Moisture as a determinant of habitat quality for a nonbreeding neotropical migratory songbird

Identifying the determinants of habitat quality for a species is essential for understanding how populations are limited and regulated. Spatiotemporal variation in moisture and its influence on food availability may drive patterns of habitat occupancy and demographic outcomes. Nonbreeding migratory birds in the neotropics occupy a range of habitat types that vary with respect to moisture. Using carbon isotopes and a satellite-derived measure of habitat moisture, we identified a moisture gradient across home ranges of radio-tracked Northern Waterthrush (Seiurus noveboracensis). We used this gradient to classify habitat types and to examine whether habitat moisture correlates with overwinter mass change and spring departure schedules of Northern Waterthrush over the late-winter dry season in the tropics. The two independent indicators of moisture revealed similar gradients that were directly proportional to body mass change as the dry season progressed. Birds occupying drier habitats declined in body mass over the study period, while those occupying wetter habitats increased in body mass. Regardless of habitat, birds lost an average of 7.6% of their mass at night, and mass recovery during the day trended lower in dry compared with wet habitats. This suggests that daily incremental shortfalls in mass recovery can lead to considerable season-long declines in body mass. These patterns resulted in consequences for the premigratory period, with birds occupying drier habitats having a delayed rate of fat deposition compared with those in wet habitats. Taken together with the finding that males, which are significantly larger than females, are also in better condition than females regardless of habitat suggests that high-quality habitats may be limited and that there may be competition for them. The habitat-linked variation in performance we observed suggests that habitat limitation could impact individual and population-level processes both during and in subsequent periods of the annual cycle. The linkage between moisture and habitat quality for a migratory bird indicates that the availability of high-quality habitats is dynamic due to variation in precipitation among seasons and years. Understanding this link is critical for ascertaining the impact of future climate change, particularly in the Caribbean basin, where a much drier future is predicted.     

Parasite transmission in a migratory multiple host system

A transmission model was devised for trichostrongyloid nematodes of saiga antelopes and domestic sheep in Kazakhstan. The framework extends previous models by including seasonal migration of saigas, contact with separate populations of sheep, and climate-driven stochasticity in herbage biomass and in the development, survival and migration onto herbage of free-living larvae. The model was parameterised for the contrasting life histories of Marshallagia, Haemonchus and Nematodirus, three important parasites of saigas and sheep in the region, and was successful at predicting broad qualitative patterns of infection dynamics in sheep and saigas. Parasite transmission between saigas and sheep was predicted to be most important for Marshallagia (from sheep to saigas in the south in winter, and onward transmission to sheep in the north in summer) and Haemonchus (from sheep in the north in summer via saigas to sheep further south in autumn). Model predictions for winter transmission of Marshallagia infection in saigas were consistent with field data, which showed that saigas culled before they have grazed the winter range carry lower burdens of this parasite than older animals. The model provides a mechanistic explanation for its predictions, which will assist hypothesis formation, and further the epidemiological basis of efforts to control parasite transmission between wildlife and livestock in both directions. A similar modelling approach could prove useful in other situations where detailed mechanistic models of parasite transmission are inappropriate in the face of parameter uncertainty and spatio-temporal variation in climate and host density. This is likely to include the majority of wildlife-parasite systems.     

The nature of the migration route shapes physiological traits and aerodynamic properties in a migratory songbird

Migration distance is supposed to represent an important selection pressure shaping physiological and morphological properties. Previous work has focussed on this effect, while the importance of ecological barriers in this context has been rarely considered. We studied two subspecies of a migratory songbird, the northern wheatear (Oenanthe oenanthe oenanthe and O. o. leucorhoa L.), on an island in the North Sea. The former subspecies reaches their Scandinavian breeding areas after a short sea crossing, whereas leucorhoa northern wheatears cross the North Atlantic towards Iceland, Greenland or Canada. Physiological traits (fuel deposition rate) and wings’ aerodynamic properties (wing pointedness independent of body size), both affecting migration speed, were hypothesized to be more pronounced in leucorhoa than in oenanthe northern wheatears. Within subspecies, the physiological and aerodynamic properties were hypothesized to explain arrival date at the stopover site with “fast migrants” arriving early. Physiological and aerodynamic properties in leucorhoa northern wheatears lead to a faster and less costly migration, favouring a sea crossing, but in trade-off lower flight manoeuvrability than in oenanthe birds. Wings’ aerodynamic properties affected the seasonal occurrence of leucorhoa females, whereas the physiological traits significantly influenced arrival date in oenanthe individuals. The less risky migration route in oenanthe birds with few short sea crossings may have favoured higher flight manoeuvrability for foraging (less pointed wings), in trade-off an energetically more costly flight. Hence, not the migration distance itself, but the presence/absence of a sea barrier presents an important selection pressure in migratory land birds favouring low flight costs.     

Delayed maturation in plumage colour: Evidence for the female-mimicry hypothesis in the kestrel

Summary In many sexually dichromatic species, young males have female-like plumage during their first potential breeding year. The female-mimicry hypothesis (FMH) supposes that by possessing female-like plumage young males deceive older conspicuous males into believing that they are females, thus reducing competition from adult males. The status-signalling hypothesis (SSH) supposes that adult males can distinguish sex, but postulates that young males reduce competition from adult males by reliably signaling low status with their dull plumage. We tested these hypotheses in the European kestrel (Falco tinnunculus). Female-like young males settled to breed closer to adult males than did other adult males (Figs. 1a, b). By settling near adult males, young males seemed to increase their chance of mating with adult females. Adult female-young male pairs had better reproductive success than yearling-yearling pairs. These results suggest that there is an adaptive value in possessing a female-like plumage colour in the breeding season. To test the FMH, we measured sexual preference of adult males when adult females and young males were simultaneously shown in an aviary. Adult males were unable to recognize sex, because in half the cases they preferred young males (Fig. 3). However, when adult males and females were shown simultaneously, males preferred females (Fig. 2). Our results support the FMH rather than the SSH, because young males successfully deceived older males by their plumage.     

Pigment allocation to eggs decreases plasma antioxidants in a songbird

Although evidence is accumulating on the adaptive function of female ornamentation, very little is known about maternal allocation decisions involving sexual signaling and other reproductive functions. Blue egg coloration has been suggested as a sexually selected signal of female quality to males, and some recent studies are in accordance with this hypothesis. Blue eggshell coloration results from the deposition of biliverdin pigment by laying females, which is a potent antioxidant. Thus, egg pigmentation should be costly in terms of antioxidants, an assumption of the signaling hypothesis that has not been tested yet. We induced increased reproductive effort in a set of female pied flycatchers Ficedula hypoleuca through nest removal and measured egg pigmentation and plasma antioxidant levels in relation with a control group. Experimental females showed a negative association between egg color and plasma antioxidant levels, while there was no relationship for control birds. This supports that egg pigmentation is costly in terms of general antioxidant defenses and suggests a tradeoff between the allocations to both traits. Furthermore, experimental females with more colorful eggs raised more fledglings, especially when breeding early. Controls did not show a relationship between egg color and reproductive success. Females laying more colorful eggs could have shifted their allocation decisions towards current reproduction, at the expense of their own antioxidant defenses. Our results highlight that blue egg coloration is a life-history trait, subject to tradeoffs with other attributes, and seems to be especially informative under harsh breeding conditions.     

Delayed plumage maturation in Lazuli buntings: tests of the female mimicry and status signalling hypotheses

The evolutionary importance of delayed plumage maturation (DPM) in passerines, the condition when more than 1 year is required to achieve adult-like coloration, remains highly contentious. Adaptive hypotheses propose that aggression from after 2nd-year (ASY) males or predation favors DPM in 2nd-year (SY) males, thereby increasing SY male survivorship or reproductive success. However, each hypothesis suggests a distinct selective mechanism explaining “how” this is accomplished. Alternatively, DPM may be a consequence of a nonadaptive molt constraint. We tested the female mimicry and status signalling hypotheses in territorial ASY male lazuli buntings (Passerinaamoena) using three sets of model presentation experiments. The female mimicry hypothesis proposes that dull SY male plumage deceptively mimics female plumage, and predicts that ASY males can not distinguish SY male from female plumage. The status signalling hypothesis proposes that dull SY male plumage honestly signals low competitive threat, and predicts that ASY males respond less aggressively to dull versus bright, ASY-like plumage. Contrary to the female mimicry hypothesis, ASY males distinguished between SY male and female plumage, as they were aggressive to SY male models exclusively and attempted to copulate with female models. Supporting the status signalling hypothesis, ASY males were significantly less aggressive to SY versus ASY male plumage. While DPM may result from a physiological constraint on bright SY male plumage, our results support the idea that dull plumage in an SY male's first breeding season may be maintained by selection to reduce aggression from ASY males, serving as a signal of competitive status. Received: 21 February 1997 / Accepted after revision: 16 June 1997     

Delayed plumage maturation in the orchard oriole (Icterus spurius): tests of winter adaptation hypotheses

Summary Explanations of delayed plumage maturation (DPM) in passerines have focused on potential breeding season advantages for non-definitive (subadult) plumage. In contrast, the molt constraint hypothesis (Rohwer and Butcher 1988) proposes that subadult plumage is a winter adaptation, increasing winter survivorship by decreasing intraspecific aggression (the winter status signaling hypothesis) or predation (the winter crypsis hypothesis). Under the molt constraint hypothesis, nondefinitive breeding plumage is non-adaptive, resulting from species-specific constraints on the number of feathers replaced during the pre-alternate (spring) molt. In studies conducted on an overwintering population of orchard orioles in Panama, I tested predictions of both the winter status signaling and molt constraint hypotheses. Contrary to predictions of winter status signaling, I found no evidence that subadult plumage reduces adult male aggression toward subadults. Agonistic encounters occurred at random with respect to plumage and the intensity of adult male/subadult male encounters was not lower than the intensity of encounters occurring within age classes. Contrary to the molt constraint hypothesis, I found no evidence that the number of feathers exchanged by subadults in their pre-alternate molt is the sole constraint on the development of subadult breeding plumage. The majority of feathers grown by molting subadult orioles during January and February were of non-definitive coloration. These results, together with results of earlier breeding season experiments which tested summer communication hypotheses for DPM in this species, suggest that subadult plumage in the orchard oriole may be non-adaptive and the result of constraints on plumage development. They also indicate, however, that the extent of the pre-alternate molt is not the sole source of that constraint.     

Testosterone upregulates lipoprotein status to control sexual attractiveness in a colorful songbird

A salient feature of many secondary sexual characteristics in animals is that their expression is controlled by sex-steroid hormones. However, for only a few types of ornaments do we know the precise molecular mechanism by which androgens like testosterone (T) enhance trait production. We studied the red carotenoid-based beak of male zebra finches (Taeniopygia guttata), which serves as a sexually selected trait and is thought to be T dependent. In previous research, we demonstrated that the extent to which male finches produce lipoproteins in the bloodstream, which bind carotenoid pigments acquired from the diet and transport them to peripheral tissues, regulates the accumulation of carotenoids in the body and beak pigmentation. Here we show that T acts to upregulate lipoprotein production and allows male zebra finches to display flashy, sexually attractive coloration. Levels of circulating T in blood positively and significantly predicted lipoprotein profile (as measured by cholesterol levels), blood carotenoid concentration, and beak color. Exogenous T administration elevated cholesterol and carotenoid status as well as beak redness. Last, experimental inhibition of T (using the anti-androgen flutamide) downregulated lipoprotein production and carotenoid circulation and faded the beak. This androgen- and lipoprotein-mediated system represents one of the more detailed physiological mechanisms underlying the development of a sex-steroid-dependent trait in animals.     

Independent effects of food and predator-mediated processes on annual fecundity in a songbird

We investigated the relative importance and interaction of ecological processes affecting annual fecundity in birds by simultaneously manipulating food availability and nest predation risk in a small songbird, the Wrentit (Chamaea fasciata). From 2000 to 2002 we provided supplemental food to individual Wrentit territories, and during 2002 we altered nest predation risk by providing supplemental food to their principal predators, Western Scrub-Jays (Aphelocoma californica). These experiments were conducted during a period of high interannual variation in rainfall, with 2002 being one of the driest years on record. Food-supplemented Wrentits in a normal predation environment produced an average of 0.54 more fledglings per year than control pairs over the three breeding seasons. During the feeding plus predation manipulation experiment, Wrentit food supplementation and lowered nest predation risk each independently increased the probability that a Wrentit pair would fledge young; however, the interaction between food supplementation and altered nest predation risk was not significant. Thus, even in an extreme drought year, both food and nest predation had equal but independent effects on reproductive success and annual fecundity. Combining supplemental food with reduced nest predation did not result in a synergistic increase in annual fecundity, primarily because Wrentits did not produce multiple broods. Our results suggest that whether food and predation have additive or synergistic effects on reproductive success depends on the life history of the species and the environment in which they live.     

Experimental evidence of a testosterone-induced shift from paternal to mating behaviour in a facultatively polygynous songbird

Previous studies have suggested that testosterone (T) profiles of male birds reflect a trade-off between mate attraction behaviours (requiring high T levels) and parental care activities (requiring low T levels). In this study, we experimentally elevated T levels of monogamous males in the facultatively polygynous European starling (Sturnus vulgaris), and compared mate attraction and paternal behaviour of T-treated males with those of controls (C-males). T-males significantly reduced their participation in incubation and fed nestlings significantly less often than C-males. Females paired to T-treated males did not compensate for their mate’s lower paternal effort. The observed reduction in a male’s investment in incubating the eggs was accompanied by an increased investment in typical female-attracting behaviours: T-males spent a significantly higher proportion of their time singing to attract additional females. They also occupied more additional nestboxes than C-males, although the differences just failed to be significant, and carried significantly more green nesting materials into an additional nestbox (a behaviour previously shown to serve a courtship function). T-males also behaved significantly more aggressively than C-males. During the nestling period, the frequency of mate-attracting behaviours by T-treated and control males no longer differed significantly. Despite the reduced paternal effort by T-males and the lack of compensation behaviour by females, hatching and breeding success did not differ significantly between T- and C-pairs. Received: 7 February 2000 / Revised: 10 August 2000 / Accepted: 3 September 2000     

Seasonal variation in male-female competition,cooperation and selfish hoarding in a monogamous songbird

Both cooperation and conflict between the sexes are commonplace in monogamous mating systems. However, little is known about how cooperation and competition varies seasonally in monogamous species that maintain permanent territories. We presented territorial pairs of male and female New Zealand robins (Petroica australis) with a large supply of insect prey at monthly intervals for 2 years. Behavioural observations after food presentation were then made to quantify seasonal and sexual differences in aggressive interactions over prey, prey acquisition rates, mate provisioning, offspring provisioning, selfish food hoarding and cache retrieval. Data were used to evaluate sex-specific behavioural strategies of mediating competition for food. Results showed that males aggressively excluded females from experimental food sources year-round. Females only accessed food sources when males left them unattended. Consequently, females acquired fewer prey than males. After controlling for differences in prey acquisition, both sexes consumed similar amounts of prey in the non-breeding season. Even though males aggressively excluded females from accessing food sources directly, males fed large amounts of prey to females during the breeding season. Both sexes provisioned young at similar rates. Males cached less prey than females in the breeding season but more prey than females in the non-breeding season. Females showed similar caching intensities year-round. Although males tried to defend their hoards, females frequently retrieved male-made caches. Overall, results showed that although New Zealand robins cooperate to raise offspring during the breeding season, conflict between the sexes occurs year-round. Males and females display different behavioural strategies to gain access to experimental food sources, which appear to lessen male–female competition for food and evenly distribute food resources between the sexes.     

Unusually high levels of extrapair paternity in a duetting songbird with long-term pair bonds

Previous studies suggest that extrapair young are very rare or absent in socially monogamous avian species that produce vocal duets. These results are generally consistent with functional hypotheses suggesting that duets may signal commitment between partners, or aid males as a paternity guard to ensure genetic as well as social monogamy. Additionally, species that exhibit social monogamy with the same partner across multiple breeding seasons tend to exhibit low levels of extrapair paternity, so duetting species that mate for life may be particularly likely to exhibit genetic monogamy. This study examined the social and genetic mating systems of California towhees (Pipilo crissalis), a duetting species thought to have life-long pair bonds. Observation of a color-banded population confirmed that California towhees exhibit long-term social monogamy. Known social families were genotyped at four microsatellite loci with high allelic diversity. Unexpectedly, paternity exclusion indicated that at least 13 of 31 (42%) nests contained extrapair young. All chicks exhibited maternal alleles, but 21 of 81 (26%) young were not the offspring of social fathers. Thus, in contrast to previous work, this study documents high frequencies of extrapair young among socially monogamous duetting birds with long-term pair bonds.     

Tracking migration routes and the annual cycle of a trans-Sahara songbird migrant

Movement ecology studies have highlighted the importance of individual-based research. As tracking devices have not been applicable for identifying year-around movements of small birds until recently, migration routes of such species relied on visual observations and ring recoveries. Within the Palaearctic–African migration system, loop migration seems to be the overall migration pattern. The interindividual variations within species-specific migration routes are, however, unknown. Here, we track the individual migration routes and annual cycles of male Northern Wheatears Oenanthe oenanthe, a trans-Sahara songbird migrant from a German breeding population with light-level geolocators. Two migrated most likely via Spain towards western Africa but returned via Corsica/Sardinia, while two others seemed to migrate via Sardinia and Corsica in autumn and via Spain and France in spring (loop migration). The fifth took presumably the same route via France and the Balearics in both seasons. All birds wintered in the Sahel zone of western Africa. Overall migration distances for autumn and spring were similar (about 4,100?km), whereas the overall migratory speed was generally higher in spring (126?km?day^?1) than in autumn (88?km?day^?1). Birds spent about 130?days at the breeding area and 147?days at the wintering grounds.     

Phenotype-dependent arrival time and its consequences in a migratory bird

Arrival times for migratory animals can be viewed as the result of an optimization process of costs and benefits of early arrival, and when the cost and benefit functions of early arrival depend on phenotypic quality, this will result in phenotype-dependent optimal arrival times. This hypothesis was tested for males of the migratory and sexually size-dimorphic barn swallow Hirundo rustica. The major cost of early arrival is poor environmental conditions which resulted in mortality of short-tailed early-arriving males in one year. The major benefits of early arrival are higher mating success, enhanced reproductive success, improved recruitment rates for offspring, and enhanced quality of the mate acquired. Annual variation in male arrival date is related to weather conditions at the breeding grounds, but also to some extent to weather conditions in the African winter quarters. Individual variation in arrival time can be explained by phenotype-dependent cost and benefit functions of early arrival. Male barn swallows with long tail ornaments arrived earlier than short-tailed males. The costs of early arrival should be particularly high under poor environmental conditions and lead to a stronger negative relationship between arrival date and phenotypic quality in years with poor environmental conditions. This prediction was confirmed by a stronger negative relationship between male tail length and date of arrival in years when arrival was relatively late because of poor weather. A female preference for early-arriving males may result in acquisition of good genes for optimal migratory behaviour, if migratory direction and extent have a genetic basis as shown in a number of different bird species.     

A genoscape-network model for conservation prioritization in a migratory bird

Migratory animals are declining worldwide and coordinated conservation efforts are needed to reverse current trends. We devised a novel genoscape-network model that combines genetic analyses with species distribution modeling and demographic data to overcome challenges with conceptualizing alternative risk factors in migratory species across their full annual cycle. We applied our method to the long distance, Neotropical migratory bird, Wilson's Warbler (Cardellina pusilla). Despite a lack of data from some wintering locations, we demonstrated how the results can be used to help prioritize conservation of breeding and wintering areas. For example, we showed that when genetic, demographic, and network modeling results were considered together it became clear that conservation recommendations will differ depending on whether the goal is to preserve unique genetic lineages or the largest number of birds per unit area. More specifically, if preservation of genetic lineages is the goal, then limited resources should be focused on preserving habitat in the California Sierra, Basin Rockies, or Coastal California, where the 3 most vulnerable genetic lineages breed, or in western Mexico, where 2 of the 3 most vulnerable lineages overwinter. Alternatively, if preservation of the largest number of individuals per unit area is the goal, then limited conservation dollars should be placed in the Pacific Northwest or Central America, where densities are estimated to be the highest. Overall, our results demonstrated the utility of adopting a genetically based network model for integrating multiple types of data across vast geographic scales and better inform conservation decision-making for migratory animals.     

Juvenile dispersal affects straying behaviors of adults in a migratory population

The resilience of organisms to large-scale environmental and climatic change depends, in part, upon the ability to colonize and occupy new habitats. While previous efforts to describe homing, or natal site fidelity, of migratory organisms have been hindered by the confounding effects of fragmented landscapes and management practices, realistic conservation efforts must include considerations of the behavioral diversity represented by animal movements and dispersal. Herein, we quantify straying away from natal origins by adult chinook salmon (Oncorhynchus tshawytscha) in a wild population that inhabits a pristine wilderness basin. Using natural isotopic signatures (7Sr/86Sr) to reconstruct the migratory behaviors of unhandled individuals over their entire life cycle, we identified ecological and behavioral factors influencing the propensity to stray. Our results indicate that natal site fidelity is scale dependent, ranging from 55% at -1-km distances to 87% at longer (> 10-km scale) distances, and juvenile dispersal and sex highly influence straying occurrence. These findings lend support for the conservation of behavioral diversity for population persistence, and we propose straying as a mechanism for maintaining genetic diversity at low population densities.     

Elevated testosterone advances onset of migratory restlessness in a nearctic-neotropical landbird

The primary cue for initiation of spring migratory restlessness (Zugunruhe) in landbird migrants is photoperiod. Gonadal hormones are known to have a role in the regulation of migratory disposition; however, the extent of their effect is not well understood. We examined the impact of exogenous testosterone on the onset of migratory restlessness in gray catbirds (Dumetella carolinensis). Catbirds were stratified by sex and randomly assigned to two rooms; individuals in one room were photoadvanced to initiate migratory restlessness and the other room was maintained on a 12:12 light/dark photoperiod. Each room had three groups (n?=?10/group); males with testosterone implants, males with empty implants, and females. We predicted that in the photoadvanced room males with testosterone implants would initiate migratory activity earlier than empty-implanted males. We found that in the photoadvanced group, testosterone-implanted males initiated migration 2 weeks prior to empty-implanted males, and 3 weeks prior to females. In the non-photoadvanced males, the testosterone-implanted males initiated migration at the same time as the corresponding group in the photoadvanced room, while the empty-implanted males and females did not exhibit Zugunruhe. Our results illustrate that elevated testosterone can advance the onset of Zugunruhe, even in the absence of an extended photoperiod. Additionally, the onset of migratory restlessness observed in the photoadvanced, non-testosterone-implanted males and females further supports the importance of photoperiod as a cue for migratory restlessness. An interesting observation was the intersexual differences in the onset of migratory activity in gray catbirds, a species not previously known to exhibit protandry.     

Characterizing the spatial structure of songbird cultures

Recent advances have shown that human-driven habitat transformations can affect the cultural attributes of animal populations in addition to their genetic integrity and dynamics. Here I propose using the song of oscine birds for identifying the cultural spatial structure of bird populations and highlighting critical thresholds associated with habitat fragmentation. I studied song variation over a wide geographical scale in a small and endangered passerine, the Dupont's Lark Chersophilus duponti, focusing on (1) cultural population structure, to determine a statistical representation of spatial variation in song and identify cultural units, and (2) the minimum patch size needed for an individual to develop a stable repertoire. I found that overall song diversity depends on variation among populations (beta-cultural diversity). Abrupt thresholds occurred in the relationships between individual song dissimilarity and geographic distance, and between individual song diversity and patch area. Spatial autocorrelation analysis showed that populations located as little as 5 km apart may have independently evolved their song traditions. Song diversity stabilized in patches as small as 100 ha supporting as few as 8-20 males. Song repertoires of smaller patches were significantly poorer. Almost one-quarter of the study populations inhabited patches <100 ha, and their cultural traditions appear to have eroded. The analysis of spatial patterns in birdsong may be a useful tool for detecting subpopulations prone to extinction.     

Sexual selection and cuckoldry in a monogamous songbird: implications for sexual selection theory

Sexual selection is generally assumed to be weaker in monogamous than in polygynous animals. Recently, though, extra-pair fertilizations have been hailed as an important force in generating variance in reproductive success among males in socially monogamous species, thereby increasing the intensity of sexual selection. To see if extra-pair copulations contribute to variance in male reproductive success in the house finch (Carpodacus mexicanus), we used DNA fingerprinting to determine the paternity of chicks from 35 nests. This species is a socially monogamous passerine in which plumage brightness serves as a sexually selected indicator of male quality. Out of 119, nestlings 10 (8.3%) were fathered by a male other than the attending male, but cuckoldry occurred randomly with respect to the plumage colouration, size, or age of the attending male. Thus extra-pair fertilizations do not generate variance in male reproductive success with respect to plumage colour. On the other hand, a strongly male-biased sex ratio and asynchronous breeding by females may generate substantial variance in male reproductive success and could explain the evolution of ornamental colouration.     

Cowbird removals unexpectedly increase productivity of a brood parasite and the songbird host

Generalist brood parasites reduce productivity and population growth of avian hosts and have been implicated in population declines of several songbirds of conservation concern. To estimate the demographic effects of brood parasitism on Bell's Vireos (Vireo bellii), we removed Brown-headed Cowbirds (Molothrus ater) in a replicated switchback experimental design. Cowbird removals decreased parasitism frequency from 77% and 85% at unmanipulated plots to 58% and 47% at removal plots in 2004 and 2005, respectively. Vireo productivity per pair was higher at cowbird removal plots when years were pooled (mean = 2.6 +/- 0.2 [SE] young per pair) compared to unmanipulated plots (1.2 +/- 0.1). Nest desertion frequency was lower at cowbird removal plots (35% of parasitized nests) compared to unmanipulated plots (69%) because removal of host eggs was the proximate cue for nest desertion, and vireos experienced lower rates of egg loss at cowbird removal plots. Nest success was higher among unparasitized than parasitized nests, and parasitized nests at cowbird removal plots had a higher probability of success than parasitized nests at unmanipulated plots. Unexpectedly, cowbird productivity from vireo pairs was higher at cowbird removal plots (mean = 0.3 +/- 0.06 young per pair) than at unmanipulated plots (0.1 +/- 0.03) because fewer parasitized nests were deserted and the probability of nest success was higher. Our study provides the first evidence that increases in cowbird productivity may be an unintended consequence of cowbird control programs, especially during the initial years of trapping when parasitism may only be moderately reduced. Thus, understanding the demographic impacts of cowbird removals requires an informed understanding of the behavioral ecology of host-parasite interactions.