Comparative hatching characteristics of nonparasitic and parasitic icterids: is the hatching of cowbird young constrained by an unusually thick eggshell?

Studies of avian brood parasitism have provided some of the best evidence of coevolutionary arms races. One of the parasitic adaptations exhibited by cuckoos (family Cuculidae) and cowbirds (family Icteridae) is the development of a thick eggshell, presumably to avoid damage from puncture ejection by some hosts and/or accidental damage during egg-laying in the host nest. However, it is unknown whether this trait constrains the hatching of parasitic young. The differences in hatching characteristics between the host red-winged blackbird (“redwing;” Agelaius phoeniceus) and the parasitic brown-headed cowbird (Molothrus ater) were examined. Prehatched cowbird young were found to spend more time hatching than pre-hatched redwing young and to emit click sounds at a greater rate in relation to pulmonary respiration than pre-hatched redwing young. However, cowbird hatchlings appear to have evolved other hatching-related traits that may compensate for the greater hatching effort, such as body parts and an egg tooth that are relatively large compared with those of redwing hatchlings.     

Partial host fidelity in nest selection by the shiny cowbird (Molothrus bonariensis), a highly generalist avian brood parasite

Obligate avian brood parasites can be host specialists or host generalists. In turn, individual females within generalist brood parasites may themselves be host specialists or generalists. The shiny cowbird Molothrus bonariensis is an extreme generalist, but little is known about individual female host fidelity. We examined variation in mitochondrial control region sequences from cowbird chicks found in nests of four common Argentinean hosts. Haplotype frequency distributions differed among cowbird chicks from nests of these hosts, primarily because eggs laid in nests of house wrens Troglodytes aedon differed genetically from those laid in nests of the other three hosts (chalk-browed mockingbird Mimus saturninus, brown-and-yellow marshbird Pseudoleistes virescens, and rufous-collared sparrow Zonotrichia capensis). These differences in a maternally inherited marker indicate the presence of a nonrandom laying behaviour in the females of this otherwise generalist brood parasite, which may be guided by choice for nest type, as house wrens nest in cavities whereas the other three species are open cup nesters.     

Brood‐parasite‐induced female‐biased mortality affects songbird demography: negative implications for conservation

Parasites, of all sorts, can profoundly affect host population dynamics. Parasites commonly cause sex‐biased mortality and this can add to their impact. Female‐biased mortality in particular can destabilize dynamics and promote population collapse. We previously reported in a correlative study that brown‐headed cowbird Molothrus ater brood parasitism of song sparrows Melospiza melodia appears to cause female‐biased host nestling mortality. Here, we report results from ‘infestation’ and ‘de‐infestation’ experiments designed to test whether brood parasitism causes female‐biased mortality, and we document the resulting demographic impact using a simulation model. Experimental cowbird infestation of song sparrow nests halved the proportion of female host nestlings (0.31±0.07 vs 0.59±0.06; infested vs unparasitized nests at day 6) replicating the halving reported in naturally cowbird‐parasitized nests (0.28±0.01 vs 0.57±0.05; parasitized vs unparasitized). De‐infestation of naturally cowbird‐parasitized nests in turn wholly eliminated any effect on the proportion of female host nestlings (0.53±0.13 vs 0.54±0.06; de‐infested vs unparasitized) confirming that brood parasitism is the cause. This halving of the proportion of females fledging is likely to be as significant as nest predation in affecting population dynamics, based on the elasticities derived from our demographic model (–0.50 vs –0.59). Experimental infestation reduced the testosterone levels, begging behaviour, and body mass of six day old female host nestlings, whereas males were largely unaffected, suggesting that it is the exacerbation of intra‐brood competition that may be primarily responsible for the resulting female‐biased mortality. The brown‐headed cowbird is invasive in most of North America and has been implicated in regional population declines of many native species. We suggest that female‐biased host offspring mortality is likely to be commonplace among the 144 host species the cowbird successfully parasitizes, and we discuss the negative implications for songbird conservation, given the projected demographic impact.     

Seasonal changes in song nuclei and song behavior in Gambel's white-crowned sparrows

In males of several songbird species, the morphology of forebrain nuclei that control song changes seasonally. The only seasonally breeding songbird in which seasonal changes in the structure of song control nuclei have been reported not to occur is the nonmigratory Nuttall's subspecies of white-crowned sparrow. In the present study, we manipulated photoperiod and plasma testosterone concentrations in captive male white-crowned sparrows of the migratory Gambel's subspeices. Males exposed to photoperiods and plasma testosterone concentrations typical of those experienced by wild breeding males had larger song control nuclei than males held on a winter photoperiod. We also found seasonal change in stereotypy of spectral and temporal parameters of song in wild Gambel's white-crowned sparrows. We hypothesize that seasonal changes in song control nuclei may correlate with seasonal changes in song stereotypy. © 1995 John Wiley & Sons, Inc.     

Comparison of eggshell porosity and estimated gas flux between the brown‐headed cowbird and two common hosts

The brown‐headed cowbird Molothrus ater is a brood‐parasite that lays eggs in nests of a wide range of host species, including the closely‐related red‐winged blackbird Agelaius phoeniceus and the dickcissel Spiza americana. Although cowbird eggs have accelerated development and hatch sooner than similar‐sized host eggs, this development takes place within a thickened eggshell that could impede gas flux to the developing embryo. We tested the hypothesis that the accelerated development of the cowbird embryo relative to hosts is enabled by an increase in eggshell porosity that allows increased fluxes of respiratory gases to and from the developing embryo. We found cowbird eggshell thickness was significantly greater than the eggshells of these two common hosts. Although the number of pores per egg was similar among all three species, the total pore area per egg in cowbirds was significantly greater than that of either host, despite having a smaller eggshell surface area than the red‐winged blackbird. Cowbird egg pore area was 1.9×larger than that of the red‐winged blackbird. Cowbird eggshells had a significantly greater gas flux than those of the red‐winged blackbird and the dickcissel. When conductance was normalized to published values of egg mass, cowbird eggs had a higher mass‐specific conductance than red‐winged blackbird or dickcissel eggs. These results are consistent with the hypothesis that the rapid development of brown‐headed cowbird embryos is facilitated by increased eggshell porosity, and that changes in eggshell porosity represent an adaptation that enables cowbird eggs to hatch earlier than equivalently‐sized host eggs.     

Maternally derived yolk testosterone enhances the development of the hatching muscle in the red-winged blackbird Agelaius phoeniceus

Hatching asynchrony in avian species often leads to the formation of a size hierarchy that places last-hatched nestlings at a significant disadvantage. The hatching muscle (musculus complexus) is responsible for breaking the shell during hatching and for dorsal flexion of the neck during begging. An increase in its strength in last-hatched nestlings could mitigate the effects of hatching asynchrony by reducing the time required for hatching or enhancing the effectiveness of begging for parentally delivered food or both. We have previously found that yolk testosterone concentration increases with laying order in the red-winged blackbird Agelaius phoeniceus. In this study, we investigated the hypothesis that yolk testosterone has anabolic effects on the development of the complexus, thereby influencing competition among asynchronously hatched nestlings. We found that both yolk testosterone concentration and relative complexus mass (complexus mass/nestling body mass) increased with laying order and that these two variables were positively correlated in both newly hatched nestlings and in two-day-old broods. Moreover, direct injections of testosterone into egg yolks resulted in an increase in relative complexus mass, while injections of flutamide, a testosterone antagonist, resulted in a decrease in relative complexus mass. Neither yolk testosterone concentration nor relative complexus mass differed between male and female nestlings.     

Widespread dispersal of Borrelia burgdorferi-infected ticks collected from songbirds across Canada

Millions of Lyme disease vector ticks are dispersed annually by songbirds across Canada, but often overlooked as the source of infection. For clarity on vector distribution, we sampled 481 ticks (12 species and 3 undetermined ticks) from 211 songbirds (42 species/subspecies) nationwide. Using PCR, 52 (29.5%) of 176 Ixodes ticks tested were positive for the Lyme disease spirochete, Borrelia burgdorferi s.l. Immature blacklegged ticks, Ixodes scapularis , collected from infested songbirds had a B. burgdorferi infection prevalence of 36% (larvae, 48%; nymphs, 31%). Notably, Ixodes affinis is reported in Canada for the first time and, similarly, Ixodes auritulus for the initial time in the Yukon. Firsts for bird-parasitizing ticks include I. scapularis in Quebec and Saskatchewan. We provide the first records of 3 tick species cofeeding on passerines (song sparrow, Swainson's thrush). New host records reveal I. scapularis on the blackpoll warbler and Nashville warbler. We furnish the following first Canadian reports of B. burgdorferi-positive ticks: I. scapularis on chipping sparrow, house wren, indigo bunting; I. auritulus on Bewick's wren; and I. spinipalpis on a Bewick's wren and song sparrow. First records of B. burgdorferi-infected ticks on songbirds include the following: the rabbit-associated tick, Ixodes dentatus, in western Canada; I. scapularis in Quebec, Saskatchewan, northern New Brunswick, northern Ontario; and Ixodes spinipalpis (collected in British Columbia). The presence of B. burgdorferi in Ixodes larvae suggests reservoir competency in 9 passerines (Bewick's wren, common yellowthroat, dark-eyed junco, Oregon junco, red-winged blackbird, song sparrow, Swainson's thrush, swamp sparrow, and white-throated sparrow). We report transstadial transmission (larva to nymph) of B. burgdorferi in I. auritulus. Data suggest a possible 4-tick, i.e., I. angustus, I. auritulus, I. pacificus, and I. spinipalpis, enzootic cycle of B. burgdorferi on Vancouver Island, British Columbia. Our results suggest that songbirds infested with B. burgdorferi-infected ticks have the potential to start new tick populations endemic for Lyme disease. Because songbirds disperse B. burgdorferi-infected ticks outside their anticipated range, health-care providers are advised that people can contract Lyme disease locally without any history of travel.     

Hatching asynchrony, nestling competition, and the cost of interspecific brood parasitism

All parental hosts of heterospecific brood parasites must paythe cost of rearing non-kin. Previous research on nest parasitismby brown-headed cowbirds (Molothrus ater) concluded that competitivesuperiority of the typically more intensively begging and largercowbird chick leads to preferential feeding by foster parentsand causes a reduction in the hosts' own brood. The larger sizeof cowbird nestlings can be the result of at least two causes:(1) cowbirds preferentially parasitize species with smallernestlings and lower growth rates; and/or (2) cowbirds hatchearlier than hosts. I estimated the cost of cowbird parasitismfor each of 29 species by calculating the difference betweenhosts' published brood sizes in nonparasitized and parasitizednests and using clutch size to standardize values. In this analysis,greater incubation length and lower adult mass, surrogate measuresof the hatching asynchrony and size difference between parasiteand hosts, were both related to greater costs of cowbird parasitismwithout bias owing to phylogeny. To establish causality, I manipulatedclutch contents of eastern phoebes (Sayornis phoebe) and examinedwhether earlier hatching by a single cowbird or phoebe egg reducesthe size of the rest of the original host brood. As predicted,greater hatching asynchrony increased the proportion of theoriginal phoebe brood that was lost. This measure of the costof parasitism was partially owing to increased hatching failureof the original eggs in asynchronous broods but was not at allrelated to the size differences of older and younger conspecificnestmates. However, proportional brood loss owing to an earlierhatching conspecific was consistently smaller than brood lossowing to asynchronous cowbirds in both naturally and experimentallyparasitized phoebe nests. These results imply that althoughhatching asynchrony is an important cause of the reduction ofhost broods in parasitized clutches, competitive features ofcowbird nestlings remain necessary to explain the full extentof hosts' reproductive costs caused by interspecific brood parasitism.     

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.     

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.      

Social status affects the degree of sex difference in the songbird brain

It is thought that neural sex differences are functionally related to sex differences in the behaviour of vertebrates. A prominent example is the song control system of songbirds. Inter-specific comparisons have led to the hypothesis that sex differences in song nuclei size correlate with sex differences in song behaviour. However, only few species with similar song behaviour in both sexes have been investigated and not all data fit the hypothesis. We investigated the proposed structure-function relationship in a cooperatively breeding and duetting songbird, the white-browed sparrow weaver (Plocepasser mahali). This species lives in groups of 2-10 individuals, with a dominant breeding pair and male and female subordinates. While all male and female group members sing duet and chorus song, a male, once it has reached the dominant position in the group, sings an additional type of song that comprises a distinct and large syllable repertoire. Here we show for both types of male-female comparisons a male-biased sex difference in neuroanatomy of areas of the song production pathway (HVC and RA) that does not correlate with the observed polymorphism in song behaviour. In contrast, in situ hybridisation of mRNA of selected genes expressed in the song nucleus HVC reveals a gene expression pattern that is either similar between sexes in female-subordinate male comparisons or female-biased in female-dominant male comparisons. Thus, the polymorphic gene expression pattern would fit the sex- and status-related song behaviour. However, this implies that once a male has become dominant it produces the duetting song with a different neural phenotype than subordinate males.     

Egg coloration and selection for crypsis in open-nesting blackbirds

High variation in egg coloration among birds has traditionally been explained as adaptation for camouflage. We tested this hypothesis by conducting reciprocal clutch exchanges (n=301) among Brewer's blackbirds Euphagous cyanocephalus , red-winged blackbirds Agelaius phoeniceus , and yellow-headed blackbirds Xanthocephalus xanthocephalus . We predicted that clutches placed against their natural nest backgrounds would have higher survival rates than heterospecific clutches. Intraspecific clutch exchanges were used as a control. Clutch survival was monitored for a 9-d period at all nests, during which time incubation rhythm and nest defense were quantified. Intraspecific clutch exchanges did not influence incubation or nest defense. For two of the species, intraspecific clutch exchanges did not influence clutch survival; in red-winged blackbirds, however, intraspecifically exchanged clutches had somewhat depressed survival curves relative to control clutches (P=0.08). The effect of interspecific clutch exchanges differed by host species. In Brewer's nests, eggs of the yellow-headed blackbird had lower survival than Brewer's eggs (P=0.02), but survival of red-winged blackbird eggs did not differ from Brewer's eggs (P=0.50). In nests of red-winged blackbirds, all three clutch types had approximately equal survival. In yellow-headed blackbird nests, eggs of the red-winged blackbird had lower survival than yellow-headed blackbird eggs (P=0.06), and survival of Brewer's eggs did not differ from yellow-headed blackbird eggs (P=0.31). These findings support a role for egg coloration as camouflage in two of the three species studied.     

Rates of parasitism,but not allocation of egg resources,vary among and within hosts of a generalist avian brood parasite

Brown-headed cowbirds (Molothrus ater) deposit their eggs into the nests of other birds, which then raise the cowbird chick. Female cowbirds thus have limited options for impacting their offspring’s development via maternal effects compared to most other passerines. Cowbirds can impact their offspring’s phenotype by choosing among potential host nests, and by adjusting egg resources based on host characteristics. To examine whether cowbirds exhibit either or both of these strategies, we investigated rates of cowbird parasitism and egg investment (egg size, yolk-to-albumen ratio, and yolk testosterone and androstenedione) among and within host species in a shrubland bird community. We found that the probability of being parasitized by cowbirds, controlling for host status as a cowbird egg accepter or rejecter and ordinal date, varied significantly among host species, indicating an apparent preference for some hosts. Parasitism rates did not differ with host size, however, and despite variation in cowbird egg size among host species, this variation was not related to host size or cowbird preference. Among host species with eggs that are larger than those of the cowbird, cowbirds were significantly more likely to parasitize nests with relatively smaller eggs, whereas parasitism rates did not vary with relative egg size in host species with smaller eggs. There was no evidence for variation in cowbird egg components among or within host species. Our data indicate that cowbirds discriminate among host nests, but do not appear to adjust the composition of their eggs based on inter- or intraspecific host variation.     

Does the house sparrow Passer domesticus represent a global model species for egg rejection behavior?

Conspecific brood parasitism (CP) is a facultative breeding tactic whereby females lay their eggs in the nests of conspecifics. In some species, potential hosts have evolved the ability to identify and reject foreign eggs from their nest. Previous studies suggest that the ubiquitous house sparrow Passer domesticus in Spain and South Africa employs both CP and egg rejection, while a population in China does not. Given the species’ invasive range expansions, the house sparrow represents a potentially excellent global model system for parasitic egg rejection across variable ecological conditions. We examined the responses of house sparrows to experimental parasitism at three geographically distinct locations (in Israel, North America, and New Zealand) to provide a robust test of how general the findings of the previous studies are. In all three geographic regions egg rejection rates were negligible and not statistically different from background rates of disappearance of control eggs, suggesting that the house sparrow is not a suitable model species for egg rejection experiments on a global scale.     

Stable isotopes identify the natal origins of a generalist brood parasite, the brown-headed cowbird Molothrus ater

Identifying the natal origins of brood parasites is a major challenge that usually requires labor-intensive searching for nests of host species. Stable isotope analysis of feathers and other body tissues of parasitic young could be a possible tool for determining natal origins if tissues reflect the isotopic composition of the diet fed to nestlings. We measured the carbon (¹³C) and nitrogen (¹⁵N) isotope compositions of feathers for two age-classes of brown-headed cowbirds Molothrus ater at the Konza Prairie Biological Station near Manhattan, Kansas: nestlings raised by five species of songbird hosts in two different habitats, and juveniles captured after independence. Isotope values from cowbird nestlings did not differ among host species and we were unable to assign juvenile cowbirds to their natal hosts. However, nestlings raised in grassland habitat had feathers that contained significantly higher δ¹³C values and lower δ¹⁵N values than nestlings raised in shrub habitats. In addition, independent juveniles had isotopic signatures that were similar to cowbird nestlings raised on shrub habitats. Although dickcissel Spiza americana comprised the majority of samples from shrub habitats, our conclusions reflect the natural pattern of parasitism at the site and should be representative of cowbirds raised at Konza. We conclude that stable isotope analysis of feathers is effective for determining the natal origins of parasitic young if isotope values from nestlings are isotopically distinct among habitats.     

Does nesting habitat predict hatch synchrony between brood parasitic brown‐headed cowbirds Molothrus ater and two host species?

Nestling brown-headed cowbirds Molothrus ater typically hatch earlier and grow faster than young of the many host species of this generalist obligate brood parasite. However, a cowbird chick also benefits from the presence of some host nest mates as the parasite is provisioned disproportionately more with increasing brood size. Since asynchronous hatching affects both cowbird and host nestlings' growth and survival, mechanisms that optimize the timing of egg-laying by female parasites should be prevalent. Several habitat features might facilitate optimal timing of parasitic egg-laying and we examined whether aspects of host nesting habitat predicted cowbird hatching synchrony. We tested whether synchronous nests were less concealed, closer to perches, and located in areas of higher host density than asynchronous nests using a broad-scale information theoretic approach. There was no support for these predictions regarding song sparrow ( Melospiza melodia ; n=55) or yellow warbler ( Dendroica petechia ; n=67) nests parasitized by brown-headed cowbirds at Mono Lake, USA. For example, the best statistical models for predicting hatching synchrony in yellow warbler nests included nesting-patch width and nest-substrate shrub species. However, these relationships were relatively weak: both synchronous and asynchronous nests were in patches with statistically indistinguishable widths and the two dominant shrub species at our site contained similar proportions of synchronous and asynchronous nests. We conclude that the variability of host nesting habitats does not contribute to a biologically consistent effect on hatching synchrony by this generalist brood parasite.     

Differential reproductive success favours strong host preference in a highly specialized brood parasite

Obligate avian brood parasites show dramatic variation in the degree to which they are host specialists or host generalists. The screaming cowbird Molothrus rufoaxillaris is one of the most specialized brood parasites, using a single host, the bay-winged cowbird (Agelaioides badius) over most of its range. Coevolutionary theory predicts increasing host specificity the longer the parasite interacts with a particular avian community, as hosts evolve defences that the parasite cannot counteract. According to this view, host specificity can be maintained if screaming cowbirds avoid parasitizing potentially suitable hosts that have developed effective defences against parasitic females or eggs. Specialization may also be favoured, even in the absence of host defences, if the parasite's reproductive success in alternative hosts is lower than that in the main host. We experimentally tested these hypotheses using as alternative hosts two suitable but unparasitized species: house wrens (Troglodytes aedon) and chalk-browed mockingbirds (Mimus saturninus). We assessed host defences against parasitic females and eggs, and reproductive success of the parasite in current and alternative hosts. Alternative hosts did not discriminate against screaming cowbird females or eggs. Egg survival and hatching success were similarly high in current and alternative hosts, but the survival of parasitic chicks was significantly lower in alternative hosts. Our results indicate that screaming cowbirds have the potential to colonize novel hosts, but higher reproductive success in the current host may favour host fidelity.     

Peripheral auditory processing changes seasonally in Gambel’s white-crowned sparrow

Song in oscine birds is a learned behavior that plays important roles in breeding. Pronounced seasonal differences in song behavior and in the morphology and physiology of the neural circuit underlying song production are well documented in many songbird species. Androgenic and estrogenic hormones largely mediate these seasonal changes. Although much work has focused on the hormonal mechanisms underlying seasonal plasticity in songbird vocal production, relatively less work has investigated seasonal and hormonal effects on songbird auditory processing, particularly at a peripheral level. We addressed this issue in Gambel’s white-crowned sparrow (Zonotrichia leucophrys gambelii), a highly seasonal breeder. Photoperiod and hormone levels were manipulated in the laboratory to simulate natural breeding and non-breeding conditions. Peripheral auditory function was assessed by measuring the auditory brainstem response (ABR) and distortion product otoacoustic emissions (DPOAEs) of males and females in both conditions. Birds exposed to breeding-like conditions demonstrated elevated thresholds and prolonged peak latencies when compared with birds housed under non-breeding-like conditions. There were no changes in DPOAEs, however, which indicates that the seasonal differences in ABRs do not arise from changes in hair cell function. These results suggest that seasons and hormones impact auditory processing as well as vocal production in wild songbirds.     

Seasonal expression of androgen receptors, estrogen receptors, and aromatase in the canary brain in relation to circulating androgens and estrogens

Songbirds have a complex neural network for learning and production of song, namely the neural song system. Several nuclei of the song system contain androgen receptors (AR), and the neostriatal nucleus HVc also contains alpha type estrogen receptors (ER). Many songbird species show seasonal changes in both song and the neural song system that are correlated with seasonal variations in the circulating levels of gonadal steroids. However, there is increasing evidence that the sensitivity of the song system to gonadal steroids also changes seasonally. This could involve changes in the expression and activity of steroid receptors and steroid-metabolizing enzymes, such as the estrogen-synthesizing enzyme aromatase (AROM). The seasonal regulation of brain AR, ER, and AROM has not been studied before in the same individual songbirds. In this work, we compared plasma levels of androgens and estrogens, the expression level of AR-, ER-, and AROM-mRNA in the telencephalon, and brain AROM activity in male canaries between autumn (November) and spring (April) periods of high singing activity. Plasma levels of androgens and estrogens were higher in April than in November. The expression level of ER in HVc was higher in November than in April. In contrast, the expression level of AROM in the caudomedial neostriatum was higher in April than in November. However, we found no seasonal differences in the level of expression of AR and the volume of HVc as delimited by AR expression. Thus, AR expression in HVc was not correlated with circulating androgen levels. This study shows that both steroid-dependent and -independent seasonal factors regulate the action of gonadal hormones on the song system. In addition, we report a new site of AROM expression in the songbird brain, the nucleus interfacialis.