Effects of Shiny Cowbird Molothrus bonariensis parasitism on different components of House Wren Troglodytes aedon reproductive success

Avian brood parasites, including cuckoos and cowbirds, have multiple negative effects on their hosts. We analysed the effects of Shiny Cowbird Molothrus bonariensis parasitism on different components (e.g. egg losses, hatching success, chick survival and nest abandonment) of House Wren Troglodytes aedon reproductive success. We also conducted an experiment to discriminate between two mechanisms that may reduce hatching success in parasitized clutches: lower efficiency of incubation due to the increase in clutch volume and disruption of host incubation by the early hatching of Cowbirds. Egg puncturing by Shiny Cowbirds reduced host clutch size at hatching by 10–20%, and parasitized nests had a decrease in hatching success of 40–80%. Egg losses and hatching failures were positively associated with the intensity of parasitism. Brood reduction was greater in parasitized nests, but the growth rate of the chicks that fledged was similar to that in unparasitized nests. The combined effects of egg losses, hatching failures and brood reduction decreased the number of fledged chicks by 80%. In addition, egg puncturing increased the likelihood of nest abandonment by Wrens. Experimental data showed that hatching failures occurred when there was a combination of: (1) an increase in the volume of the clutch by the addition of the Cowbird egg without removal of host eggs, and (2) the addition of the Cowbird egg before the onset of incubation. This was relatively common in House Wren nests, as Cowbirds generally parasitize before the onset of incubation. Our results indicate that Shiny Cowbird parasitism imposes a major impact on House Wrens, as it affects all components of the Wren's reproductive success.     

History of Shiny Cowbird Molothrus bonariensis brood parasitism in Trinidad and Tobago

In attempting to evaluate the evolutionary stability of the reproductive strategies of an avian brood parasite, it is important to know whether or not the observed levels of parasitism on host species have changed through time and whether or not new host species are being used in the region under study. In this regard, the islands of Trinidad and Tobago provide an excellent opportunity because they are ornithologically well known and the expansion of the parasitic Shiny Cowbird Molothrus bonariensis into the West Indian region is thought to have originated from Trinidad and Tobago. On the whole, host choice has remained remarkably stable over the past 50–60 years. The three hosts (House Wren Troglodytes aedon, Red-breasted Blackbird Leistes militaris and Yellow-hooded Blackbird Agelaius icterocephalus) known to be heavily parasitized historically remain heavily parasitized today. Only one instance of parasitism of a new host (Tropical Kingbird Tyrannus melancholicus) is recorded. In the case of one intensively studied host, the Yellow-hooded Blackbird, cowbird parasitism had a minimal negative effect on the host's reproductive success. Reproductive equilibrium in this study seemed to be maintained by behavioural adaptations of host and parasite to each other. The stable pattern of host choice by Shiny Cowbirds in Trinidad and Tobago may not develop elsewhere in the West Indian region. Before the cowbird's expansion in the West Indies, potential hosts had no prior experience of brood parasites. Consequently, the island bird populations are unlikely to have evolved defensive strategies and thus may be more vulnerable to reproductive failure as a result of cowbird parasitism, precluding the development of a stable system at this point in time.     

Brood parasitism of White-rumped Swallows by Shiny Cowbirds

ABSTRACT.   Swallows and martins are infrequent hosts of the generalist brood parasite Shiny Cowbird ( Molothrus bonariensis ). We monitored 50 nesting attempts by White-rumped Swallows ( Tachycineta leucorrhoa ) over a two-year period in Argentina and detected low rates of brood parasitism (three nests, or 6%). Of the three nests parasitized, cowbirds ( N = 1 per nest) successfully fledged from two. Eight of 13 swallow eggs in these three nests were punctured by cowbirds, and all but one swallow nesting starved at the two parasitized nests. At least two factors may contribute to the low frequency of parasitism of White-rumped Swallows by Shiny Cowbirds, including the inability of the larger cowbirds to enter some nest cavities and the aggressive nest defense behavior of adult swallows.     

Shiny Cowbird parasitism of a low quality host: effect of host traits on a parasite's reproductive success

ABSTRACT.   The reproductive success of parasitic cowbirds ( Molothrus spp.) varies among host species and is influenced by the degree of synchronization in timing of egg laying, the duration of parasite and host incubation periods, and the ability of hosts to incubate and rear parasite young. We studied the reproductive success of Shiny Cowbirds ( Molothrus bonariensis ) that parasitized the nests of Creamy-bellied Thrushes ( Turdus amaurochalinus ) in the Monte desert region of Argentina. Shiny Cowbirds frequently parasitized Creamy-bellied Thrush nests (60%), and most cowbirds synchronized egg laying with that of thrushes (79%). Most parasitic eggs (80%) hatched within 1 d of the hatching of the first host egg, and more than 91% of the eggs survived until the end of the incubation. However, only 60% of the cowbird eggs hatched and 52% of young survived. The proportion of Shiny Cowbirds eggs laid in Creamy-bellied Thrush nests that resulted in fledged young was 0.03, including eggs and young lost due to predation or desertion. Despite this low reproductive success, Creamy-bellied Thrushes were heavily parasitized by Shiny Cowbirds in our study area. Shiny Cowbirds may continue to parasitize these thrushes because of diffuse selection or because Shiny Cowbird chicks are more likely to fledge from Creamy-bellied Thrush nests in years or areas with greater food availability when brood reduction does not occur.     

Do Hosts Discriminate between Sexually Dichromatic Male and Female Brown‐headed Cowbirds?

Female brood parasites are recognized as threats to reproductive success by many host species. Male brood parasites may accompany females while they search for nests to parasitize and males depredate nests throughout the nesting cycle. Hence, selection may also favour recognition of males. We examined whether two common host species perceive male brown-headed cowbirds ( Molothrus ater ) as brood parasites, as nest predators, or neither. We quantified visits of male cowbirds to nests of yellow warblers ( Dendroica petechia ) and red-winged blackbirds ( Ageliaus phoeniceus ) to assess the frequency with which these host species interact with male cowbirds. Males were observed near nests during hosts' laying and incubating stages, although less frequently than female cowbirds. No visits by cowbirds occurred while parents cared for nestlings. We then presented models of male and female cowbirds plus a non-threatening control to yellow warblers and red-winged blackbirds during laying and nestling periods. If hosts perceive males and females similarly, they should respond more intensely to the cowbird models during the laying period, when nests are most likely to be parasitized. Both species responded similarly to male and female cowbird models during laying, which suggests that hosts view cowbirds of both sexes as threats. The responses of yellow warblers with nestlings to male cowbirds were strongly influenced by the order of model presentation. Warblers first presented with the male cowbird gave much reduced anti-parasite responses than those that first interacted with the female then the male cowbird. These results suggest that yellow warblers recognized male vs. female cowbirds, but that discrimination was not expressed during laying. By contrast, red-winged blackbirds did not discriminate between male and female cowbirds at either nesting stage.     

Molecular tracking of individual host use in the Shiny Cowbird – a generalist brood parasite

Generalist parasites exploit multiple host species at the population level, but the individual parasite's strategy may be either itself a generalist or a specialist pattern of host species use. Here, we studied the relationship between host availability and host use in the individual parasitism patterns of the Shiny Cowbird Molothrus bonariensis, a generalist avian obligate brood parasite that parasitizes an extreme range of hosts. Using five microsatellite markers and an 1120‐bp fragment of the mtDNA control region, we reconstructed full‐sibling groups from 359 cowbird eggs and chicks found in nests of the two most frequent hosts in our study area, the Chalk‐browed Mockingbird Mimus saturninus and the House Wren Troglodytes aedon. We were able to infer the laying behavior of 17 different females a posteriori and found that they were mostly faithful to a particular laying area and host species along the entire reproductive season and did not avoid using previously parasitized nests (multiple parasitism) even when other nests were available for parasitism. Moreover, we found females using the same host nest more than once (repeated parasitism), which had not been previously reported for this species. We also found few females parasitizing more than one host species. The use of an alternative host was not related to the main hosts' nest availability. Overall, female shiny cowbirds use a spatially structured and host species specific approach for parasitism, but they do so nonexclusively, resulting in both detectable levels of multiple parasitism and generalism at the level of individual parasites.     

Why brown-headed cowbirds do not influence red-winged blackbird parent behaviour

Brown-headed cowbirds, Molothrus ater, frequently parasitize red-winged blackbirds,Agelaius phoeniceus . The presence of a brood parasite, unrelated to both host nestlings and parents, has provoked speculation regarding within-brood food allocation and parental provisioning. This study is the first to compare directly the effect of brood parasitism on host parent and offspring behaviour in younger and older broods. We videotaped 28 unparasitized red-winged blackbird broods and compared them to 22 parasitized broods. Red-winged blackbird nestling begging appears largely unaffected by cowbird parasitism. The presence of the cowbird in the nest affected neither the latency nor duration of host nestling begging, but stimulated more frequent begging by red-winged blackbird nestlings following food distribution. Begging by cowbirds was unique in two ways: (1) cowbirds maintained a consistent begging effort throughout the nestling period (but did not receive a consistent food share); and (2) cowbirds begged longer and more frequently following the allocation of food. Persistent begging by the cowbird following the allocation of food has implications for the division of parental care, if by doing so the brood parasite is able to provoke the foster parent to increase provisioning, at the expense of brooding. We found no evidence for the adjustment of parental care. Neither the foraging rates nor the lengths of the parental feeding visits differed markedly between parasitized and unparasitized broods. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

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.     

Brood parasitism increases provisioning rate, and reduces offspring recruitment and adult return rates, in a cowbird host

Interspecific brood parasitism in birds presents a special problem for the host because the parasitic offspring exploit their foster parents, causing them to invest more energy in their current reproductive effort. Nestling brown-headed cowbirds (Molothrus ater) are a burden to relatively small hosts and may reduce fledgling quality and adult survival. We documented food-provisioning rates of one small host, the prothonotary warbler (Protonotaria citrea), at broods that were similar in age (containing nestlings 8–9 days old), but that varied in composition (number of warbler and cowbird nestlings) and mass, and measured the effect of brood parasitism on offspring recruitment and adult returns in the host. The rate of food provisioning increased with brood mass, and males and females contributed equally to feeding nestlings. Controlling for brood mass, the provisioning rate was higher for nests with cowbirds than those without. Recruitment of warbler fledglings from unparasitized nests was 1.6 and 3.7 times higher than that of fledglings from nests containing one or two cowbirds, respectively. Returns of double-brooded adult male and female warblers decreased with an increase in the number of cowbirds raised, but the decrease was more pronounced in males. Reduced returns of warbler adults and recruitment of warbler fledglings with increased cowbird parasitism was likely a result of reduced survival. Cowbird parasitism increased the warblers’ investment in current reproductive effort, while exerting additional costs to current reproduction and residual reproductive value. Our study provides the strongest evidence to date for negative effects of cowbird parasitism on recruitment of host fledglings and survival of host adults.     

Effects of nest-site characteristics and parental activity on cowbird parasitism and nest predation in Brown-and-yellow Marshbirds

ABSTRACT Nest‐site selection and nest defense are strategies for reducing the costs of brood parasitism and nest predation, two selective forces that can influence avian nesting success and fitness. During 2001–2002, we analyzed the effect of nest‐site characteristics, nesting pattern, and parental activity on nest predation and brood parasitism by cowbirds (Molothrus spp.) in a population of Brown‐and‐yellow Marshbirds (Pseudoleistes virescens) in the Buenos Aires province, Argentina. We examined the possible effects of nest detectability, nest accessibility, and nest defense on rates of parasitism and nest predation. We also compared rates of parasitism and nest predation and nest survival time of marshbird nests during the egg stage (active nests) with those of the same nests artificially baited with passerine eggs after young fledged or nests failed (experimental nests). Most nests (45 of 48, or 94%) found during the building or laying stages were parasitized, and 79% suffered at least one egg‐predation event. Cowbirds were responsible for most egg predation, with 82 of 107 (77%) egg‐predation events corresponding to eggs punctured by cowbirds. Nests built in thistles had higher rates of parasitism and egg predation than nests in other plant, probably because cowbirds were most active in the area where thistles were almost the only available nesting substrate. Parasitism rates also tended to increase as the distance to conspecific nests increased, possibly due to cooperative mobbing and parental defense by marshbirds. The proportion of nests discovered by cowbirds was higher for active (95%) than for experimental (29%) nests, suggesting that cowbirds used host parental activity to locate nests. Despite active nest defense, parental activity did not affect either predation rates or nest‐survival time. Thus, although nest defense by Brown‐and‐yellow Marshbirds appears to be based on cooperative group defense, such behavior did not reduce the impact of brood parasites and predators.     

Sex and seasonal differences in hippocampal volume and neurogenesis in brood‐parasitic brown‐headed cowbirds (Molothrus ater)

Brown‐headed cowbirds (Molothrus ater) are one of few species in which females show more complex space use than males. Female cowbirds search for, revisit, and parasitize host nests and, in a previous study, outperformed males on an open field spatial search task. Previous research reported a female‐biased sex difference in the volume of the hippocampus, a region of the brain involved in spatial memory. Neurons produced by adult neurogenesis may be involved in the formation of new memories and replace older neurons that could cause interference in memory. We tested for sex and seasonal differences in hippocampal volume and neurogenesis of brood‐parasitic brown‐headed cowbirds and the closely related non‐brood‐parasitic red‐winged blackbird (Agelaius phoeniceus) to determine whether there were differences in the hippocampus that reflected space use in the wild. Females had a larger hippocampus than males in both species, but hippocampal neurogenesis, measured by doublecortin immunoreactivity (DCX+), was greater in female than in male cowbirds in the absence of any sex difference in blackbirds, supporting the hypothesis of hippocampal specialization in female cowbirds. Cowbirds of both sexes had a larger hippocampus with greater hippocampal DCX+ than blackbirds. Hippocampus volume remained stable between breeding conditions, but DCX+ was greater post‐breeding, indicating that old memories may be lost through hippocampal reorganization following breeding. Our results support, in part, the hypothesis that the hippocampus of cowbirds is specialized for brood parasitism. © 2016 Wiley Periodicals, Inc. Develop Neurobiol 76: 1275–1290, 2016     

Community-level patterns of population recruitment in a generalist avian brood parasite, the brown-headed cowbird

The brown-headed cowbird (Molothrus ater) is a generalist brood parasite that typically parasitizes many host species in a single bird community. Population recruitment in a generalist parasite should be diverse with respect to host species; however, host-specific rates of cowbird recruitment have not been reported in any host community, and the determinants of host quality are poorly known. We studied the combined influence of parasitism level, nest abundance, and host quality on community-level patterns of cowbird recruitment in New Mexico, USA. Our objectives were to: (1) evaluate patterns of host use and quality; (2) compare cowbird egg investment and recruitment among host species; (3) identify host species of most importance to cowbird recruitment. Cowbirds parasitized 11 host species, with five “major” hosts experiencing high parasitism levels (≥1 cowbird egg/nest) and six minor hosts experiencing low parasitism levels (<0.3 cowbird eggs/nest). Parasitism level was not correlated with host species abundance, host mass, host nestling period length, or host success at fledging cowbirds. However, tree-nesting hosts were parasitized more than ground-nesters, and foliage-gleaners more than sally-foragers and ground-foragers. Average estimated survival to fledging of cowbird eggs laid in active host nests was 0.19. Cowbird recruitment was diverse with respect to hosts but was less evenly distributed across the host community than was cowbird egg investment because western tanagers (Piranga ludovicianus) fledged cowbirds more successfully than other hosts. This success in western tanagers was due to high cowbird survivorship in tanager nests and may be associated with the larger body size of tanagers relative to other hosts.     

Multiple effects of brood parasitism reduce the reproductive success of prothonotary warblers, Protonotaria citrea

Avian brood parasitism often has multiple negative effects on the reproductive success of the host. Most studies have focused on one or two of these effects, but rarely have they all been studied simultaneously for one species. I studied prothonotary warblers to quantify the effects of different intensities of (i.e. multiple) brood parasitism by brown-headed cowbirds, Molothrus ater, on the production of host and cowbird young and on the between-year returns of adult warblers. Host clutch size decreased with an increase in the number of cowbird eggs laid in nests. The hatching success of warbler and cowbird eggs decreased with increased cowbird eggs in nests, but was always higher for cowbird eggs than warbler eggs. The survival of warbler nestlings, but not cowbird nestlings, decreased with increased cowbird nestlings in the brood. An increase in the number of cowbird nestlings in broods resulted in a reduction in the average mass of warbler nestlings but not cowbird nestlings. The number of cowbird eggs or nestlings present did not affect nest predation, and the fledging of cowbirds did not influence the renesting interval of female warblers. In addition, the between-year returns of adult warblers were not negatively affected by brood parasitism. Decreased hatching success and nestling survival reduced the reproductive output of the warblers the most. These effects were substantial and appear to favour the evolution of behavioural responses that reduce the effects of brood parasitism on prothonotary warblers. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour      

Brood parasitism by cowbirds: risks and effects on reproductive success and survival in indigo buntings

We observed brood parasitism by brown-beaded cowbirds (Molothrusater) on indigo buntings (Passerina cyanea) and estimated dieimpact of parasitism on the success of the individual buntingsin their current nests and in their future survival and reproduction.Rates of parasitism over 8 years were 26.6% in 1040 nests and19.8% in 693 nests in two areas in southern Michigan. Risk ofparasitism was high early in the season; half the bunting nestswere begun after the end of the cowbird season. Risk was independentof female age, plant containing the nest, or habitat The immediatecost of parasitism was 1.19 and 1.26 fewer buntings fledgedper nest. Bunting success was lower in parasitized nests withcowbird eggs (nests were more likely to be deserted or predated),lower when the cowbird nestling failed (nests were more likelyto be predated), and lower when the cowbird fledged (fewer buntingsfledged) compared to nonparasitized nests. Costs were due toremoval of a bunting egg when die cowbird laid its own egg andto competition for parental care of the cowbird and buntingnestlings. Buntings that fledged from nests where a cowbirdalso fledged were only 18% as likely to survive and return totheir natal area in the next year as buntings from nests wherea cowbird did not fledge. Long-term effects of cowbird parasitismon adult breeding later in the season, survival to the nextseason, and reproductive success in the next season were negligiblewhen compared between birds that reared a cowbird and birdsthat reared only a bunting brood, or between birds that wereparasitized and birds that escaped parasitism. The results indicatelittle long-term cost of brood parasitism on individual fitnessof adult buntings beyond the impact on the current nest andthe survival of buntings that fledge from it; nearly all costis to the parasitized brood.     

Roosting behaviour is related to reproductive strategy in brood parasitic cowbirds

Shiny Cowbirds Molothrus bonariensis and Screaming Cowbirds Molothrus rufoaxillaris are closely related brood parasites but the former is socially polygynous or promiscuous and an extreme host generalist, whereas the latter is socially monogamous and parasitizes almost exclusively one host. Females of both species lay in relative darkness, before dawn, relying for host nest location on previous days’ prospecting activity, or possibly on following better‐informed roost associates. We studied the temporal and spatial patterns of roosting behaviour in these species to test the hypothesis that roosting behaviour of cowbirds is related to their breeding strategy (brood parasitism) and reflects differences in strategies between species. We recorded fidelity to a roost, location fidelity within a roost, inter‐individual spatial associations and timing of roost departures and parasitic events, using tagged individuals. Female Shiny Cowbirds and both sexes of Screaming Cowbirds showed marked fidelity in roosting location, and roost departures occurred both during and after the known time window for parasitism, with earlier departures probably corresponding to laying days. Screaming Cowbird females and males that were trapped together and showed high levels of association during the day, also showed high levels of association in the roost. We describe the spatial and temporal patterns of a relatively poorly known aspect of avian ecology in general and the behaviour of brood parasites in particular.     

Experiments and observations of prothonotary warblers indicate a lack of adaptive responses to brood parasitism

It has been suggested that prothonotary warblers, Protonotaria citrea, respond adaptively to brood parasitism by brown-headed cowbirds, Molothrus ater, even though they lack historical habitat and range overlap with cowbirds. I studied behaviours functioning as potential defences against brood parasitism in the prothonotary warbler, a cavity-nesting host species. Opening sizes preferred by prothonotary warblers were not small enough to exclude cowbirds, and warblers were parasitized heavily in nests with larger openings. Male and female prothonotary warblers were always away from their nests before sunrise when cowbirds laid eggs in their nests. Prothonotary warblers infrequently (∼6% of 560 nests) deserted nests that were parasitized during the egg-laying period, but frequently (56% of 151 nests) deserted nests that were parasitized before a female warbler laid her first egg. Prothonotary warblers also deserted 60-70% of nests where a cowbird egg, warbler egg or die were experimentally added before egg laying. However, the experimental addition of one of these three objects during the egg-laying period did not elicit desertion. The desertion of parasitized nests was not affected by nest site availability as has been reported elsewhere in the literature. This lack of a response to brood parasitism by prothonotary warblers may be an example of evolutionary lag, because it is likely that they have only recently been exposed to widespread parasitism, and they accept parasitism at a high cost to their own reproductive success. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

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.     

Reproductive ecology of blackbirds ( Turdus merula maximus) in a high-altitude location, Tibet

Reproductive data of high-altitude blackbirds (Turdus merula maximus) were collected at an alpine environment covered with scrub vegetation in Lhasa mountains, Tibet. The blackbird nests were found from 3,800 to 4,500 m at a breeding density of 0.07 pairs/ha. Egg-laying occurred from early May to late July, with a peak (73.0% of all nesting attempts) between mid-May and mid-June. Most (79.5%) of the nests were sited in six different plant species with Cotoneaster microphyllus being most preferred. A few (20.5%) nests were against rocky walls. Bushes used for nest sites were significantly higher. Compared with lowland blackbirds, montane blackbirds experienced a shorter breeding season (2.5 months), laid smaller clutches (2–4 eggs, averaging 2.86), but larger eggs (mean volume index 16,348), produced only one brood per year, had a slightly shorter incubation period (12–13 days), but a longer nestling period (16–18 days), and enjoyed a higher breeding success (59.0%). In the alpine environment where climate is harsh and food supply poor, the strategy of increased egg size with decreased clutch size suggests that the high-altitude blackbirds improve their breeding success by investing more reproductive energy in fewer offspring in response to the constraints of time and resources. However, the environmental constraints on reproduction result in a low annual reproductive output.This revised version was published online in December 2004. The german abstract has been included.     

Density-dependent habitat selection by brown-headed cowbirds (<Emphasis Type="Italic">Molothrus ater</Emphasis>) in tallgrass prairie

Local distributions of avian brood parasites among their host habitats may depend upon conspecific parasite density. We used isodar analysis to test for density-dependent habitat selection in brown-headed cowbirds (Molothrus ater) among tallgrass prairie adjacent to wooded edges, and prairie interior habitat (>100 m from wooded edges) with and without experimental perches. Eight study sites containing these three habitat treatments were established along a geographical gradient in cowbird abundance within the Flint Hills region of Eastern Kansas and Oklahoma, USA. The focal host species of our study, the dickcissel (Spiza americana), is the most abundant and preferred cowbird host in the prairie of this region. Cowbird relative abundance and cowbird:host abundance ratios were used as estimates of female cowbird density, whereas cowbird egg density was measured as parasitism frequency (percent of dickcissel nests parasitized), and parasitism intensity (number of cowbird eggs per parasitized nest). Geographical variation in cowbird abundance was independent of host abundance. Within study sites, host abundance was highest in wooded edge plots, intermediate in the experimental perch plots, and lowest in prairie interior. Cowbirds exhibited a pattern of density-dependent selection of prairie edge versus experimental perch and interior habitats. On sites where measures of cowbird density were lowest, all cowbird density estimates (female cowbirds and their eggs) were highest near (100 m) wooded edges, where host and perch availability are highest. However, as overall cowbird density increased geographically, these density estimates increased more rapidly in experimental perch plots and prairie interiors. Variation in cowbird abundance and cowbird:host ratios suggested density-dependent cowbird selection of experimental perch over prairie interior habitat, but parasitism levels on dickcissel nests were similar among these two habitats at all levels of local cowbird parasitism. The density-dependent pattern of cowbird distribution among prairie edge and interior suggested that density effects on perceived cowbird fitness are greatest at wooded edges. A positive relationship between daily nest mortality rates of parasitized nests during the nestling period with parasitism intensity levels per nest suggested a density-dependent effect on cowbird reproductive success. However, this relationship was similar among habitats, such that all habitats should have been perceived as being equally suitable to cowbirds at all densities. Other unmeasured effects on cowbird habitat suitability (e.g., reduced cowbird success in edge-dwelling host nests, cowbird despotism at edges) might have affected cowbird habitat selection. Managers attempting to minimize cowbird parasitism on sensitive cowbird hosts should consider that hosts in otherwise less-preferred cowbird habitats (e.g., habitat interiors) are at greater risk of being parasitized where cowbirds become particularly abundant.     

THE DECLINE OF AN ADAPTATION IN THE ABSENCE OF A PRESUMED SELECTION PRESSURE

The colonial nesting Village Weaver (Ploceus cucullatus) lays eggs that vary in ground color and pattern, but individual females lay similar eggs each time. Tests on captive African stocks have shown that females reject eggs of other cohorts if such eggs are sufficiently different. The Village Weaver may have evolved rejection behavior and variable eggs in response to cuckoo parasitism in Africa. The Village Weaver was introduced into Hispaniola from Africa as early as the 18th century. Before the arrival of the Shiny Cowbird (Molothrus bonariensis) in the early 1970's, there were no brood parasites on Hispaniola. Furthermore, in an experimental parasitism study, Hispaniolan Village Weavers accepted both dummy eggs and dissimilar Village Weaver eggs. The Village Weaver may have decreased the egg-rejection behavior in the absence of the selective pressure of brood parasitism. Now Hispaniolan populations of the Village Weaver are parasitized by the Shiny Cowbird, which lays eggs dissimilar to those of the weaver. Brood parasitism by the Shiny Cowbird exerts a detrimental impact on the Village Weaver by reducing nest success and productivity.