherited marker indicate the presence of a nonrandom laying behaviour in the females of this otherwise generalist brood parasite, which may be guided by choice for nest type, as house wrens nest in cavities whereas the other three species are open cup nesters.     

Is Female Visual Signaling to Male Song Socially Regulated in Brown‐headed Cowbirds?

This research focused on how adult female brown‐headed cowbirds, Molothrus ater, regulate social feedback on a group level to shape the development of male song. Specifically, females produce rapid wing movements in response to male song, termed ‘wing strokes,’ which have been shown to shape male song and predict song quality. These effects have been documented in captive dyads and triads, but not in more naturalistic flocks, where song development actually occurs. Here, we studied wing stroking in small seminatural flocks of differing female‐to‐male ratios. Despite differences in the number of females and their social selectivity, the same pattern of female feedback emerged in seven of eight flocks: One female produced the majority of wing strokes to male song, making her the primary wing stroker in her flock. Previous studies on large flocks have demonstrated females to facilitate male song improvisation and development if they exhibited higher social selectivity by approaching immature males less. Here, we found that primary wing strokers were indeed more socially selective than non‐primary wing strokers. This research is the first to document social stimulation being facilitated at the group level to ensure that more highly selective females deliver the most feedback.     

Host–parasite coevolution beyond the nestling stage? Mimicry of host fledglings by the specialist screaming cowbird

Egg mimicry by obligate avian brood parasites and host rejection of non-mimetic eggs are well-known textbook examples of host-parasite coevolution. By contrast, reciprocal adaptations and counteradaptations beyond the egg stage in brood parasites and their hosts have received less attention. The screaming cowbird (Molothrus rufoaxillaris) is a specialist obligate brood parasite whose fledglings look identical to those of its primary host, the baywing (Agelaioides badius). Such a resemblance has been proposed as an adaptation in response to host discrimination against odd-looking young, but evidence supporting this idea is scarce. Here, we examined this hypothesis by comparing the survival rates of young screaming cowbirds and non-mimetic shiny cowbirds (Molothrus bonariensis) cross-fostered to baywing nests and quantifying the similarity in plumage colour and begging calls between host and cowbird fledglings. Shiny cowbirds suffered higher post-fledging mortality rates (83%) than screaming cowbirds (0%) owing to host rejection. Visual modelling revealed that screaming cowbirds, but not shiny cowbirds, were indistinguishable from host young in plumage colour. Similarly, screaming cowbirds matched baywings' begging calls more closely than shiny cowbirds. Our results strongly support the occurrence of host fledgling mimicry in screaming cowbirds and suggest a role of visual and vocal cues in fledgling discrimination by baywings.     

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.     

Edge effects reduce the nesting success of Acadian Flycatchers in a moderately fragmented forest

ABSTRACT.   Forest fragmentation can create negative edge effects that reduce the reproductive success of birds nesting near the forest/nonforest interface, and threaten bird populations deeper in remnant forest habitats. Negative edge effects may be more pronounced in landscapes that are moderately fragmented, particularly where agriculture is the primary land-use fragmenting forests. Information about the extent and strength of edge effects at a site can help guide conservation actions, and determine their effectiveness. We examined edge effects for birds breeding in a nearly contiguous forest fragmented by relatively narrow agricultural corridors in Illinois (USA). We measured rates of nest predation and brood parasitism for Acadian Flycatchers ( Empidonax virescens ) over a continuum of distances from the edge of an agricultural inholding. Nest predation and brood parasitism were highest near the edge and decreased with increasing distance from the edge. Given the cumulative effects of nest predation and brood parasitism on reproductive success, we determined that forest within 600 m of the inholding was sink habitat. We found, however, that deeper forest interior areas currently serve as source habitat, and that conversion of the entire 205 ha agricultural corridor to forest would add 1350 ha of source habitat for Acadian Flycatchers. Such results provide support for a local conservation strategy of forest consolidation and establish baseline measures necessary to determine the relative effectiveness of any subsequent reforestation efforts.