Egg size and asymmetric sibling rivalry in red-winged blackbirds

How big to make an egg is a life history decision that in birds is made coincident with a series of other similar decisions (how many eggs to have, whether to fortify them with maternally derived hormones or immune system boosters, whether to hatch the eggs synchronously or asynchronously). Though within-population variation in egg size in birds has been well studied, its adaptive significance, if any, is unclear. Here we examine within-population variation in egg size in relation to asymmetric sibling rivalry in a 17-year study of red-winged blackbirds (Agelaius phoeniceus), an altricial songbird. Egg mass showed a twofold range of variation, with roughly 80% of the variation occurring across clutches. By commencing incubation before the clutch is complete, mothers create advantaged core and disadvantaged marginal elements within their brood. Previous work on this system has shown that sibling competition is asymmetric, and that core offspring enjoy priority access to food, and as a consequence show higher growth and lower mortality than marginal offspring. Here we examine the effect of initial egg size on nestling growth and survival in relation to these competitive asymmetries. Egg mass was strongly linked to hatchling mass, and remained significantly related to the mass of both core and marginal nestlings; the effect of egg size was stronger for core offspring early in the nestling period, but the disparity between core and marginal nestlings narrowed as they approached fledging age, and slower growing marginals fell victim to brood reduction. The effect of egg mass on survival differed dramatically between core and marginal nestlings. Egg mass was significantly related to the survival of marginal but not core nestlings: below average egg mass was associated primarily with very early mortality. Asymmetric sibling competition is clearly a strong determinant of the consequences of egg size variation.     

Experimental analyses of intra- and intersexual competition in red-winged blackbirds

The roles of intra- and intersexual selection in determiningharem size for many polygynous birds have proved difficult toseparate. We designed experiments to feed only one sex to distinguishbetween intra- and intersexual selection in polygynous red-wingedblackbirds. We then tested whether choices made by females benefitfemales. Our experiments were conducted on a number of differentmarines, so we could evaluate whether females were choosingmales within marshes or whether they are choosing their breedingsituation primarily by making choices between different marshes.Females choosing between marshes strongly preferred marsheswith males that had supplemental food and experimentally increaseddisplay rates. At the marsh level, some male displays were predictorsof larger harem size and of greater nesting success. But whilefemale choice was very strong among marshes, we detected onlyweak female preference for individual male traits within marshes.These results suggest that problems of scale probably affectstudies of mate choice in avian breeding systems in ways that,to the best of our knowledge, have not previously been investigatedby behavioral ecologists     

Male red-winged blackbirds distrust unreliable and sexually attractive neighbours

In many species, territorial neighbours fight to establish their mutual border and then develop a truce, known as the dear-enemy phenomenon, characterized by reduced vigilance and aggression along the border. We present evidence that among male red-winged blackbirds (Agelaius phoeniceus) the dear-enemy relationship is a form of reciprocal conditional cooperation that is stabilized, at least in part, by retaliation against cheaters. Simulated intrusions by randomly chosen neighbours were punished by a targeted increase in vigilance and aggression that persists for days. We interpret this increase in vigilance towards trespassers as a manifestation of distrust. The conditional decrease in vigilance and aggression is tempered by each neighbour's probability of cuckolding the focal male. Male red-winged blackbirds maintained greater vigilance and aggression towards sexually attractive neighbours that were more successful at extra-pair fertilizations (EPFs). It is unlikely that males directly observed neighbours copulating with their mates. They were more likely to assess a neighbour's ability to achieve extra-pair copulations using surrogate cues that correlate with success at EPFs, including body size. Our results suggest that red-winged blackbirds use rules that incorporate their neighbour's behaviour and quality in their territorial interactions with one another. Our results expand our understanding of cooperation for animals and for humans as well.     

Breeding synchrony and extrapair fertilizations in two populations of red-winged blackbirds

We tested the relationship between synchrony of breeding andthe frequency of extrapair fertilizations (EPFs) in two populationsof red-winged blackbirds known to differ in female extrapairbehavior. We found no association between the number of simultaneouslyfertilizable females (temporal neighbors) and EPF rate in eitherpopulation, although a significant difference between populationsin the direction of this relationship (positive where femalesinitiated extrapair copulations and negative where males initiatedthem) suggested a modest difference in the influence of synchrony.Males losing offspring to EPFs tended to have more fertilizablefemales at that time than the actual sires in some analysesbut not in others. We also tested several assumptions underlyingtwo competing hypotheses for the effects of synchrony. We foundno evidence that females pursued extrapair copulations moreoften when other females were synchronous. Rather, females weremore likely to gain EFFs with exirapatr males whose social mateswere not yet building their nests. Synchrony also did not consistentlyaffect male pursuit of exirapair copulations or achievementof EPFs. These results suggest that timing of breeding has someeffects on extrapair activity, but that those effects are bothrelatively weak and influenced by other factors that vary betweenyears or populations.     

The effects of androgen and antiandrogen on dominance and aggressiveness in male red-winged blackbirds

Male red-winged blackbirds were given implants of testosterone or flutamide in order to manipulate the level of androgen reaching receptor sites. Implants of cholesterol were given as controls. Radioimmunoassay showed higher levels of testosterone and DHT in the plasma of testosterone-implanted birds than in the plasma of the pooled flutamide-implanted and cholesterol-implanted birds. Dominance was significantly correlated with effective androgen level. Androgen level was not correlated with rates of supplantations per subordinate but was correlated with rates of escalated attacks per subordinate. Thus androgen level influences dominance in male red-winged blackbirds by influencing one component of aggressiveness.     

A test of the good-genes-as-heterozygosity hypothesis using red-winged blackbirds

Brown recently proposed that the "good genes" that females pursuewhen choosing mates may be individual heterozygosity becausemore heterozygous mates sire offspring with higher fitness.Further, because heterozygosity might enhance developmentalstability, males with more heterozygosity are recognized bythe reduced fluctuating asymmetry (FA) of their bilaterallypaired traits. We used a point sample of 67 male red-winged blackbirds(Agelaius phoeniceus) to test two predictions of this hypothesis:(1) males with more heterozygosity have higher fitness, and(2) males with more heterozygosity have lower FA. We identified7 polymorphic loci from an initial screening of 16 enzymes;32 individuals were completely homozygous, and 35 individualswere heterozygous at at least 1 locus. Larger and older malesrealized higher mating success in this population, but neither sizenor age was related to heterozygosity. Heterozygous males werenot in better condition than homozygous males, nor were theyless infected by hematozoa, lice, or mites. Among 1-year oldmales, epaulet length did not differ between homozygotes andheterozygotes, but among older males, heterozygotes did havelonger epaulets. Homozygotes and heterozygotes did not differin their mean FA scores for nine individual characters. Althoughthe two groups of males did differ in composite FA, heterozygousmales were less symmetrical. Interestingly, this differencewas attributable to a single allele at the PGM-3 locus. Combinedwith previous results showing that FA was generally unrelatedto male health, viability, parental care, social dominance,or mating success, the present results indicate that Brown's hypothesisdoes not explain mate choice or male quality in this populationof red-winged blackbirds.     

Reproductive consequences of bilateral asymmetry for individual male red-winged blackbirds

We used morphological and breeding data from a 2-year fieldstudy of red-winged blackbirds (Agelaius phoeniceus) to testthe hypothesis that males characterized by low levels of bilateralasymmetry (i.e., high developmental competence) realize a reproductiveadvantage. Specifically, we evaluated each of several distinctcomponents of male reproductive success relative to asymmetrymeasures made on five bilaterally paired characters. Resultsof a male removal experiment generally failed to support theprediction that symmetry would be associated with success incompetition for access to breeding territories: establishedterritory owners and nonterritorial replacement males were effectivelyindistinguishable in this regard. Similarly, there was no indicationthat symmetrical males were more likely to establish territoriesin high-quality marsh habitat than in marginal upland fieldhabitat. Finally, monitoring of breeding activity in high-qualityhabitat revealed that male symmetry was generally unrelatedto recruitment of social mates (i.e., harem size), the productivityof those mates (average female reproductive success), withinpairpaternity (assessed using DNA-based analysis of parentage),or extrapair mating success. Collectively these results indicatethat symmetry is not an important determinant of reproductivesuccess among individual male red-winged blackbirds. This observation,in combination with the results of several other recent investigations,suggests that the fitness consequences of subtle departuresfrom perfect symmetry may be less significant and/or less ubiquitousthan initially suggested.     

Tests of spatial and temporal factors influencing extra-pair paternity in red-winged blackbirds

Extra-pair paternity (EPP) is a widespread and highly variable reproductive phenomenon in birds. We tested the effects of habitat, spatial factors, and timing of breeding on the occurrence of EPP in red-winged blackbirds (Agelaius phoeniceus). We used PCR-amplified microsatellites to assess the paternity of 1479 nestlings from 537 broods on 235 territories over four breeding seasons. Over 4 years, 40% of nestlings were extra-pair. At least 27% of actual sires were non-neighbours, suggesting that males or females interacted over longer distances than in other populations of red-winged blackbirds. The level of EPP was significantly clumped within broods and males but not within females across broods. EPP was negatively related to the area of a male's territory. The spatial proximity of a female's nest to the territory boundary had no effect on total EPP, but tended to increase the probability of an EPP by a nearby male. We found no influence on EPP of the type of habitat on the territory or the level of nesting activity nearby. The time in the season when a nest was started and the synchrony of breeding also had no significant effect on the level of EPP. The age of the male, the age of his neighbours, and the interaction between the two had no effect on total EPP. However, older males were less likely to have an offspring sired by a neighbour on their territory. Males with older neighbours were also less likely to have offspring sired by a neighbour, particularly if they were new territory owners. The high variability in who gained and lost paternity, and the limited impact of spatial and temporal factors influencing it, have some interesting implications for theories seeking to explain mating patterns.     

Experimental confirmation of the polygyny threshold model for red-winged blackbirds

The polygyny threshold model assumes that polygynous mating is costly to females and proposes that females pay the cost of polygyny only when compensated by obtaining a superior territory or male. We present, to the authors' knowledge, the first experimental field test to demonstrate that females trade mating status against territory quality as proposed by this hypothesis. Previous work has shown that female red-winged blackbirds (Agelaius phoeniceus) in Ontario prefer settling with unmated males and that this preference is adaptive because polygynous mating status lowers female reproductive success. Other evidence suggests that nesting over water increases the reproductive success of female red-winged blackbirds. Here we describe an experiment in which females were given choices between two adjacent territories, one owned by an unmated male without any over-water nesting sites and the other by an already-mated male with over-water sites. Females overwhelmingly preferred the already-mated males, demonstrating that superior territory quality can reverse preferences based on mating status and supporting the polygyny threshold model as the explanation for polygyny in this population.     

Polygyny and extrapair fertilizations in eastern red-winged blackbirds (Agelaius phoeniceus)

Parentage of nestling red-winged blackbirds (Agelaius phoeniceus)from an eastern population was determinedusing DNA fingerprintingtechniques. Of 235 nestlings surveyed, 58 had fingerprints excludingthemale, but none excluded the female tending the nest. Data onpairing status during the female's fertilizable period was availablefor 232 offspring; 55 (25%of 1988 nestlings, 23% of 1989 nestlings)of those were sired through extrapair copulations. Of these55 offspring, 33 could be assigned to nearby territory holders;16 of the remaining nestlings may have been sired by nearbymales that were not captured. During both years, 44% of territorialmales had more than one female nesting simultaneously on theirterritory. The number of extrapair fertilizations gained bymales increased significandy with harem size in 1 year. Paternity(die proportion of nesdings on the territory sired by die territoryholder) showed a positive but nonsignificant increase widi haremsize in bodi years. There was no apparent cost in paternityfor males guarding two or more fertilizable females at the sametime. The broods of females that were fertilizable at die sametime anodier female was setding on die same territory tendedto have a greater proportion of extrapair fertilizations (0.37)than did die broods of odier females within harems (0.15). Establishedfertilizable females were chased significantly more by die territoryowner and by extrapair males when a new female was setding.There were no associations between a male's paternity or successat gaining extrapair fertilizations and his age or color-bandcombination. Overall, extrapair fertilizations had litde effecton die relationship between fledgling success and harem sizeand appeared to have a minimal impact on die overall intensityof sexual selection on males.     

Secondary sexual traits, parasites, and polygyny in red-winged blackbirds, Agelaius phoeniceus

Data from a 3-year study of red-winged blackbirds (Agelaiusphoeniceus) were used to test the hypothesis that parasites(in this case, haematozoa) reduce male fitness and cause diminishedexpression of secondary sexual traits, which, in turn, are usedby females to select parasite-free males as mates. There wasno evidence indicating a fitness cost to being parasitized becauseparasitized males were as likely as unparasitized males to acquirea territory and to survive from one year to the next. Similarly,parasitized and unparasitized females did not differ with regardto how early they started nesting, how many eggs they laid,or their year-to-year survival. Secondary sexual traits, particularlyintrasexual aggression, did reliably (>80%) reveal the parasitestatus of males. Plumage and morphological traits also alloweddiscrimination of parasitized and unparasitized females. However,apparent mating patterns were unrelated to either the males'or the females' parasite status. Only if genetic analyses revealthat unparasitized males actually realize higher productivesuccess will these results potentially provide support for theparasite hypothesis of sexual selection.     

Resource availability, breeding site selection, and reproductive success of red-winged blackbirds

Red-winged blackbirds are polygynous and show strong breeding site preferences, but it is unclear which environmental factors regulate their reproductive success and are ultimately responsible for shaping their patterns of habitat selection and their mating system. We evaluated the effect of variation in insect emergence rates on the reproductive success of male and female redwings nesting on replicate ponds. The number of male and female redwings that settled on a pond varied two- to three-fold among ponds, but was not related to insect emergence rates. Insect emergence rates had a positive effect on the number of nestlings successfully fledged by females, the number of nestlings fledged from male territories, and on the mass of nestlings at fledging. Typha stem density also varied widely among ponds, and was positively related to male and female settling density and mass of nestlings at fledging, but not to the number of nestlings fledged by females or males. We conclude that alternative breeding sites differ in their ability to support redwing reproduction, and that the availability of emerging odonates is an important environmental factor influencing the reproductive success of both male and female red-winged blackbirds. Received: 31 March 1997 / Accepted: 3 July 1997     

Alterations in tissue aerobic capacity may play a role in premigratory fattening in shorebirds

Migratory shorebirds show regulated seasonal increases in body mass (BM) even in captivity, consisting primarily, but not exclusively, of fat. We examined whether captive red knot (Calidris canutus) exhibited seasonal alterations in mitochondrial volume (liver, pectoral muscle) and/or succinate dehydrogenase (SDH) activity (liver, pectoral muscle, heart, small intestine) during three distinct life-cycle stages: stable BM, spring peak in BM, and as BM rapidly declined after the spring peak. Mitochondrial volume in liver and pectoral muscle and SDH activity in liver and heart did not alter with life-cycle stage. However, red knot undergoing premigratory fattening exhibited significantly lower pectoral muscle SDH activity in concert with significantly elevated activity in the small intestine compared with the other two time-points, suggesting that tissue metabolic rate alters with life-cycle stage. The increased intestinal SDH activity may indicate an elevation in energy assimilation at a time when intestine hypertrophy occurs, thus maximizing BM increase prior to putative migration. The concomitant decrease in pectoral muscle activity may act to reduce overall metabolic rate, or at least help counter the elevation in intestinal mass-specific metabolic rate. Both tissues hypertrophy prior to migration in wild red knot, but hypertrophy of the intestine precedes that of pectoral muscle. Indeed, it appears that the intestinal mass undergoes atrophy by the time pectoral muscle hypertrophy occurs in wild red knot. Thus, physiological adjustments in tissue metabolism may be an important factor in the life-history strategies of migrating shorebirds.     

The hormonal regulation of de-etiolation

De-etiolation involves a number of phenotypic changes as the plants shift from a dark-grown (etiolated) to a light-grown (de-etiolated) morphology. Whilst these light-induced, morphological changes are thought to be mediated by plant hormones, the precise mechanism/s are not yet fully understood. Here we provide further direct evidence that gibberellins (GAs) may play an important role in de-etiolation, because a similar light-induced reduction in bioactive GA levels was detected in barley (Hordeum vulgare L.), Arabidopsis (Arabidopsis thaliana L.), and pea (Pisum sativum L.). This is indicative of a highly conserved, negative-regulatory role for GAs in de-etiolation, in a range of taxonomically diverse species. In contrast, we found no direct evidence of a reduction in brassinosteroid (BR) levels during de-etiolation in any of these species. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.