Intraspecific nest parasitism in the grey starling (Sturnus cineraceus)

We studied intraspecific nest parasitism in the grey starling (Sturnus cineraceus) in 1992 and 1993. We used three criteria to detect nest parasitism: (i) the appearance of more than one egg per day while the host was laying; (ii) the appearance of extra eggs after the host completed its clutch; and (iii) the appearance of eggs which were of a different shape, size and color to other eggs in the clutch. There were 290 nests (157 nests in 1992; 133 nests in 1993) in which the clutch was completed early (clutches initiated before May 10). Twenty-nine (1992) and 32 (1993) nests contained at least one parasitic egg. Parasitic eggs hatched if they were laid during the laying period and early in the incubation period of their host, and a few of them fledged. Fledging success of parasitic eggs was not different from that of eggs in non-parasitized nests if parasitic eggs were laid during the host's laying period. However, fledging success of all parasitic eggs was fewer than that of eggs in non-parasitized nests. By comparison, fledging success of parasitized nests was not a great as that of non-parasitized nests.     

Relative reproductive success of female moorhens using conditional strategies of brood parasitism and parental care

In a population of moorhens (Gallinula chloropus), at least27% of netting females laid one or more eggs in a neighbor'snest Females laid parasitically under three conditions: 56%of parasitic eggs were from nesting females that preceded layinga dutch in their own nest by a parasitic laying bout, 19% werefrom females whose nests were depredated before clutch completionand that laid the following egg parasiticaDy, and 25% were froma small number of females without territories, "non-nesting"parasites, that each laid a series of parasitic eggs. Clutchsizes varied greatly between females, but nesting females eachlaid a consistent clutch size both within and between seasonsfor a given mate and territory. Nesting females that employeda dual strategy of brood parasitism and parental care producedextra eggs that they laid in the nests of neighbors before layinga dutch in their own nests. Two out of ten females whose dutchesI experimentally removed during the laying period were successfullyinduced to lay their next egg in the nest of a neighbor. Nestingfemales that laid parasitically selected their hosts opportunisticallyfrom among the nests dosest to their territories. An experimentin which parasitic eggs were removed and hosts left to rearonly their own young showed that parasites did not choose hoststhat were better parents than pairs with contemporary neststhat were not parasitized. Females that only laid parasiticaDywithin a given season timed their parasitic laying bouts poorlyand achieved no reproductive success. Parasitic young rarelyfledged, and the mean seasonal reproductive success of nestingbrood parasites did not differ from that of nonparasitic females.However, the variance in reproductive success of nesting broodparasites was significantly higher than that of nonparasiticfemales.     

Small hosts infrequently disrupt laying by Brown-headed Cowbirds and Bronzed Cowbirds

ABSTRACT Brood parasites often must overcome host defenses that may include behaviors that serve other functions, such as deterrence of predators and nest attendance during laying and incubation. Host use by brood parasites may also be influenced by competitors in areas where more than one parasitic species occurs. We identified the degree to which behavior of potential hosts and potential competitors affected laying by Brown‐headed Cowbirds (Molothrus ater) and Bronzed Cowbirds (M. aeneus) at a site in south Texas where they co‐occur. We watched potential host nests during the presunrise period to record cowbird laying and document nest visitation, laying, cowbird‐host encounters, and nest attentiveness by hosts. Hosts were frequently at their nests when cowbirds laid eggs (83% of 121 watches among nests of five host species) and cowbirds regularly encountered hosts (43–74% and 40–77% of watches per species of host for Brown‐headed and Bronzed cowbirds, respectively). Host nest defense infrequently interfered with cowbird laying and cowbirds rarely interacted with one another during laying. Overall, 12% of the 42 cowbird laying attempts that elicited host nest defense failed, resulting in cowbird eggs either laid atop hosts as they sat in nests or laid outside the nest cup. We clearly documented that relatively small hosts can thwart parasitism by cowbirds. Thus, the potential for successful defense of nests should be considered when assessing the evolution of behaviors to deter the removal of host eggs by cowbirds and mechanisms leading to nest abandonment. Regarding the latter, the presence of a cowbird at a nest would be a poor indicator for parasitism as some laying attempts were thwarted and unparasitized broods were reared at those nests. Despite the potential for nest defense to affect host use by cowbirds, we did not detect an effect of nest defense. Because most host defense was ineffective, we examined hypotheses for the timing of cowbird laying and host nest attendance. Our analysis of time of day of laying by Brown‐headed Cowbirds at our site and data compiled from the literature suggests that laying time is best predicted by the time of civil twilight (first light) rather than sunrise.     

Host-parasite relatedness in wood ducks: patterns of kinship and parasite success

We investigated the role of kinship in intraspecific nest parasitismof wood ducks (Aix sponsa). Among waterfowl, female philopatrycreates the potential for female relatives to nest in proximity.Costs of intraspecific nest parasitism to host females may bereduced if parasites lay eggs with kin. However, previous observationsof marked wood ducks indicated that females avoided parasitizingclutch mates or the female that incubated them. To further examinethe role of kinship, we determined the genotypes of 27 host-parasitepairs at five microsatellite loci. Average relatedness betweenhosts and all females laying parasitic eggs was only 0.04 ±0.03. Parasites appeared to choose hosts randomly with respectto kinship from among females with nests in the neighborhoodand those within the entire study area. However, host relatednessto the parasite with the greatest number of young leaving thenest was 0.11 ± 0.03, which was greater than expectedif eggs were accepted randomly from neighboring females or fromfemales present on the entire study area (p = .03 and p = .02,respectively). These patterns may reflect parasitism of randomlyselected nests followed by differential acceptance by hosts,differential hatching success of related parasites (e.g., dueto greater laying synchrony), or a mixture of parasitic strategies,one with a focus on related hosts and the other on unrelatedhosts. Genetic data revealed that social relationships did notalways reflect true relatedness and that success of primaryparasites was associated with kinship to hosts.     

Molecular identification of brood‐parasitic females reveals an opportunistic reproductive tactic in ruddy ducks

In many taxa, females lay eggs in the nests of other conspecifics. To determine the conditions under which conspecific brood parasitism develops, it is necessary to identify parasitic offspring and the females who produce them; however, for most systems parasitism can be difficult to observe and most genetic approaches have relatively low resolving power. In this study, we used protein fingerprinting from egg albumen and 10 microsatellite loci to genetically match parasitic ducklings to their mothers in a population of ruddy ducks (Oxyura jamaicensis). We found that 67% of nests contained parasitic offspring, and we successfully identified their mothers in 61% of the cases. Of the parasitic females identified, 77% also had nests of their own (i.e. a dual tactic, where females both nest and lay parasitically), and we found no evidence that parasitic females pursued a specialist (parasitism only) tactic. We also found that parasitic egg laying was not influenced by nest loss, predation or female condition. Thus, in contrast to most waterfowl studied to date, female ruddy ducks appear to lay parasitic eggs whenever the opportunity arises.     

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.     

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.     

Function of egg punctures by Shiny Cowbirds in parasitized and nonparasitized Creamy‐bellied Thrush nests

ABSTRACT Avian brood parasites usually remove or puncture host eggs. Several hypotheses have been proposed to explain the function of these behaviors. Removing or puncturing host eggs may enhance the efficiency of incubation of cowbird eggs (incubation‐efficiency hypothesis) or reduce competition for food between cowbird and host chicks in parasitized nests (competition‐reduction hypothesis) and, in nonparasitized nests, may force hosts to renest and provide cowbirds with new opportunities for parasitism when nests are too advanced to be parasitized (nest‐predation hypothesis). Puncturing eggs may also allow cowbirds to assess the development of host eggs and use this information to decide whether to parasitize a nest (test‐incubation hypothesis). From 1999 to 2002, we tested these hypotheses using a population of Creamy‐bellied Thrushes (Turdus amaurochalinus) in Argentina that was heavily parasitized by Shiny Cowbirds (Molothrus bonariensis). We found that 56 of 94 Creamy‐bellied Thrush nests (60%) found during nest building or egg laying were parasitized by Shiny Cowbirds, and the mean number of cowbird eggs per parasitized nest was 1.6 ± 0.1 (N= 54 nests). At least one thrush egg was punctured in 71% (40/56) of parasitized nests, and 42% (16/38) of nonparasitized nests. We found that cowbird hatching success did not differ among nests where zero, one, or two thrush eggs were punctured and that the proportion of egg punctures associated with parasitism decreased as incubation progressed. Thus, our results do not support the incubation‐efficiency, nest‐predation, or test‐incubation hypotheses. However, the survival of cowbird chicks in our study was negatively associated with the number of thrush chicks. Thus, our results support the competition‐reduction hypothesis, with Shiny Cowbirds reducing competition between their young and host chicks by puncturing host eggs in parasitized nests.     

Parasitism of maternal investment selects for increased clutch size and brood reduction in a host

The allocation of resources to young that will ultimately beleft to die appears counterintuitive. Yet obligate brood reductionhas evolved in a number of species, despite the waste of reproductiveinvestment this may incur. Here we test whether brood parasitismcould be one factor leading to the evolution of obligate broodreduction because surplus eggs in the nest during incubationoffer some protection from the costs of parasitism. Surpluseggs could benefit females in two ways. First, additional eggsmay protect against the direct costs of parasitism by facilitatingrecognition and removal of parasitic eggs with greater accuracy.Second, additional eggs may protect against the indirect costsof parasitism as parasites often damage or remove host eggswhen entering the host nest; surplus eggs may be an essentialinsurance strategy against this damage. We test these possibilitiesin the Montezuma Oropendola (Psarocolius Montezuma), a speciesexperiencing high levels of parasitism by Giant Cowbirds (Scaphiduraoryzivora) throughout their range. Overall rejection rates ofcowbird eggs were high (72%), and experimental addition of parasiticeggs to empty, one-, and two-egg nests demonstrated that recognitionsuccess was unaffected by the presence of additional host eggsfor comparison. However, the value of surplus eggs when oneegg was removed or damaged by a parasite was high; 31.6% ofsuccessful two-egg clutches lost a single egg during incubationand would have failed to produce a chick without a second egg.This was directly attributable to parasitism in at least 33%of all cases. Therefore, despite highly developed host defensesagainst direct costs of parasitism (recognition and removalof parasitic eggs), the associated indirect costs (egg damageand removal) could play an important role in selection for aclutch size that results in more chicks than can be raised.     

Host activity and the risk of nest parasitism by brown-headed cowbirds

Proportions of nests parasitized by brown-headed cowbirds (Molothrus ater) vary greatly among host species, but factors underlyingthis variation remain poorly understood. Cowbirds are believedto find nests by watching host behavior. We tested the hypothesisthat the activity of hosts during nest building correlateswith the probability of parasitism among and within four sympatrichosts: dusky flycatchers (Empidonax oberholseri), warblingvireos (Vireo gilvus), yellow warblers (Dendroica petechia),and American redstarts (Setophaga ruticilla). Daily probabilityof parasitism varied substantially among these species, from3% for dusky flycatchers to more than four times that for warblingvireos. The four species did not differ in the proportion ofcowbirds fledged from their nests. Differences in nest placementdid not explain differences in probability of parasitism amongor within species. Parasitism frequencies increased among speciesthat made longer nest-building visits, had a greater propensityto perch during nest approach, spent more time near their nests,and had males that vocalized more often near nests. Within species, females that visited their nests less often, spent more timeon the nest per visit, and males that sang more and were activein a smaller area around their nests were more likely to beparasitized by cowbirds. These data support the hypothesisthat cowbirds use the activity of hosts during nest buildingto locate nests.     

Colony kin structure and host‐parasite relatedness in the barnacle goose

Conspecific brood parasitism (CBP), females laying eggs in the nest of other ‘host’ females of the same species, is a common alternative reproductive tactic among birds. For hosts there are likely costs of incubating and rearing foreign offspring, but costs may be low in species with precocial chicks such as waterfowl, among which CBP is common. Waterfowl show strong female natal philopatry, and spatial relatedness among females may influence the evolution of CBP. Here we investigate fine‐scale kin structure in a Baltic colony of barnacle geese, Branta leucopsis, estimating female spatial relatedness using protein fingerprints of egg albumen, and testing the performance of this estimator in known mother‐daughter pairs. Relatedness was significantly higher between neighbour females (nesting ≤ 40 metres from each other) than between females nesting farther apart, but there was no further distance trend in relatedness. This pattern may be explained by earlier observations of females nesting close to their mother or brood sisters, even when far from the birth nest. Hosts and parasites were on average not more closely related than neighbour females. In 25 of 35 sampled parasitized nests, parasitic eggs were laid after the host female finished laying, too late to develop and hatch. Timely parasites, laying eggs in the host’s laying sequence, had similar relatedness to hosts as that between neighbours. Females laying late parasitic eggs tended to be less related to the host, but not significantly so. Our results suggest that CBP in barnacle geese might represent different tactical life‐history responses.     

Antagonistic antiparasite defenses: nest defense and egg rejection in the magpie host of the great spotted cuckoo

Brood parasites dramatically reduce the reproductive successof their hosts, which therefore have developed defenses againstbrood parasites. The first line of defense is protecting thenest against adult parasites. When the parasite has successfullyparasitized a host nest, some hosts are able to recognize andreject the eggs of the brood parasite, which constitutes the secondline of defense. Both defense tactics are costly and would be counteractedby brood parasites. While a failure in nest defense implies successfulparasitism and therefore great reduction of reproductive successof hosts, a host that recognizes parasitic eggs has the opportunityto reduce the effect of parasitism by removing the parasiticegg. We hypothesized that, when nest defense is counteractedby the brood parasite, hosts that recognize cuckoo eggs shoulddefend their nests at a lower level than nonrecognizers becausethe former also recognize adult cuckoos. Magpie (Pica pica) hoststhat rejected model eggs of the brood parasitic great spottedcuckoo (Clamator glandarius) showed lower levels of nest defensewhen exposed to a great spotted cuckoo than when exposed toa nest predator (a carrion crow Corvus corone). Moreover, magpiesrejecting cuckoo eggs showed lower levels of nest defense againstgreat spotted cuckoos than nonrecognizer magpies, whereas differencesin levels of defense disappeared when exposed to a carrion crow.These results suggest that hosts specialize in antiparasitedefense and that different kinds of defense are antagonistically expressed.We suggest that nest-defense mechanisms are ancestral, whereasegg recognition and rejection is a subsequent stage in the coevolutionaryprocess. However, host recognition ability will not be expressedwhen brood parasites break this second line of defense.     

Brood parasitic European starlings do not lay high-quality eggs

Chicks of obligate brood parasites employ a variety of morphologicaland behavioral strategies to outcompete nest mates. Elevatedcompetitiveness is favored by natural selection because parasiticchicks are not related to their host parents or nest mates.When chicks of conspecific brood parasites (CBPs) are unrelatedto their hosts, they and their parents would also benefit fromtraits that enhance competitiveness. However, these traits mustbe inducible tactics in CBPs, since conspecific brood parasitism(CBP) is facultative. Such tactics could be induced by resourcespassed to offspring through the egg. Thus, females engagingin CBP should allocate to their eggs resources that will enhanceoffspring competitiveness. We tested this prediction in a populationof European starlings (Sturnus vulgaris) breeding in southernSweden. Previous research showed that almost all CBPs in thispopulation are floater females that have yet to breed in thecurrent season. We identified putative brood parasitic eggsthrough monitoring egg laying and verified parasitism usingprotein fingerprinting. We then determined whether parasiticeggs were larger, larger-yolked, or had higher concentrationsof yolk testosterone or androstenedione than control eggs. The14 brood parasitic eggs laid in active nests (those with clutchesof at least two eggs that were eventually incubated) did notdiffer from controls in any of these characteristics. Ten dumpedeggs, laid in nonactive nest-boxes or on the ground, were smallerand smaller-yolked than control eggs but did not differ in yolkandrogen concentrations. The failure of our prediction couldbe the result of high costs of investing in eggs, lack of competition-basedbenefits for chicks, or physiological constraints on egg manipulation.     

Evolution of host egg mimicry in a brood parasite, the great spotted cuckoo

Brood parasitism in birds is one of the best examples of coevolutionary interactions in vertebrates. Coevolution between hosts and parasites is assumed to occur because the parasite imposes strong selection pressures on its hosts, reducing their fitness and thereby favouring counter-adaptations (e.g. egg rejection) which, in turn, select for parasite resistance (e.g. egg mimicry). Great spotted cuckoos ( Clamator glandarius ) are usually considered a brood parasite with eggs almost perfectly mimicking those of their host, the magpie ( Pica pica ). However, Cl. glandarius also exploits South African hosts with very different eggs, both in colour and size, while the Cl. glandarius eggs are similar to those laid in nests of European hosts. Here, we used spectrophotometric techniques for the first time to quantify mimicry of parasitic eggs for eight different host species. We found: (1) non-significant differences in appearance of Cl. glandarius eggs laid in nests of different host species, although eggs laid in South Africa and Europe differed significantly; (2) contrary to the general assumption that Cl. glandarius eggs better mimic those of the main host in Europe ( P. pica ), Cl. glandarius eggs more closely resembled those of the azure-winged magpie ( Cyanopica cyana ), a potential host in which there is no evidence of recent parasitism; (3) the appearance of Cl. glandarius eggs was not significantly related to the appearance of host eggs. We discuss three possible reasons why Cl. glandarius eggs resemble eggs of some of their hosts. We suggest that colouration of Cl. glandarius eggs is an apomorphic trait, and that variation between eggs laid in South African and European host nests is due to genetic isolation among these populations and not due to variation in colouration of host eggs.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79 , 551–563.     

The evolution of egg rejection by cuckoo hosts in Australia and Europe

Exploitation of hosts by brood parasitic cuckoos is expectedto stimulate a coevolutionary arms race of adaptations and counteradaptations.However, some hosts have not evolved defenses against parasitism.One hypothesis to explain a lack of host defenses is that thelife-history strategies of some hosts reduce the cost of parasitismto the extent that accepting parasitic eggs in the nest is evolutionarilystable. Under this hypothesis, it pays hosts to accept cuckooeggs if (1) the energetic cost of raising the cuckoo is low,(2) there is time to renest, and (3) clutch size is small. Weparasitized the nests of host and nonhost species with nonmimeticmodel eggs to test whether the evolution of egg recognitionby cuckoo hosts could be explained by life-history variablesof the host. The most significant factor explaining rates ofrejection of model eggs was whether or not a species was a cuckoohost, with hosts rejecting model eggs at a higher rate thannonhosts. Egg-rejection rates were also explained by visibilitywithin the nest and by cuckoo mass. We found little supportfor the life-history model of egg rejection. Our results suggestthat parasitism is always sufficiently costly to select forhost defenses and that the evolution of defenses may be limitedby proximate constraints such as visibility within the nest.     

Nest destruction elicits indiscriminate con‐ versus heterospecific brood parasitism in a captive bird

Following nest destruction, the laying of physiologically committed eggs (eggs that are ovulated, yolked, and making their way through the oviduct) in the nests of other birds is considered a viable pathway for the evolution of obligate interspecific brood parasitism. While intraspecific brood parasitism in response to nest predation has been experimentally demonstrated, this pathway has yet to be evaluated in an interspecific context. We studied patterns of egg laying following experimental nest destruction in captive zebra finches, Taeniopygia guttata, a frequent intraspecific brood parasite. We found that zebra finches laid physiologically committed eggs indiscriminately between nests containing conspecific eggs and nests containing heterospecific eggs (of Bengalese finches, Lonchura striata vars. domestica), despite the con‐ and heterospecific eggs differing in both size and coloration. This is the first experimental evidence that nest destruction may provide a pathway for the evolution of interspecific brood parasitism in birds.     

Parasitism strategy of the grey starling, Sturnus cineraceus: Selection based on host characters and nest location

The strategy used by the intraspecific brood parasite, the grey starling, Sturnus cineraceus (Temminck) and the degree to which this strategy reflected the sources of mortality for parasite eggs were examined. Approximately 74% of all parasite eggs failed to produce young that survived to fledglings. Most of this mortality was due to two factors: (i) laying in a nest that had been deserted by a host during its nesting cycle (19%); and (ii) mismatched timing of laying in the hosts nesting cycle (38%). It is important for parasites to select a suitable host in order to avoid this mortality and increase their reproductive success. However, grey starlings did not select hosts on the basis of nest location, host characteristics, or laying date. Lack of attention to these factors implies a failure on the part of the grey starlings to recognize cues that could direct them to select host nests that would provide the best environment for their eggs. Although some egg loss and egg replacement occurred before clutch initiation by hosts, no evidence was found that parasitic birds removed host eggs after clutch initiation by hosts. These results suggest that parasites did not adopt a successful strategy for enhancing the survival rate of their own eggs.     

Recognizing odd smells and ejection of brood parasitic eggs. An experimental test in magpies of a novel defensive trait against brood parasitism

One of the most important defensive host traits against brood parasitism is the detection and ejection of parasitic eggs from their nests. Here, we explore the possible role of olfaction in this defensive behaviour. We performed egg‐recognition tests in magpie Pica pica nests with model eggs resembling those of parasitic great spotted cuckoos Clamator glandarius. In one of the experiment, experimental model eggs were exposed to strong or moderate smell of tobacco smoke, whereas those of a third group (control) were cleaned with disinfecting wipes and kept in boxes containing odourless cotton. Results showed that model eggs with strong tobacco scent were more frequently ejected compared with control ones. In another experiment, models were smeared with scents from cloacal wash from magpies (control), cloacal wash or uropygial secretions from cuckoos, or human scents. This experiment resulted in a statistically significant effect of treatment in unparasitized magpie nests in which control model eggs handled by humans were more often rejected. These results provide the first evidence that hosts of brood parasites use their olfactory ability to detect and eject foreign eggs from their nests. These findings may have important consequences for handling procedures of experimental eggs used in egg‐recognition tests, in addition to our understanding of interactions between brood parasites and their hosts.     

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     

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