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The phenomenon of cuckoos' brood parasitism is well known and can be investigated using applied mathematical techniques. Among adaptive features of this phenomenon are certain egg parameters that ensure their shortened incubation period (I) and thus the successful survival of their offspring. In particular, the volume of a cuckoo egg is not less than, or exceeds, that of the host species, which should, in theory, increase I. Also, cuckoo eggs have thicker shell than that of nest hosts. Here, we analyzed the available geometric dimensions of eggs in 447 species and found an inverse correlation (-0.585, p < 0.05) between I and the shell thickness-to-egg surface area ratio (T/S). A mathematical relationship was derived to calculate I depending on T/S. This premise was confirmed by comparative calculations using egg images of two parasitic species, common (Cuculus canorus) and plaintive cuckoo (Cacomantis merulinus) and their hosts: great reed warbler (Acrocephalus arundinaceus), European robin (Erithacus rubecula), rufescent prinia (Prinia rufescens), and common tailorbird (Orthotomus sutorius). An average calculated I value for cuckoo eggs was one day less than that for host eggs. Our findings unravel additional details of how cuckoos adapt to brood parasitism and specific host-parasite relationships.

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The shiny cowbird Molothrus bonariensis parasitizes many species with different life-history traits and has a detrimental effect on the survival of the progeny of the hosts. In response, hosts have developed numerous antiparasitic defenses. Here, we examined the effects of brood parasitism by shiny cowbird on the clutch and brood sizes (83 nests) in a small host, the black-backed water tyrant Fluvicola albiventer. We also studied whether the death of parasite nestlings was related to the care of the foster parents and whether the host had any antiparasitic defense against the shiny cowbird. Our results indicate that brood parasitism significantly decreased the host hatching and fledging successes. The majority of nest failures (57%) were caused by brood parasitism. Shiny cowbird parasitism occurred in 52% of nests and the intensity of parasitism was 1.23 ± 0.53 eggs per parasitized nest. Of the total host eggs, 54% were damaged. During the incubation stage, 20 nests (47%) were abandoned because of egg punctures by shiny cowbirds females. Only two parasitic fledglings were recorded, while the remaining nestlings either died from starvation (n = 12) or predation (n = 3). Foster parents abandoned parasitic nestlings between 5 and 10 days old. Our findings demonstrate that the shiny cowbird has very low rates of fledging success when parasitizing black-backed water tyrant. Also, parasitism had a high reproductive cost in the black-backed water tyrant because a very low proportion (7%) of the parasitized nests (n = 43) were successful.

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BACKGROUND: Animal bacterial symbionts are established early in life, either through vertical transmission and/or by horizontal transmission from both the physical and the social environment, such as direct contact with con- or heterospecifics. The social environment particularly can influence the acquisition of both mutualistic and pathogenic bacteria, with consequences for the stability of symbiotic communities. However, segregating the effects of the shared physical environment from those of the social interactions is challenging, limiting our current knowledge on the role of the social environment in structuring bacterial communities in wild animals. Here, we take advantage of the avian brood-parasite system of Eurasian magpies (Pica pica) and great spotted cuckoos (Clamator glandarius) to explore how the interspecific social environment (magpie nestlings developing with or without heterospecifics) affects bacterial communities on uropygial gland skin. RESULTS: We demonstrated interspecific differences in bacterial community compositions in members of the two species when growing up in monospecific nests. However, the bacterial community of magpies in heterospecific nests was richer, more diverse, and more similar to their cuckoo nest-mates than when growing up in monospecific nests. These patterns were alike for the subset of microbes that could be considered core, but when looking at the subset of potentially pathogenic bacterial genera, cuckoo presence reduced the relative abundance of potentially pathogenic bacterial genera on magpies. CONCLUSIONS: Our findings highlight the role of social interactions in shaping the assembly of the avian skin bacterial communities during the nestling period, as exemplified in a brood parasite-host system.

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The Diederik cuckoo, Chrysococcyx caprius, is a small Afrotropical bird in the family Cuculidae. It is taxonomically related to 13 other species within the genus Chrysococcyx and is migratory in sub-Saharan Africa. It has a unique breeding behaviour of being a brood parasite: Breeding pairs lay their eggs in the nests of a host species and hatchlings expel the eggs of the host species. The aim of the present study was to investigate diversity in two circadian clock genes, Clock and Adcyap1, to probe for a relationship between genetic polymorphisms and their role in circannual timing and habitat selection (phenology) in intra-African migrants. DNA extracted from blood was used for the PCR amplification and sequencing of clock genes in 30 Diederik cuckoos. Three alleles were detected for Clock with similar genotypes between individuals from the Northern and Southern breeding ranges while 10 alleles were detected for Adcyap1, having shorter alleles in the North and longer alleles in the South. Population genetic analyses, including allele frequency and zygosity analysis, showed distinctly higher frequencies for the most abundant Clock allele, containing 10 polyglutamine repeats, as well as a high degree of homozygosity. In contrast, all individuals were heterozygous for Adcyap1 and alleles from both regions showed distinct differences in abundance. Comparisons between both clock genes and phenology found several phenotypic correlations. This included evidence of a relationship between the shorter alleles and habitat selection as well as a relationship between longer alleles and timing. In both instances, evidence is provided that these effects may be sex-specific. Given that these genes drive some of the synchronicity between environments and the life cycles of birds, they provide valuable insight into the fitness of species facing global challenges including climate change, urbanisation and expanding agricultural practices.

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Social monogamy is the prevalent mating system in birds, but alternative strategies of extra-pair paternity (EPP) and conspecific brood parasitism (CBP) occur in many species. Raptors are virtually absent in discussions of broad taxonomic reviews regarding EPP and CBP likely because these strategies are mostly absent or at low frequency; CBP is unreported in solitary nesting raptors. In contrast, we found high frequencies of EPP (16%-31%) and CBP (15%-26%) nests among three populations of Cooper's Hawks (Accipiter cooperii) across the northern breeding range of this solitary nesting, socially monogamous species. EPP and CBP combined occurred in 42%-46% of all nests among populations and hence unexpectedly were nearly equivalent to proportions of genetically monogamous nests. Select covariates failed to predict presence of EPP and CBP in part because virtually all extra-pair adults were uncaught and likely were floaters. We found no support for the hypothesis that territorial females traded copulations for food to maximize energy intake for increased production. Our unique discoveries enhance knowledge of the extent and diversity of alternative breeding strategies among groups of avian and other animal species.

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Four new inquiline social parasites are described in the dolichoderine ant genus Tapinoma from the Nearctic region, and keys are provided for queens and males of the Nearctic Tapinoma species. The new social parasite species represent the first inquiline species in the genus Tapinoma and the first confirmed inquilines known from the ant subfamily Dolichoderinae. The four new species appear to be workerless inquilines that exploit a single host, Tapinomasessile (Say), and they represent at least two distinct life history syndromes. Tapinomaincognitum Cover & Rabeling, sp. nov. is highly derived morphologically and is a host-queen-tolerant inquiline. In contrast, T.inflatiscapus Cover & Rabeling, sp. nov. shows a lesser degree of morphological modification and appears to be a host-queen-intolerant social parasite. The life history of T.pulchellum Cover & Rabeling, sp. nov. is presently unknown, but its close similarity to T.incognitum suggests that it is also a host-queen-tolerant inquiline. The life history of T.shattucki Cover & Rabeling, sp. nov. is still uncertain. Our findings provide novel insights into the complex biology of ant inquiline life history syndromes.

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The existence of adult sexual dimorphism is typically explained as a consequence of sexual selection, yet coevolutionary drivers of sexual dimorphism frequently remain untested. Here, I investigate the role of sexual dimorphism in host-parasite interactions of the brood parasitic diederik cuckoo, Chrysococcyx caprius. Female diederik cuckoos are more cryptic in appearance and pose a threat to the clutch, while male diederik cuckoos are conspicuous and not a direct threat. Specifically, I examine whether sexual dimorphism in diederik cuckoos provokes threat-level sensitive responses in Southern red bishop, Euplectes orix, hosts. I use experimentally simulated nest intrusions to test whether hosts have the capacity to differentially (i) detect, and/or (ii) discriminate between, male and female diederik cuckoos, relative to harmless controls. Overall, I found no evidence that diederik cuckoos differ in detectability, since both sexes are comparable to controls in the probability and speed of host detection. Furthermore, neither male nor female hosts discriminate between sexually dimorphic diederik cuckoos when engaging in frontline nest defences. However, hosts that witnessed a male diederik cuckoo during the trial were more likely to reject odd eggs. Moreover, experimental eggs were significantly more likely to be rejected when female bishops observed a male compared to a female diederik cuckoo. While the cryptic appearance of female diederik cuckoos does not reduce detection by hosts, it does provide the benefit of anonymity given the egg rejection costs of conspicuous male-like appearance in the nest vicinity. These findings have implications for the evolution and maintenance of sexual dimorphism across the Cuculidae, and highlight the value of testing assumptions about the ecological drivers of sexual dimorphism.

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In predator-prey interactions, the prey faces extreme challenges from predation, which drives the evolution of defense or anti-predator mechanisms. Compared with adult birds, nestlings are more vulnerable but not helpless. However, data on whether nestlings eavesdrop on the danger signals transmitted by other prey nestlings and the mechanisms of eavesdropping remain limited. In brood parasitism, common cuckoo (Cuculus canorus) nestlings, raised by host adults who are not closely related, offer an instructive system for studying the transmission and recognition of danger signals among nestlings of different species that share special relationships. We played back the distress calls of common cuckoo nestlings to nestlings of three sympatric host species (the oriental reed warbler Acrocephalus orientalis, which is a primary host of the common cuckoo, the reed parrotbill Paradoxornis heudei, an occasional host, and the vinous-throated parrotbill Sinosuthora webbiana, which is not parasitized in the study area) to investigate whether the host nestlings reduced their begging behavior. We also quantified the degree of inhibition toward begging behavior for these nestlings. The results revealed that, in response to the distress calls, the three sympatric host species markedly suppressed their begging behavior. This response can likely be attributed to the innate response of host nestlings caused by the general characteristics of distress calls, rather than the acoustic similarity and phylogenetic relationship between host nestlings and cuckoo nestlings. Furthermore, we observed that upon hearing the distress calls of cuckoo nestlings, the oriental reed warbler nestlings exhibited the greatest reduction in the total number of calls compared to the other two host species, potentially owing to stronger predation and parasitic pressures. This study suggests that host nestlings can detect danger signals emitted by parasitic nestlings; however, further investigation is needed to determine whether they can respond to distress calls from unfamiliar nestlings in different regions.

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Cleptoparasitism, also known as brood parasitism, is a widespread strategy among bee species in which the parasite lays eggs into the nests of the host species. Even though this behavior has significant ecological implications for the dynamics of several species, little is known about the molecular pathways associated with cleptoparasitism. To shed some light on this issue, we used gene expression data to perform a comparative analysis between two solitary neotropical bees: Coelioxoides waltheriae, an obligate parasite, and their specific host Tetrapedia diversipes. We found that ortholog genes involved in signal transduction, sensory perception, learning, and memory formation were differentially expressed between the cleptoparasite and the host. We hypothesize that these genes and their associated molecular pathways are engaged in cleptoparasitism-related processes and, hence, are appealing subjects for further investigation into functional and evolutionary aspects of cleptoparasitism in bees.

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When host nests are scarce, avian brood parasites would benefit from behaviours that increase the availability of suitable nests. Several studies reported ejection of host nestlings from nests by brood parasites; however, whether brood parasites do so to induce the host to re-nest and thus increase opportunities for future parasitism (i.e. 'farming' behaviour) remains unclear. Here, we report observational evidence of farming behaviour by a common cuckoo Cuculus canorus female in a Daurian redstart Phoenicurus auroreus population: (1) the cuckoo destroyed a host nest by ejecting all nestlings, (2) the host then produced a new nest and (3) the cuckoo successfully parasitized the replacement nest. We suggest that farming behaviour may be more common, but often goes undetected because it requires intense nest monitoring.

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The role of species interactions, as well as genetic and environmental factors, all likely contribute to the composition and structure of the gut microbiome; however, disentangling these independent factors under field conditions represents a challenge for a functional understanding of gut microbial ecology. Avian brood parasites provide unique opportunities to investigate these questions, as brood parasitism results in parasite and host nestlings being raised in the same nest, by the same parents. Here we utilized obligate brood parasite brown-headed cowbird nestlings (BHCO; Molothrus ater) raised by several different host passerine species to better understand, via 16S rRNA sequencing, the microbial ecology of brood parasitism. First, we compared faecal microbial communities of prothonotary warbler nestlings (PROW; Protonotaria citrea) that were either parasitized or non-parasitized by BHCO and communities among BHCO nestlings from PROW nests. We found that parasitism by BHCO significantly altered both the community membership and community structure of the PROW nestling microbiota, perhaps due to the stressful nest environment generated by brood parasitism. In a second dataset, we compared faecal microbiotas from BHCO nestlings raised by six different host passerine species. Here, we found that the microbiota of BHCO nestlings was significantly influenced by the parental host species and the presence of an inter-specific nestmate. Thus, early rearing environment is important in determining the microbiota of brood parasite nestlings and their companion nestlings. Future work may aim to understand the functional effects of this microbiota variability on nestling performance and fitness.

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Egg recognition and rejection are the most common and effective anti-parasitic strategies against avian brood parasitism in terms of maintaining stability over time and plasticity in response to environmental cues. Conversely, parasites have evolved multiple counter-adaptations to the anti-parasitic defenses of hosts. Among them, the crypsis hypothesis suggests that eggs that appear more cryptic in color and are closely matched to the environment are helping to counter the egg recognition strategy of the host. In this study, we investigated whether the egg recognition ability of Oriental reed warblers (Acrocephalus orientalis), a common host of common cuckoos (Cuculus canorus), changed during different reproductive stages by using model egg experiments. The effect of the crypsis hypothesis on the egg recognition ability of the hosts was also investigated by controlling the color contrast between the inside of the experimental nests and the model eggs. The results showed that the Oriental reed warbler retained strong egg recognition abilities, which were similar to the incubation stage (GLMMs: F 1,27 = 0.424, p = .521), even after entering the nestling stage and preferentially rejected model eggs with distinct contrasting colors (binomial test: Fisher's exact, p = .016). These results are consistent with the crypsis hypothesis. The present study suggests that the host retains a strong egg recognition ability even during the nestling stage and that cryptic-colored eggs that are closely matched with the breeding nest environment help counter the host's egg recognition abilities and increase the chances of successful parasitism by cuckoos. However, the effectiveness of cryptic egg may be weaker than mimic egg in countering egg recognition and rejection by hosts with open-cup nests.

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Egg rejection is an effective and widespread antiparasitic defense to eliminate foreign eggs from the nests of hosts of brood parasitic birds. Several lines of observational and critical experimental evidence support a role for learning by hosts in the recognition of parasitic versus own eggs; specifically, individual hosts that have had prior or current experience with brood parasitism are more likely to reject foreign eggs. Here we confirm experimentally the role of prior experience in altering subsequent egg-rejection decisions in the American robin Turdus migratorius, a free-living host species of an obligate brood parasite, the brown-headed cowbird Molothrus ater. We then model the coevolutionary trajectory of both the extent of mimicry of host eggs by parasitic eggs and the host's egg rejection thresholds in response to an increasing role of learning in egg recognition. Critically, with more learning, we see the evolution of both narrower (more discriminating) rejection thresholds in hosts and greater egg mimicry in parasites. Increasing host clutch size (number of eggs/nest) and increasing parasite load (parasitism rate) also have narrowing effects on the egg-rejection threshold. Together, these results suggest that learning from prior experience with egg rejection may play an important role in the coevolution of egg-mimetic lineages of brood parasites and the refined egg rejection defenses of hosts.

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Brood parasitism by cuckoos relies on manipulating hosts to raise their offspring and has evolved stunning adaptations to aid in their deception. The fact that cuckoos usually but not always, remove one or two host eggs while laying their eggs has been a longstanding focus of intensive research. However, the benefit of this behavior remains elusive. Moreover, the recently proposed help delivery hypothesis, predicting that egg removal by cuckoos may decrease the egg-laying duration in the parasitism process caused by biting action, lacks experimental verification. Therefore, in this study, we examined the effects of egg removal/biting on the egg-laying speed in the common cuckoo (Cuculus canorus) to experimentally test this hypothesis. We compared the duration of cuckoo egg-laying in empty nests, nests with host eggs, and nests with artificial blue stick models to test whether cuckoos biting an egg/stick can significantly hasten the egg-laying speed than no biting action. Our results showed that biting an egg or an object is associated with cuckoos laying approximately 37% faster than when they do not bite an egg or an object. This study provides the first experimental evidence for the help delivery hypothesis and demonstrates that when cuckoos bite eggs or other objects in the nest, they lay eggs more quickly and thereby avoid suffering the hosts' injurious attack.

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Avian brood parasitism is a model system for understanding coevolutionary arms races, and the great reed warbler (Acrocephalus arundinaceus, hereafter 'warbler') and its parasite the common cuckoo (Cuculus canorus, hereafter 'cuckoo') are prime examples of this coevolutionary struggle. Here, warblers select for egg colour mimicry by rejecting poorly matched cuckoo eggs. Contrary to long-held assumptions, recent work showed that warblers tend to reject lighter and browner eggs but tended to accept darker and bluer eggs rather than basing rejection decisions solely on perceived colour differences (i.e. the degree of mimicry). This counterintuitive, colour-biased rejection behaviour would select for bluer and darker cuckoo eggs, but would only be adaptive if cuckoos were consistently lighter and browner than warbler eggs. Therefore, we tested whether warbler eggs were consistently bluer and darker than cuckoo eggs. To do so, we re-analysed eggshell reflectance spectra of warblers and the cuckoos that parasitized them in the Czech Republic. As expected, we found that warbler eggs were significantly bluer and darker than the cuckoo eggs at the population level. Thus, we demonstrate imperfect mimicry in a long-coevolved cuckoo host-race and provide insights for exploring the coevolutionary interactions among hosts and their brood parasites.

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AbstractHosts and brood parasites are a classic example of conflict. Parasites typically provide no offspring care after laying eggs, imposing costs on hosts. Female subsocial wasps (Ammophila pubescens) alternate between initiating their own nests and using an "intruder" tactic of replacing eggs in nests of unrelated conspecifics. Hosts can respond by substituting new eggs of their own, with up to eight reciprocal replacements. Remarkably, intruders usually provision offspring in host nests, often alongside hosts. We used field data to investigate why intruders provision and to understand the basis of interactions. We found that intruders could not increase their fitness payoffs by using the typical brood parasite tactic of not provisioning offspring. Intruders using the typical tactic would benefit when hosts provisioned in their stead, but their offspring would starve when hosts failed to provision. Although some hosts obtained positive payoffs when intruders mistakenly provisioned their offspring, on average utilizing a conspecific nest represents parasitism: hosts pay costs while intruders benefit. Hosts and intruders used the same tactic of egg replacement, but intruders more often laid the final egg. Selection should favor better discrimination of offspring, which could lead to repeated cycles of costly egg replacement.

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Comparative studies of egg recognition and rejection between various sympatric hosts provide insight into the coevolutionary history of the hosts and parasites, as well as the degree of antagonism between the species. Although buntings are widely considered to be a suitable host taxon for cuckoos, there has been relatively little research on this example of parasitism and host antiparasitic behaviour. Here we provided the first report on brood parasitism and egg recognition in three sympatric ground-nesting bunting hosts of the common cuckoo (Cuculus canorus), namely the yellow-throated bunting (Emberiza elegans), south rock bunting (E. yunnanensis), and crested bunting (E. lathami). The results show that for the five breeding seasons during 2018-2022, the parasitism rate by common cuckoos was 0.87% and 0.45% in yellow-throated buntings and south rock buntings, respectively, whereas the parasitism rate by an unidentified parasite was 4% during 2018-2023 in the crested bunting. The rejection rates of the three bunting hosts for blue non-mimetic eggs were 89.3%, 88.9%, and 100% for yellow-throated buntings, south rock buntings, and crested buntings, respectively. The rejection rates for red non-mimetic eggs by yellow-throated buntings and south rock buntings were lower at 76.9% and 82.4%, respectively. All three sympatric bunting hosts examined had high levels of egg recognition and egg rejection, suggesting that it may have been subjected to high parasitic history and that egg recognition ability was retained after the loss of parasitism, which needs to be further verified by future experiments.

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Host-parasite dynamics involve coevolutionary arms races, which may lead to host specialization and ensuing diversification. Our general understanding of the evolution of host specialization in brood parasites is compromised by a restricted focus on bird and insect lineages. The cuckoo catfish (Synodontis multipunctatus) is an obligate parasite of parental care of mouthbrooding cichlids in Lake Tanganyika. Given the ecological and taxonomic diversity of mouthbrooding cichlids in the lake, we hypothesized the existence of sympatric host-specific lineages in the cuckoo catfish. In a sample of 779 broods from 20 cichlid species, we found four species parasitized by cuckoo catfish (with prevalence of parasitism of 2%-18%). All parasitized cichlids were from the tribe Tropheini, maternal mouthbrooders that spawn over a substrate (rather than in open water). Phylogenetic analysis based on genomic (ddRAD sequencing) and mitochondrial (Dloop) data from cuckoo catfish embryos showed an absence of host-specific lineages. This was corroborated by analyses of genetic structure and co-ancestry matrix. Within host species, parasitism was not associated with any individual characteristic we recorded (parent size, water depth), but was costly as parasitized parents carried smaller clutches of their own offspring. We conclude that the cuckoo catfish is an intermediate generalist and discuss costs, benefits and constraints of host specialization in this species and brood parasites in general.

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Co-parasitism is ubiquitous and has important consequences for the ecology and evolution of wild host populations. Studies of parasite co-infections remain limited in scope, with few experimental tests of the fitness consequences of multiple parasites, especially in natural populations. We measured the separate and combined effects of Philornis seguyi nest flies and shiny cowbirds Molothrus bonariensis on the fitness of a shared host, the chalk-browed mockingbird (Mimus saturninus) in Argentina. Using a two-factor experimental approach, we manipulated the presence of nest flies and cowbirds in mockingbird nests and assessed their effects on mockingbird haemoglobin levels, begging and provisioning rates, body size, and fledging success. We also monitored rates of nest predation in relation to parasitism by flies and cowbirds. Nest flies reduced the haemoglobin concentration, body size, and fledging success of mockingbirds, likely because mockingbirds did not compensate for parasitism by begging more or feeding their nestlings more. Cowbirds also reduced the fledging success of mockingbirds, even though they had no detectable effect on haemoglobin or body size. Nests with cowbirds, which beg more than mockingbirds, attracted more nest predators. There was no significant interaction between the effects of flies and cowbirds on any component of mockingbird fitness. The combined effects of nest flies and cowbirds were strictly additive. In summary, we show that nest flies and cowbirds both reduce host fitness, but do not have interactive effects in co-parasitized nests. Our results further suggest that predators exacerbate the effects of nest flies and cowbirds on their hosts. Our study shows that the fitness consequences of co-parasitism are complex, especially in the context of community-level interactions.

El co-parasitismo es ubicuo y tiene consecuencias importantes para la ecología y la evolución de las poblaciones de hospedadores silvestres. Los estudios de coinfecciones por parásitos poseen un alcance limitado, con pocos trabajos experimentales que analicen las consecuencias del parasitismo múltiple sobre la eficacia biológica, especialmente en poblaciones naturales. Medimos los efectos separados y combinados de las moscas de los nidos Philornis seguyi y el tordo renegrido (Molothrus bonariensis) sobre la eficacia biológica de un hospedador compartido, la calandria grande (Mimus saturninus) en Argentina. Utilizando un enfoque experimental de dos factores, manipulamos la presencia de moscas de nido y tordos en los nidos de calandrias y evaluamos sus efectos sobre los niveles de hemoglobina, las tasas de solicitud de alimento y aprovisionamiento, el tamaño corporal y el éxito de supervivencia de los pichones de calandria. También monitoreamos las tasas de depredación de nidos en relación con el parasitismo de moscas y tordos. Las moscas de los nidos redujeron la concentración de hemoglobina, el tamaño corporal y el éxito de supervivencia de los pichones de calandrias, probablemente porque las calandrias no compensaron el parasitismo solicitando más alimento o alimentando más a sus pichones. Los tordos también redujeron el éxito de supervivencia de las calandrias, aunque no tuvieron un efecto detectable sobre la hemoglobina o el tamaño corporal. Los nidos con tordos, quienes solicitaron más alimento más que las calandrias, atrajeron a más depredadores de nidos. El aumento de la depredación de nidos asociado a la alta intensidad de solicitud de alimento puede explicar por qué las calandrias en Argentina no solicitan más alimento en respuesta al parasitismo de las moscas. No hubo una interacción significativa entre los efectos de las moscas y los tordos en ningún componente de la eficacia biológica de las calandrias. Los efectos combinados de las moscas de los nidos y los tordos fueron estrictamente aditivos. En resumen, mostramos que las moscas de los nidos y los tordos reducen la eficacia biológica del hospedador, pero no tienen efectos interactivos en los nidos co-parasitados. Nuestros resultados sugieren además que los depredadores exacerban los efectos de las moscas de los nidos y los tordos sobre sus hospedadores. Nuestro estudio muestra que las consecuencias del co-parasitismo sobre la eficacia biológica son complejas, especialmente en el contexto de las interacciones a nivel comunitario.

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Most mimicry systems involve imperfect mimicry, whereas perfect and high-fidelity mimicry are rare. When the fidelity of mimicry is high, mimics might be expected to have the upper hand against their antagonists. However, in coevolving systems, diversification of model phenotypes may provide an evolutionary escape, because mimics cannot simultaneously match all model individuals in the population. Here we investigate high-fidelity mimicry in a highly specialized, Afrotropical brood parasite-host system: the African cuckoo and fork-tailed drongo. Specifically, we test whether host egg polymorphisms are an effective defence against such mimicry. We show, using a combination of image analysis, field experiments and simulations, that: (1) egg colour and pattern mimicry of fork-tailed drongo eggs by African cuckoos is near-perfect on average; (2) drongos show fine-tuned rejection of foreign eggs, exploiting unpredictable pattern differences between parasitic eggs and their own; and (3) the high degree of interclutch variation (polymorphic egg 'signatures') exhibited by drongos gives them the upper hand in the arms race, with 93.7% of cuckoo eggs predicted to be rejected, despite cuckoos mimicking the full range of drongo egg phenotypes. These results demonstrate that model diversification is a highly effective defence against mimics, even when mimicry is highly accurate.

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