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Detailed osteological descriptions of the craniomandibular complex of passerine birds are lacking for most species, limiting our understanding of their diversity and evolution. Cowbirds (genus Molothrus) are a small but widespread group of New World nine-primaried songbirds, well-known for their unique brooding parasitic behavior. However, detailed osteological data for cowbirds and other Icteridae are currently scarce and several features of their skulls remain undescribed or poorly known. To address this issue, a detailed comparative osteology of cowbird skulls is presented here for the first time based on data from x-ray microcomputed tomography, dry skeletal data, and multivariate analyses of linear morphometric data. Cowbird skulls offer some functional insights, with many finch-like features probably related to a seed-rich diet that distinguishes them from most other icterids. In addition, features previously overlooked in earlier studies might provide valuable phylogenetic information at different levels of passerine phylogeny (Passerida, Emberizoidea, Icteridae, and Agelaiinae), including some of the otic region and nasal septum. Comparisons among cowbirds show that there is substantial cranial variation within the genus, with M. oryzivorus being the most divergent cowbird species. Within the genus, distantly related species share similar overall skull morphology and proportions, but detailed osteological data allow species identification even in cases of strong convergence. Further efforts are warranted to furnish baseline data for future studies of this iconic group of Neotropical birds and to fully integrate it into phylogenetic comparative frameworks.

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Generalist brood parasites that share nests with host nestlings can optimize resource acquisition from host parents by balancing the benefits that host nest-mates provide, including attracting increased provisions to the nest, against the costs of competing with the same host young over foster parental resources. However, it is unclear how parasitic chicks cope when faced with more nest-mates than are optimal for their survival upon hatching. We suggest that, in the obligate brood parasitic brown-headed cowbird (Molothrus ater), chicks use a niche construction strategy and reduce larger, more competitive host broods to maximize the parasites' survival to fledging. We experimentally altered brood sizes to test for Goldilocks principle patterns (i.e. a 'just right' intermediate brood size) of cowbird survival in nests of prothonotary warbler (Protonotaria citrea) hosts. We found that intermediate brood sizes of two host nestlings maximized cowbird fledging success relative to 0 or 4 host nest-mates at hatching. Specifically, cowbird nestlings lowered host brood sizes towards this optimum when placed in broods with more host nestlings. The results suggest that cowbirds reduce, but do not eliminate, host broods as a niche construction mechanism to improve their own probability of survival.

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Acoustic signalling is vital to courtship in many animals, yet the role of female vocalizations is understudied. Here, we combine observational and experimental methods to assess the courtship function of the female chatter call in brown-headed cowbirds, Molothrus ater. While the chatter call is likely multifunctional, it is frequently used in social interactions and overlapping duets with males during the breeding season. Based on a combination of focal- and scan-sampling data from large naturalistic aviaries, we did not find support for the hypothesis that the chatter call elicits male attention or encourages continued courtship. However, we did find evidence that the chatter call plays a role in pair bond formation, as females preferentially chattered in response to songs from pair-bond males in the 2 weeks leading up to the median date of first copulation. Females were less selective in male-directed chatter use after copulations began. We also found support for the hypothesis that chatter is used to signal-jam male songs. Frame-by-frame video analysis revealed that the majority of female chatter calls were tightly time-locked to song, occurring less than 500 ms after male vocal onset. To test the effect of signal jamming on male song potency, we designed a laboratory experiment in which male song playbacks were jammed by various recorded stimuli. Natural chatter calls more effectively reduced female copulatory responses to song than high-pass filtered chatter calls, suggesting that the low frequencies in natural chatter (2-4 kHz) are important for interfering with male song and reducing its potency. Our results suggest that sexual conflict is operating in cowbird courtship, with signal jamming serving as a mechanism by which females guard, resist or select their mates. We also discuss ways in which cowbird vocal interactions may function cooperatively to coordinate reproduction or transition females into breeding condition.

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At the core of recognition systems research are questions regarding how and when fitness-relevant decisions made. Studying egg-rejection behavior by hosts to reduce the costs of avian brood parasitism has become a productive model to assess cognitive algorithms underlying fitness-relevant decisions. Most of these studies focus on how cues and contexts affect hosts' behavioral responses to foreign eggs; however, the timing of when the cues are perceived for egg-rejection decisions is less understood. Here, we focused the responses of American robins Turdus migratorius to model eggs painted with a thermochromic paint. This technique modified an egg's color with predictably varying temperatures across incubation: at the onset of incubation, the thermochromic model egg was cold and perceptually similar to a static blue model egg (mimicking the robin's own blue-green egg color), but by the end of an incubation bout, it was warm and similar to a static beige egg (mimicking the ground color of the egg of the robin's brood parasite, the brown-headed cowbird Molothrus ater). Thermochromic eggs were rejected at statistically intermediate rates between those of the static blue (mostly accepted) and static beige (mostly rejected) model eggs. This implies that at the population level, egg-rejection relevant cues are not perceived solely when arriving to or solely when departing from the nest. We also found that robins rejected their own eggs more often when exposed to color-changing model eggs relative to static eggs, suggesting that recognizing variable foreign eggs entails costly rejection errors for this host species.

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Social regulation of reproductive hormones is a means by which conspecific males and females orchestrate successful reproductive efforts. We investigate whether social cues modify activity within the hypothalamic-pituitary-gonadal (HPG) axis and the specificity of this response in a social parasite that is known to eavesdrop on the communication signals of other species: the brown-headed cowbird (Molothrus ater). Brown-headed cowbirds are obligate brood parasites that do not build nests or care for their own young. Instead, obligate brood parasites leave their eggs in the nest of a host species and therefore must coordinate their breeding attempts with conspecifics as well as potential heterospecific hosts. Here, we explore whether the vocal signals of potential host species can also be used as a social cue that modifies the HPG axis of female brown-headed cowbirds. Results reveal that both conspecific and heterospecific song-exposed females exhibit significantly greater circulating estradiol concentrations as compared to silence-exposed females. While conspecific song induces the greatest elevation in circulating estradiol, there is no significant difference in circulating estradiol levels in females exposed to either conspecific or heterospecific songs. This pattern suggests both song types are effective at evoking a reproductive physiological response. On the other hand, circulating progesterone concentrations did not differ among the song- and silence-exposed groups nor did the size of the female's ovarian follicles. These results indicate that heterospecific vocal communication signals can effectively be used as a social cue that simultaneously provides necessary information regarding breeding status of hosts and modifies breeding condition of the eavesdropper.

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Rothstein (Behavioral Ecology and Sociobiology, 11, 1982, 229) was one of the first comprehensive studies to examine how different egg features influence egg rejection behaviors of avian brood parasite-hosts. The methods and conclusions of Rothstein (1982) laid the foundation for subsequent experimental brood parasitism studies over the past thirty years, but its results have never been evaluated with replication. Here, we partially replicated Rothstein's (1982) experiments using parallel artificial model egg treatments to simulate cowbird (Molothrus ater) parasitism in American robin (Turdus migratorius) nests. We compared our data with those of Rothstein (1982) and confirmed most of its original findings: (1) robins reject model eggs that differ from the appearance of a natural robin egg toward that of a natural cowbird egg in background color, size, and maculation; (2) rejection responses were best predicted by model egg background color; and (3) model eggs differing by two or more features from natural robin eggs were more likely to be rejected than model eggs differing by one feature alone. In contrast with Rothstein's (1982) conclusion that American robin egg recognition is not specifically tuned toward rejection of brown-headed cowbird eggs, we argue that our results and those of other recent studies of robin egg rejection suggest a discrimination bias toward rejection of cowbird eggs. Future work on egg recognition will benefit from utilizing a range of model eggs varying continuously in background color, maculation patterning, and size in combination with avian visual modeling, rather than using model eggs which vary only discretely.

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Oceanic archipelagos are vulnerable to natural introduction of parasites via migratory birds. Our aim was to characterize the geographic origins of two Plasmodium parasite lineages detected in the Galapagos Islands and in North American breeding bobolinks (Dolichonyx oryzivorus) that regularly stop in Galapagos during migration to their South American overwintering sites. We used samples from a grassland breeding bird assemblage in Nebraska, United States, and parasite DNA sequences from the Galapagos Islands, Ecuador, to compare to global data in a DNA sequence registry. Homologous DNA sequences from parasites detected in bobolinks and more sedentary birds (e.g., brown-headed cowbirds Molothrus ater, and other co-occurring bird species resident on the North American breeding grounds) were compared to those recovered in previous studies from global sites. One parasite lineage that matched between Galapagos birds and the migratory bobolink, Plasmodium lineage B, was the most common lineage detected in the global MalAvi database, matching 49 sequences from unique host/site combinations, 41 of which were of South American origin. We did not detect lineage B in brown-headed cowbirds. The other Galapagos-bobolink match, Plasmodium lineage C, was identical to two other sequences from birds sampled in California. We detected a close variant of lineage C in brown-headed cowbirds. Taken together, this pattern suggests that bobolinks became infected with lineage B on the South American end of their migratory range, and with lineage C on the North American breeding grounds. Overall, we detected more parasite lineages in bobolinks than in cowbirds. Galapagos Plasmodium had similar host breadth compared to the non-Galapagos haemosporidian lineages detected in bobolinks, brown-headed cowbirds, and other grassland species. This study highlights the utility of global haemosporidian data in the context of migratory bird-parasite connectivity. It is possible that migratory bobolinks bring parasites to the Galapagos and that these parasites originate from different biogeographic regions representing both their breeding and overwintering sites.

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Animal populations exhibit various patterns in ectoparasite distribution across different sexes and age classes, and numerous factors can potentially contribute to ectoparasite abundance and distribution. We examined the influence of host body size and age on the distribution of a chewing louse among brown-headed cowbirds. Differences in louse prevalence (males 62.9 ± 2.8%, females 47.5 ± 4.9%) and intensity (males 15.42 ± 1.51, females 9.04 ± 1.69) were primarily driven by differences in host body mass and not host sex. Larger birds had larger louse infra-populations, which likely translated into a lower risk of local extinction, a possible explanation for higher louse prevalence observed among larger birds. Among males, younger individuals showed higher louse prevalence (70.21 ± 4.72%) compared to older males (59.36 ± 3.59). We speculate that this pattern is likely driven by behavioural difference and not body size, with young males spending relatively more time foraging in large groups, increasing their risk of louse transmission. By examining the mechanisms that underlie the sex- and age-biased infections observed in natural populations, we can better identify the hosts most responsible for parasite transmission.

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The coevolutionary relationships between brood parasites and their hosts are often studied by examining the egg rejection behaviour of host species using artificial eggs. However, the traditional methods for producing artificial eggs out of plasticine, plastic, wood, or plaster-of-Paris are laborious, imprecise, and prone to human error. As an alternative, 3D printing may reduce human error, enable more precise manipulation of egg size and shape, and provide a more accurate and replicable protocol for generating artificial stimuli than traditional methods. However, the usefulness of 3D printing technology for egg rejection research remains to be tested. Here, we applied 3D printing technology to the extensively studied egg rejection behaviour of American robins, Turdus migratorius. Eggs of the robin's brood parasites, brown-headed cowbirds, Molothrus ater, vary greatly in size and shape, but it is unknown whether host egg rejection decisions differ across this gradient of natural variation. We printed artificial eggs that encompass the natural range of shapes and sizes of cowbird eggs, painted them to resemble either robin or cowbird egg colour, and used them to artificially parasitize nests of breeding wild robins. In line with previous studies, we show that robins accept mimetically coloured and reject non-mimetically coloured artificial eggs. Although we found no evidence that subtle differences in parasitic egg size or shape affect robins' rejection decisions, 3D printing will provide an opportunity for more extensive experimentation on the potential biological or evolutionary significance of size and shape variation of foreign eggs in rejection decisions. We provide a detailed protocol for generating 3D printed eggs using either personal 3D printers or commercial printing services, and highlight additional potential future applications for this technology in the study of egg rejection.

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Brown-headed cowbirds (Molothrus ater) are obligate brood parasites. Only females search for host nests and they find host nests one or more days before placing eggs in them. Past work has shown that females have a larger hippocampus than males, but sex differences in spatial cognition have not been extensively investigated. We tested cowbirds for sex and seasonal differences in spatial memory on a foraging task with an ecologically relevant retention interval. Birds were trained to find one rewarded location among 25 after 24 h. Females made significantly fewer errors than males and took more direct paths to the rewarded location than males. Females and males showed similar search times, indicating there was no sex difference in motivation. This sex difference in spatial cognition is the reverse of that observed in some polygynous mammals and is consistent with the hypothesis that spatial cognition is adaptively specialized in this brood-parasitic species.

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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.