The importance of nest‐site and habitat in egg recognition ability of potential hosts of the Common Cuckoo Cuculus canorus

The intensity of selection exerted by brood parasites on their hosts depends on the proportion of nests that are parasitized and the fitness costs of parasitism. Nest detection by brood parasites influences the probability of parasitism, and we propose that the difficulty faced by brood parasites of finding nests on the ground may make ground‐nesting species subject to lower levels of parasitism, causing a reduction in levels of defence compared with species breeding in shrubs, trees and elsewhere above the ground. We tested the prediction that the rejection rate of Common Cuckoo Cuculus canorus eggs by hosts is inversely related to the frequency with which they build nests on the ground, both at local and at continental scales. First, we used estimates of the rejection rate of non‐mimetic model eggs experimentally introduced into the nests of 26 potential host species breeding in the Sierra Nevada Mountains of southern Spain. Most species tested in the Sierra Nevada showed high rejection rates of both mimetic and non‐mimetic eggs, whereas the European Robin Erithacus rubecula, with a low rejection rate, was the only species that was regularly parasitized. At the continental scale we used all available published information on rejection rates of non‐mimetic models by European hosts of the Common Cuckoo. The frequency of ground‐nesting explained interspecific variation in rejection rate of non‐mimetic model eggs both for the species tested in the Sierra Nevada and for all European hosts after controlling for all other life‐history variables known to affect rejection rates. An effect of the abundance of trees in a particular habitat, previously shown to affect parasitism by the Common Cuckoo, was only apparent from analyses of continental‐scale data and not from the Sierra Nevada mountains, suggesting that particular properties of mountainous areas affect Common Cuckoo parasitism. Ground‐nesting species showed lower rejection rates than species breeding in bushes or trees. These results suggest that species nesting on the ground may have suffered lower parasitism pressures in their historical coevolutionary interactions with the Common Cuckoo.     

Symbiotic nesting of birds with formidable animals: a review with applications to biodiversity conservation

Increased predation and parasitism of bird nests has become a major problem in many biological communities altered by human activities, often causing declines in bird populations. To help solve this threat to biodiversity, I propose restoring the abundance of symbiotic nest-protecting animals in habitats where birds face an increased risk from predators and parasites, so that birds there can increase their chances of reproductive success by nesting close to these protectors. The re-establishment of such protective nesting associations to increase avian reproductive success differs from other proposed solutions to this problem in that it involves point defense of bird nests themselves. Rather than diminishing the number of nest predators and brood parasites in the whole habitat or community, as proposed with other approaches, the presence, activity and success of these enemies are reduced only within the microhabitat defended by the protector. The animal protecting the nest need not be larger in size than the predators or brood parasites, and is often many times smaller. In addition, it need not be from a higher trophic position, and in many cases comes from the same or a lower trophic level. Research suggests that an informed and careful use of nest protecting animals by wildlife managers can help reverse or prevent the decline of many bird populations, especially when used in combination with other approaches such as restoration of top predator populations and habitats. Although wildlife biologists have long recognized the important role that plants play in concealing and protecting bird nests from enemies, and regularly recommend manipulation of vegetation to enhance nest survival, they have generally ignored the important role that formidable animals play in protecting bird nests, and failed to incorporate animal protectors into management strategies. Because of this neglect, a host of new studies and experiments are urgently needed to provide managers with the critical information needed to use protective nesting associations effectively in integrated strategies to preserve avian biodiversity.     

Common Cuckoos Cuculus canorus change their nest‐searching strategy according to the number of available host nests

In recent decades, numerous studies have examined factors affecting risk of host nest parasitism in well‐known avian host–parasite systems; however, little attention has been paid to the role of host nest availability. In accordance with other studies, we found that nest visibility, reed density and timing of breeding predicted brood parasitism of Great Reed Warblers Acrocephalus arundinaceus by the Common Cuckoo Cuculus canorus. More interestingly, hosts had a greater chance of escaping brood parasitism if nesting was synchronized. Cuckoo nest searching was governed primarily by nest visibility at high host‐nest density. However, even well‐concealed nests were likely to be parasitized during periods when just a few hosts were laying eggs, suggesting that Cuckoos adjust their nest‐searching strategy in relation to the availability of host nests. Our results demonstrate that host vulnerability to brood parasitism varies temporally and that Cuckoo females are able to optimize their nest‐searching strategy. Moreover, our study indicated that Cuckoos always manage to find at least some nests to parasitize. Thus, in this case, the co‐evolutionary arms race should take place mainly in the form of parasitic egg rejection rather than via frontline pre‐parasitism defence.     

Colony size and nest density predict the likelihood of parasitism in the colonial Southern Red Bishop Euplectes orix– Diderick Cuckoo Chrysococcyx caprius system

Colonial nesters are potentially susceptible to brood parasitism because they present an aggregated source of accessible nests to brood parasites. However, colonial breeding may confer fitness advantages on potential host species that outweigh the costs of parasitism, particularly if colonies are large or dense enough for corporate vigilance to deter parasitism. In addition, some studies have suggested that the spatial habitat structure near host populations is a critical determinant of parasitism rate, with cuckoos targeting those colonies close to cover. We examined the determinants of the likelihood of parasitism by Diderick Cuckoos Chrysococcyx caprius on their main southern African host, the colonially breeding Southern Red Bishop Euplectes orix , from 24 colonies and 1141 nests over two seasons. The likelihood of parasitism by Diderick Cuckoos decreased significantly with increasing colony size and nest density, but not with distance to cover from a colony, providing no support for the spatial habitat structure hypothesis. We suggest that proximate constraints, such as visibility in the semi-closed nest, limit the ability of the host to recognize the Cuckoo egg, giving rise to low rates of rejection of Cuckoo eggs. However, the Cuckoo cannot exploit this apparent weakness when the host breeding population is in large colonies that are protected by corporate vigilance. Consequently, the Diderick Cuckoo – Southern Red Bishop system may represent an evolutionary equilibrium in host and parasite defences.     

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.     

Nesting aggregation as a predictor of brood parasitism in mason bees (Osmia spp.)

1. Parasitism can be an important source of mortality for insect populations; however, we know little about the factors influencing vulnerability of wild bees to parasites. Mason bees (genus Osmia; Hymenoptera: Megachilidae) are important pollinators of crops and wild plants and are vulnerable to attack by brood parasites. High nest densities may increase rates of brood parasitism by attracting disproportionate numbers of parasites. 2. Three years of field observations from multiple sites were analysed to assess whether mason bee brood parasitism increased with host density. Mason bees were allowed to nest in artificial nesting blocks and establish natural variation in nesting density. Nest cells constructed by bees were checked for the presence of parasite eggs. 3. Parasitism of nest cells strongly increased with the number of actively nesting bees at a nesting block. Mason bees showed no preference for nesting in blocks that were occupied or unoccupied by other mason bees. Parasitism also increased with the number of days a nest was provisioned and decreased over the course of the season. Nest cells constructed last in a nest were significantly more parasitised than inner cells, despite being sealed against invasions. 4. These findings show positively density‐dependent parasitism in mason bees. They also suggest that bees terminate parasitised nests, causing parasitised cells to become outermost nest cells – a behaviour that may represent a defence against parasites. Our results have implications for the management of mason bees as agricultural pollinators, as cultivating them at high densities could reduce offspring survival.     

Effects of brood parasitism on host reproductive success: evidence from larval interactions among dung beetles

This paper investigates the effect of brood parasitism in a dung beetle assemblage in an arid region of Spain. The study was conducted during the spring season (March-May 1994-1998) using mesh cylinders buried into the ground, filled with sand and with sheep dung on top. We quantified the proportion of nests containing larvae of parasitic beetles and their effect on host larvae survival. Experiments on the effect of parasitic larvae on host-larvae survival were conducted by placing scarab brood masses (raised from captive scarabs in the laboratory) in containers with and without aphodiid larvae. During the spring, dung desiccation is rapid, preventing aphodiids nesting in the dung, and forcing these species to adopt brood parasitism as a nesting strategy. Parasitic aphodiids were found in 12-47% of scarab nests of three species. The incidence of brood parasitization was positively related with the number of brood masses contained in the nests, being also higher in the most abundant species. Field data and experiments showed that brood parasites significantly reduced host larvae survival from 74.8% in non-parasitized nests to 8.8% in parasitized nests. Because different rates of nest parasitization and mortality were caused by parasites, brood parasitism had a differential effect on different host species. Thus, brood parasitism constitutes an important mortality factor reducing the reproductive success of the host species and potentially affecting the beetle abundance in the area.     

The evolution of acceptance and tolerance in hosts of avian brood parasites

Avian brood parasites lay their eggs in the nests of their hosts, which rear the parasite's progeny. The costs of parasitism have selected for the evolution of defence strategies in many host species. Most research has focused on resistance strategies, where hosts minimize the number of successful parasitism events using defences such as mobbing of adult brood parasites or rejection of parasite eggs. However, many hosts do not exhibit resistance. Here we explore why some hosts accept parasite eggs in their nests and how this is related to the virulence of the parasite. We also explore the extent to which acceptance of parasites can be explained by the evolution of tolerance; a strategy in which the host accepts the parasite but adjusts its life history or other traits to minimize the costs of parasitism. We review examples of tolerance in hosts of brood parasites (such as modifications to clutch size and multi‐broodedness), and utilize the literature on host–pathogen interactions and plant herbivory to analyse the prevalence of each type of defence (tolerance or resistance) and their evolution. We conclude that (i) the interactions between brood parasites and their hosts provide a highly tractable system for studying the evolution of tolerance, (ii) studies of host defences against brood parasites should investigate both resistance and tolerance, and (iii) tolerance and resistance can lead to contrasting evolutionary scenarios.     

Errors in egg-laying by female Common Cuckoo Cuculus canorus in nests of its common host

Dozens of studies have documented that brood parasites are well adapted to a brood parasitic lifestyle but not all parasitism events are successful. Co-evolution between brood parasites and their hosts is a dynamic process so it is reasonable to expect that a female brood parasite may commit errors during egg deposition by laying her eggs outside the laying period of the host, with consequent impacts on her fitness. Using an extensive dataset from a long-term study, we evaluated egg-laying patterns and errors related to the timing of egg-laying in the Common Cuckoo Cuculus canorus (hereafter ‘Cuckoo’). Specifically, we tested whether the Cuckoo avoids laying before or on the day of host clutch initiation to reduce the risk of rejection of parasitic eggs, whether laying errors will be more frequent in periods with a lack of active host nests, and whether the laying errors will be more frequent in periods with intense Cuckoo parasitism and a consequent lack of suitable host nests. We found that about one-third of Cuckoo eggs were laid on the host clutch initiation day or 1 day before, and the percentage of Cuckoo eggs laid decreased thereafter. Surprisingly, the probability of Cuckoo egg acceptance by the hosts was not affected by the egg-laying stage of the host clutch. Errors in the timing of egg-laying with fatal consequences (i.e. those precluding Cuckoo hatching because of laying in incubated or deserted clutches) were recorded in about 5% of cases. Only laying date of a Cuckoo egg had a significant effect on the probability of errors, which increased during the breeding season. This may be related to the higher number of deserted and incubated host nests at the site at the end of the breeding season. Errors in egg-laying may be attributed to young and inexperienced females but also impaired body condition or intraspecific competition may cause this behaviour. Future studies, which will test these possible explanations, will help to understand better the mechanism of co-evolutionary arms races and differences between host specialist and generalist brood parasites in various host–parasite systems.     

Parasites favour intermediate nestling mass and brood size in cliff swallows

A challenge of life‐history theory is to explain why animal body size does not continue to increase, given various advantages of larger size. In birds, body size of nestlings and the number of nestlings produced (brood size) have occasionally been shown to be constrained by higher predation on larger nestlings and those from larger broods. Parasites also are known to have strong effects on life‐history traits in birds, but whether parasitism can be a driver for stabilizing selection on nestling body size or brood size is unknown. We studied patterns of first‐year survival in cliff swallows (Petrochelidon pyrrhonota) in western Nebraska in relation to brood size and nestling body mass in nests under natural conditions and in those in which hematophagous ectoparasites had been removed by fumigation. Birds from parasitized nests showed highest first‐year survival at the most common, intermediate brood‐size and nestling‐mass categories, but cliff swallows from nonparasitized nests had highest survival at the heaviest nestling masses and no relationship with brood size. A survival analysis suggested stabilizing selection on brood size and nestling mass in the presence (but not in the absence) of parasites. Parasites apparently favour intermediate offspring size and number in cliff swallows and produce the observed distributions of these traits, although the mechanisms are unclear. Our results emphasize the importance of parasites in life‐history evolution.     

Breeding success of common cuckoos Cuculus canorus parasitising four sympatric species of Acrocephalus warblers

We investigated the level of parasitism, egg mimicry and breeding success of cuckoos parasitising four sympatric species of Acrocephalus warblers in southern Moravia, Czech Republic. The parasitism rate was highest in the marsh warbler Acrocephalus palustris (44.8%) followed by great reed warbler A. arundinaceus (33.8%), sedge warbler A. schoenobaenus (26.5%) and reed warbler A. scirpaceus (11.6%). Although the cuckoo eggs showed a high level of mimicry the eggs of the marsh warbler this host species rejected 72% of the cuckoo eggs, resulting in a cuckoo breeding success of only 4.3%. Cuckoo eggs laid in great reed warbler and reed warbler nests showed a similar hatching success, but the cuckoo chicks survived better in great reed warbler nests, resulting in a breeding success of 30.4%, as compared to 16.4% in nests of the reed warbler. The relationship between the level of parasitism, host rejection of cuckoo eggs, cuckoo chick survival and breeding success is discussed for the four host species.     

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.     

Mode of development and interspecific avian brood parasitism

Avian interspecific brood parasites differ considerably in theircommitment to parasitism; 87 species are obligate brood parasites,whereas 35 species are known to be facultative brood parasites.This variation is strongly related to mode of development. Obligateparasitism is found almost exclusively in altricial species,whereas facultative interspecific parasitism is predominantin precocial birds. We propose that the association betweenmode of development and form of parasitism reflects a fundamentaldifference between altricial and precocial birds in the relativebenefits of emancipation from parental care after laying. Weargue that altricial brood parasites obtain such a large increasein realized fecundity by avoiding the costs of parental carethat obligate parasitism is favored over facultative parasitism.In contrast, precocial brood parasites gain relatively littlein terms of increased fecundity via obligate parasitism, andmuch of this increase could potentially be gained by facultativeparasitism. Thus, obligate interspecific brood parasitism willnot be favored in precocial birds. Three factors influence thisdifference between altricial and precocial species: (1) altricialbirds have relatively more energy and nutrients with which tolay additional eggs, (2) altricial birds can produce more eggsfor the same amount of energy and nutrients, and (3) altricialbirds realize a greater relative gain in fecundity for eachadditional egg laid. We suggest further that facultative interspecificparasitism in birds may originate simply through a carry overof intraspecific parasitism; 29 of 33 facultative interspecificparasites also parasitize conspecifics. Facultative parasitismof other species would provide a greater range of potentialhost nests and could be maintained as an evolutionarily stableend point by the same mechanisms that maintain intraspecificbrood parasitism. [Behav Ecol 1991;2:309–318]     

A game-theoretic model of interspecific brood parasitism with sequential decisions

The interaction between hosts and parasites in bird populations has been studied extensively. This paper uses game-theoretic methods to model this interaction. This has been done in previous papers but has not been studied taking into account the detailed sequential nature of this game. We introduce a model allowing the host and parasite to make a number of decisions which will depend on various natural factors. The sequence of events begins with the host forming a nest and laying a number of eggs, followed by the possibility that a parasite bird will arrive at the nest; if it does it can choose to destroy some of the host eggs and lay one of its own. A sequence of events follows, which is broken down into two key stages; firstly the interaction between the host and the parasite adult, and secondly that between the host and the parasite chick. The final decision involves the host choosing whether to raise or abandon the chicks that are in the nest. There are certain natural parameters and probabilities which are central to these various decisions; in particular the host is generally uncertain whether parasitism has taken place, but can assess the likelihood of parasitism based upon certain cues (e.g. how many eggs remain in its nest). We then use this methodology to model two real-world interactions, that of the Reed Warbler with the Common Cuckoo and also the Yellow Warbler with the Brown-headed Cowbird. These parasites have different methods in the way they parasitize the nests of their hosts, and the hosts can in turn have different reactions to these parasites. Our model predictions generally match the real results well, and the model also makes predictions of the effect of changes in various key parameters on the type of parasitic interactions that should occur.     

Temporal patterns of host availability, brown-headed cowbird brood parasitism, and parasite egg mass

I studied relationships between temporal patterns of host availability, brood parasitism, and egg mass for the parasitic brown-headed cowbird (Molothrus ater). At a study site consisting largely of edge habitat in north-eastern Illinois, I found 834 bird nests from 27 species. A total of 407 cowbird eggs and nestlings were found in these nests over three laying seasons. Nearly all (n= 379; 93%) were found in the nests of seven host species. For these species and all taken together, weekly nest availability generally decreased whereas parasitism frequency generally increased throughout the cowbird laying season, but the proportions of nests parasitized and the mean number of cowbird eggs in them did not. Additionally, no correlation was found between the proportions of nests parasitized and nest availability. Cowbird egg mass generally increased throughout the laying season, indicating that foraging conditions improved and that, early in the laying season, egg mass and quality may be less important than quantity. Consistently high weekly levels of parasitism indicate that cowbird reproduction was less limited by resources needed for egg production and more by the availability of suitable host nests. Fluctuating weekly host availabilities suggest that previously established, constant rates of cowbird egg laying would produce an excess of eggs during periods of low host availability. Further, the low frequency of parasitism (1%) of nests in stages too advanced for successful parasitism, and of abandoned nests, is consistent with the hypothesis that cowbirds' consistently high rate of egg production helps assure an egg is available when an appropriate nest is found. Frequently, nests were parasitized multiple times, raising the possibility that cowbirds were interfering with their own reproduction. A diverse host community increases the possibility that a decline of any one host species is unlikely to affect cowbird reproduction significantly. Received 11 July 1997 / Accepted: 31 March 1998     

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.     

Seasonal and diurnal patterns of population vocal activity in avian brood parasites

Patterns of vocal activity may involve information about vocalizations themselves as well as their function. In birds, vocal activity at the individual and population level is generally closely associated with breeding cycles, reaching the peak during territorial and mating competition, and decreasing with the onset of egg incubation and chick feeding. However, little is known about patterns of vocal activity in avian brood parasites that have unusual breeding cycles without parental care. Using passive acoustic monitoring, we determined the seasonal and diurnal patterns of population vocal activity in two avian brood parasites: the Common Cuckoo Cuculus canorus and the Lesser Cuckoo C. poliocephalus. We found that both species and both sexes showed a similarly highly structured pattern of seasonal vocal activity, reaching a sharp peak in the early breeding season when birds compete for territories and mates, although males sang more frequently than females. Likewise, the diurnal patterns of vocal activity were similar in both species and both sexes of cuckoos, with peak activity occurring around dawn. Nocturnal calls by male cuckoos were also detected in both species, but only in the early breeding season. Collectively, the observed patterns of population vocal activity may suggest that the absence of parental care may not extend the period of vocal activity in these two species of brood parasites.     

The effect of Jacobin Cuckoo Clamator jacobinus parasitism on the body mass and survival of young in a new host species

The southern African subspecies of Jacobin Cuckoo Clamator jacobinus serratus is a brood parasite of a range of host species. While Jacobin Cuckoos do not evict host young, previous research has found that host young rarely survive the nestling period. Here we provide the first records of Jacobin Cuckoo parasitism of a new host species, the Southern Pied Babbler Turdoides bicolor. We investigate rates of brood parasitism and the survival of host young. The Southern Pied Babbler is one of the largest recorded hosts for Jacobin Cuckoos and, unusually, we find that host young tend to survive the nestling period and maintain similar body mass to host young in unparasitized broods. However, host young were less likely to survive to independence than young raised in unparasitized nests, suggesting a post‐fledging reproductive cost to hosts.     

Sex-specific defence behaviour against brood parasitism in a host with female-only incubation

Nest protection against intruders is an indispensable component of avian parental care. In species with biparental care, both mates should evolve nest defence behaviour to increase their reproductive success. In most host-parasite systems, host females are predicted to have more important roles in nest defence against brood parasites, because they typically are primarily responsible for clutch incubation. Male antiparasitic behaviour, on the other hand, is often underestimated or even not considered at all. Here we investigated sex-specific roles in four aspects of great reed warbler (Acrocephalus arundinaceus) nest defence against a brood parasite—the cuckoo (Cuculus canorus), namely (1) mobbing, (2) nest attendance/guarding, (3) nest checking and (4) egg ejection. Using dummy experiments, simulating brood parasitism and by video-monitoring of host nests we found that males took the key roles in cuckoo mobbing and nest guarding, while females were responsible for nest checking and egg ejection behaviours. Such partitioning of parental roles may provide a comprehensive clutch protection against brood parasitism.