Avian malaria is associated with increased reproductive investment in the blue tit

Haemosporidians causing avian malaria are very common parasites among bird species. Their negative effects have been repeatedly reported in terms of deterioration in survival prospects or reproductive success. However, a positive association between blood parasites and avian fitness has also been reported. Here, we studied a relationship between presence of malaria parasites and reproductive performance of the host, a hole‐breeding passerine – the blue tit Cyanistes caeruleus. Since the malaria parasites might affect their hosts differently depending on environmental conditions, we performed brood size manipulation experiment to differentiate parental reproductive effort and study the potential interaction between infection status and brood rearing conditions on reproductive performance. We found individuals infected with malaria parasites to breed later in the season in comparison with uninfected birds, but no differences were detected in clutch size. Interestingly, infected parents produced heavier and larger offspring with stronger reaction to phytohemagglutinin. More importantly, we found a significant interaction between infection status and brood size manipulation in offspring tarsus length and reaction to phytohemagglutinin: presence of parasites had stronger positive effect among birds caring for experimentally enlarged broods. Our results might be interpreted either in the light of the parasite‐mediated selection or terminal investment hypothesis.     

Effect of an ectoparasite on lay date, nest-site choice, desertion, and hatching success in the great tit (Pants major)

Ectoparasites are common in most bird species, but experimentalevidence of their effects on life-history traits is scarce.We investigated experimentally the effects of the hematophagoushen flea (Ceratophyllus gallinae) on timing of reproduction,nest-site choice, nest desertion, clutch size, and hatchingsuccess in the great tit (Parus major). When great tits wereoffered a choice on their territory between an infested anda parasite-free nest-box, they chose the one without parasites.When there was no choice, the great tits in a territory containingan infested nest-box delayed laying the clutch by 11 days ascompared with the birds that were offered a parasite-free nestingopportunity. The finding that there was no difference in phenotypictraits related to dominance between the birds nesting in infestedboxes and birds nesting in parasite-free boxes suggests thatthe delay is not imposed by social dominance. Nest desertionbetween laying and shortly after hatching was significandy higherin infested nests. There was no difference between infestedand parasite-free nests in clutch size, but hatching successand hence brood size at hatching were significantly smallerin infested nests. Nest-box studies of great tits have beenseminal in the development of evolutionary, ecological, andbehavioral theory, but recently a polemic has arisen in theliterature about the validity of the conclusions drawn fromnest-box studies where the naturally occurring, detrimentalectoparasites are eliminated by the routine removal of old nestsbetween breeding seasons. Our study suggests that this criticismis valid and that the evaluation of the effects of ectoparasitesmay improve our understanding of behavioral traits, life-historytraits, or population dynamics     

Habitat-specific clutch size and cost of incubation in eiders reconsidered

The energetic incubation constraint hypothesis (EICH) for clutch size states that birds breeding in poor habitat may free up resources for future reproduction by laying a smaller clutch. The eider (Somateria mollissima) is considered a candidate for supporting this hypothesis. Clutch size is smaller in exposed nests, presumably because of faster heat loss and higher incubation cost, and, hence, smaller optimal clutch size. However, an alternative explanation is partial predation: the first egg(s) are left unattended and vulnerable to predation, which may disproportionately affect exposed nests, so clutch size may be underestimated. We experimentally investigated whether predation on first-laid eggs in eiders depends on nest cover. We then re-evaluated how nesting habitat affects clutch size and incubation costs based on long-term data, accounting for confounding effects between habitat and individual quality. We also experimentally assessed adult survival costs of nesting in sheltered nests. The risk of egg predation in experimental nests decreased with cover. Confounding between individual and habitat quality is unlikely, as clutch size was also smaller in open nests within individuals, and early and late breeders had similar nest cover characteristics. A trade-off between clutch and female safety may explain nest cover variation, as the risk of female capture by us, mimicking predation on adults, increased with nest cover. Nest habitat had no effect on female hatching weight or weight loss, while lower temperature during incubation had an unanticipated positive relationship with hatching weight. There were no indications of elevated costs of incubating larger clutches, while clutch size and colony size were positively correlated, a pattern not predicted by the ‘energetic incubation constraint’ hypothesis. Differential partial clutch predation thus offers the more parsimonious explanation for clutch size variation among habitats in eiders, highlighting the need for caution when analysing fecundity and associated life-history parameters when habitat-specific rates of clutch predation occur.     

Microsporidiosis in a Daphnia pulex population

The impact of parasitism by Thelohania sp., an intracellular microsporidian, on Daphnia pulex inhabiting a vernal pond was studied for three years. Three issues were considered: the distribution of parasites among hosts, the effect of parasites on individuals, and the impact of parasitism on population growth. Each year, Thelohania infected members of at least half of the generations of Daphnia produced in the pond. When the frequency of infection was low, parasites were found only in large adults. As infection frequency increased, parasitism spread downward through host size classes. However, parasites were rarely found in juveniles. Parasitism reduced clutch size drastically, increased mortality to a variable extent, had little impact on egg size or on per instar growth and none on molt frequency. Interaction with other stresses, such as food limitation, exacerbated some of these effects. Parasitism reduced instantaneous birth rate much more than it elevated instantaneous death rate. Population growth was reduced significantly but it is unlikely that Thelohania alone regulates the growth of this Daphnia population.     

Increased detrimental effects of ectoparasites on their bird hosts during adverse environmental conditions

Parasites are usually assumed to negatively affect fitness components of their hosts, particularly during adverse environmental conditions. We experimentally tested whether the ectoparasitic house martin bug (Oeciacus hirundinis) had larger negative effects on its house martin (Delichon urbica) host during second clutches. Conditions for reproduction decreased during the season, being less favourable for the second clutch than the first clutch. This was demonstrated by smaller clutch sizes, lower breeding success and smaller and lighter nestlings during the second clutch, as compared to the first, in the absence of parasites. Experimental manipulation of the intensity of parasitism resulted in the reproductive success of the host being reduced with increasing numbers of house martin bugs. This effect was much stronger during the second, as compared to the first, clutch. The house martin bug therefore had a greater detrimental effect on the reproduction of its host when conditions for reproduction were poor.     

Plasmodium relictum (lineage SGS1) and Plasmodium ashfordi (lineage GRW2): the effects of the co-infection on experimentally infected passerine birds

The effects of avian malaria parasites of the genus Plasmodium on their hosts are insufficiently understood. This is particularly true for malarial co-infections, which predominant in many bird populations. We investigated effects of primary co-infection of Plasmodium relictum (lineage SGS1) and Plasmodium ashfordi (GRW2) on experimentally infected naive juveniles of siskin Spinus spinus, crossbill Loxia curvirostra and starling Sturnus vulgaris. All siskins and crossbills were susceptible but starlings resistant to both these infections. A general pattern of the co-infections was that heavy parasitemia (over 35% during peaks) of both parasites developed in both susceptible host species. There were no significant effects of the co-infections on mean body mass of the majority of infected birds. Mean haematocrit value decreased approximately 1.5 and 3 times in siskins and crossbills at the peak of parasitemia, respectively. Mortality was recorded among infected crossbills. We conclude that co-infections of P. relictum and P. ashfordi are highly virulent and act synergetically during primary infections in some but not all passerine birds.     

Allocation of reproductive effort in female Arrenurus spp. water mites (Acari: Hydrachnidia; Arrenuridae)

Summary Twenty-seven species of water mites of the genus Arrenurus were compared with respect to allocation of reproductive effort. Clutch volume was positively correlated to female volume, female volume was positively correlated with clutch size and with larval volume, while clutch size and larval volume were negatively correlated. In threespace, corresponding to female volume, clutch size and larval volume, species were arranged along two trajectories representing separate reproductive strategies. The strategy characterized by small larvae, large clutches and large females corresponded to species of the subgenus Arrenurus, known to be larval parasites of adult Odonata. Members of three subgenera, known to be larval parasites of adult Diptera, followed the second strategy of small clutch size and alrge larvae. Of nine species of mites tested for intraspecific relationships, only one significant relationship was found, between female volume and clutch size for A. (Megaluracarus) bartonensis.     

Interactions between population processes in a cyclic species: parasites reduce autumn territorial behaviour of male red grouse

The causes of population cycles fascinate and perplex ecologist. Most work have focused on single processes, whether extrinsic or intrinsic, more rarely on how different processes might interact to cause or mould the unstable population dynamics. In red grouse (Lagopus lagopus scoticus), two causal mechanisms have been supported: territorial behaviour (changes in autumn aggressiveness) and parasites (parasite induced reduction in fecundity). Here, we report on how these two regulatory processes might interact, by testing whether the parasite suspected to cause the grouse cycles, the nematode Trichostrongylus tenuis, reduces male autumn territorial behaviour. We either treated males with an anthelmintic, to remove parasites (dosed or D-males), or challenged them with infective T. tenuis larvae, to increase parasite intensity (challenged or C-males). We first show that dosing was effective in removing T. tenuis parasites, while parasite intensities increased in challenged birds during the autumn. Because old males initially had more parasites than young males, the treatments generated greater differences in parasite intensity in old than in young males. We also show that various aspects of territorial behaviour (increase in testosterone-dependent comb size in autumn, territorial call rate, likelihood of winning territorial interactions and over-winter survival) were significantly higher in dosed than in challenged males, but in old birds only. Our data thus supported the hypothesis that parasites reduce male aggressiveness during the autumn territorial contests, and could thereby influence recruitment. Our results also highlight that the territorial behaviour of young males, which have fewer parasites, is not as limited by parasites as that of old, previously territorial males. We discuss the implications of these findings for our understanding of the processes regulating red grouse populations and causing their complex, unstable population dynamics.     

Consequences of chronic infections with three different avian malaria lineages on reproductive performance of Lesser Kestrels (Falco naumanni)

We studied the consequences of chronic infections by three different lineages of avian malaria, two Plasmodium (RTSR1, LK6) and one Haemoproteus (LK2), on reproductive performance of Lesser Kestrels (Falco naumanni). Malaria infections in male and female parents had no effect on clutch size, hatching success or nesting success. However, when only successful nests were considered, we found that males parasitized by LK6 raised a lower number of fledglings, suggesting that the level of parental effort by males may be limited by this particular lineage of Plasmodium. This effect was not evident in females, probably due to the higher investment of males during the chick rearing period in this species. Overall, we have found that chronic stages of specific malaria lineages have certain negative consequences on host reproductive performance, highlighting the importance of considering genetic differences among malaria parasites to study their consequences on natural bird populations.     

The phenology and clutch size of UK Blue Tits does not differ with woodland composition

The deciduous tree-herbivorous caterpillar-insectivorous bird food chain is a well-studied system for investigating the impacts of climate change across trophic levels. To date, across Europe, most attention has focused on the impacts of increasing spring temperature on changes to phenology in Oak-dominated (Quercus spp.) woodlands. Paridae species and Pied Flycatcher Ficedula hypoleuca are the most studied secondary consumers, all of which demonstrate an advancement in reproductive phenology with increases in spring temperature. Shifts in climate and phenology may also impact on reproductive investment in clutch size, and the effects of climate on phenology and clutch size may vary depending on woodland composition. To date, the effects of among-habitat variation in phenology and reproductive investment have received little attention. Insectivorous birds inhabiting woodlands that differ in tree composition may differ in the timing of breeding, due to local tree leafing phenology acting as a cue for egg-laying date and/or clutch size. Moreover, for most insectivorous birds, woodland composition within a territory is likely to be the main determinant of food availability for both adults and chicks. Consequently, if warming springs affect the temporal patterns of food availability differently across different woodland compositions, this may affect the optimal average local phenology for nesting birds. Here, using data from 34 long-term (mean 15 years) nest monitoring sites across the UK, we investigate the effect of woodland tree composition and temperature on Blue Tit Cyanistes caeruleus first egg date (FED) and clutch size. We supplemented the nest monitoring data by quantifying woodland composition, at a site level, through modified point counts. We predict that birds breeding in woodlands with greater proportions of late-leafing species, such as Oak and Ash Fraxinus excelsior, will breed later than those breeding in woodlands with greater proportions of early-leafing species, such as Birch Betula spp. and Beech Fagus sylvatica. We found no evidence for differences in Blue Tit FED or clutch size in relation to the proportion of any of the tree species investigated, after controlling for temperature and latitude (FED: −3.4 and 2.2, clutch size: −0.4 and − 0.2 eggs for one-unit increase in temperature and latitude, respectively). In recent decades and across all sites, clutch size has decreased as spring temperatures have increased, a strategy which could allow birds flexibly to adjust their breeding phenology such that nestling demand coincides with peak food availability. The lack of an effect of woodland composition on Blue Tit phenology suggests Blue Tits do not fine-tune their reproductive phenology to the local tree composition. Whether this lack of evidence for phenological divergence is due to an absence of divergent selection on breeding phenology and clutch size or to gene flow is not clear.     

Plasmodium relictum (lineage P-SGS1): effects on experimentally infected passerine birds

We evaluated the effects of Plasmodium relictum (lineage P-SGS1), which is a host generalist, to five species of passerine birds. Light infection of P. relictum was isolated from a naturally infected adult reed warbler Acrocephalus scirpaceus. The parasites were inoculated to naive juveniles of the chaffinch Fringilla coelebs, common crossbill Loxia curvirostra, house sparrow Passer domesticus, siskin Spinus spinus and starling Sturnus vulgaris. Susceptibility of these birds to the infection of P. relictum was markedly different. This parasite developed in birds belonging to the Fringillidae and Passeridae but the starlings (Sturnidae) were resistant. Only 50% of experimental house sparrows were susceptible to the infection. The intensity of parasitemia varied markedly inside and between different susceptible bird species. There were no effects of the infection on body mass or temperature of experimentally infected birds. Infection of P. relictum leads to the significant decrease of haematocrit value and hypertrophy of spleen and liver in heavily infected common crossbills and siskins. This study shows that infection of the same lineage of P. relictum causes diseases of different severity in different avian hosts; that might have different evolutionary consequences and should be taken in consideration in conservation projects.     

Elevational gradient in clutch size of Red‐faced Warblers

Our understanding of life history evolution has benefited from debates regarding the underlying causes, and geographic ubiquity, of spatial patterns in avian clutch sizes. Past studies have revealed that birds lay smaller clutch sizes at higher elevation. However, in most previous studies, investigators have failed to adequately control for elevational differences in breeding phenology. To better understand the elevational gradient in avian clutch size, we need to know how clutch size changes across the entire elevational breeding range of a species (i.e., the shape of the relationship between elevation and clutch size), and whether the elevational gradient in clutch size is merely an artifact of elevational gradients in breeding phenology or breeding season length. We examined the relationship between breeding elevation and clutch size of Red‐faced Warblers (Cardellina rubrifrons) along a 1000‐m elevational gradient in Arizona. Our objectives were to determine how clutch size changed with elevation, and if the relationship between clutch size and elevation merely reflected elevational changes in breeding season length or phenology. The proportion of 5‐egg clutches decreased and the proportion of 3‐ and 4‐egg clutches increased non‐linearly with increasing elevation, even after controlling for the elevational gradient in nest initiation date. Thus, average clutch size declined across the elevational breeding range of Red‐faced Warblers, but this decline was not due to elevational variation in breeding phenology. Timing of breeding changed, but the duration of the breeding season did not change appreciably across the elevational gradient. Hence, elevational differences in breeding season length or breeding phenology cannot explain why Red‐faced Warblers (and perhaps other birds) breeding at higher elevations have smaller clutches.     

Chronic malaria infections increase family inequalities and reduce parental fitness: experimental evidence from a wild bird population

Avian malaria parasites (Plasmodium) occur commonly in wild birds and are an increasingly popular model system for understanding host–parasite co‐evolution. However, whether these parasites have fitness consequences for hosts in endemic areas is much debated, particularly since wild‐caught individuals almost always harbour chronic infections of very low parasite density. We used the anti‐malarial drug Malarone^TM to test experimentally for fitness effects of chronic malaria infection in a wild population of breeding blue tits (Cyanistes caeruleus). Medication caused a pronounced reduction in Plasmodium infection intensity, usually resulting in complete clearance of these parasites from the blood, as revealed by quantitative PCR. Positive effects of medication on malaria‐infected birds were found at multiple stages during breeding, with medicated females showing higher hatching success, provisioning rates and fledging success compared to controls. Most strikingly, we found that treatment of maternal malaria infections strongly altered within‐family differences, with reduced inequality in hatching probability and fledging mass within broods reared by medicated females. These within‐brood effects appear to explain higher fledging success among medicated females and are consistent with a model of parental optimism in which smaller (marginal) offspring can be successfully raised to independence if additional resources become available during the breeding attempt. Overall, these results demonstrate that chronic avian malaria infections, far from being benign, can have significant effects on host fitness and may thus constitute an important selection pressure in wild bird populations.     

SHORT NOTES

Earlé, R. A. 1981. Factors governing avian breeding in Acacia savanna, Pietermaritzburg, Part 3: Breeding success, recruitment and clutch size. Ostrich 52:235-243.

The clutch size and breeding success of eight species of birds were monitored over a two year period in Acacia savanna. The mean clutch size fluctuated within the breeding season and four patterns of clutch size variation were noted. A smaller proportion of large clutches were laid in the 1979 season when less food was available. Breeding success was higher in the dry 1979 season but fewer breeding attempts were made and the overall recruitment was thus lower. At the peak of the breeding season, in mid-November, breeding success was at its lowest. Predation on nests was the greatest factor reducing breeding success although rain and strong winds affected some species.     

Geographic variation in malarial parasite lineages in the common yellowthroat (<Emphasis Type="Italic">Geothlypis trichas</Emphasis>)

Our current understanding of migration routes of many birds is limited and researchers have employed various methods to determine migratory patterns. Recently, parasites have been used to track migratory birds. The objective of this study was to determine whether haemosporidian parasite lineages detect significant geographic structure in common yellowthroats (Geothlypis trichas). We examined liver tissue or blood from 552 birds sampled from multiple locations throughout the continental United States, southern Canada, and the Bahamas. We found a 52.7% overall prevalence of haematozoan infection. We identified 86.1% of these infections to genus: 81% were Plasmodium; 5% were Haemoproteus; and 0.1% were Leucocytozoon. There were significant differences in the prevalence of different parasite genera among regions (χ² = 36.82, P < 0.0001) and in the proportion of Plasmodium infections versus other parasites among regions (χ² = 35.52, P < 0.0001). Sequence information identified three Haemoproteus lineages, two Leucocytozoon lineages, and thirteen Plasmodium lineages. Due to the low number of Haemoproteus and Leucocytozoon, only Plasmodium lineages were used in the geographic comparison of lineages. Six Plasmodium lineages were found in eight or more birds and the prevalence of these varied significantly among regions (χ² = 172.33, P < 0.0001). Additionally, 45 juvenile birds were sampled to determine what parasites could be obtained in the breeding grounds and we found only one lineage. In conclusion, parasite lineages show some geographic structure, with some lineages being more geographically specific than others, but are not useful for determining migratory connectivity in this species. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. The U.S. Government’s right to retain a non-exclusive, royalty-free license in and to any copyright is acknowledged.     

Breeding site fidelity in willow ptarmigan: the influence of previous reproductive success and familiarity with partner and territory

Summary Breeding site fidelity is high in willow ptarmigan: only 9% of males and 31% of females switched territories between years. Unpaired males were more likely to switch territories than paired males. For paired males, survival of their previous partner and reproductive success in year x did not influence probability of switching in year x+1. A female was more likely to switch territories if her previous partner disappeared. If her partner returned, she had a higher probability of switching if she did not produce chicks the previous year. Most hens moved to the territories of older males, although hens paired with unfamiliar older males did not have higher reproductive success than those paired with yearlings. Individuals that paired with their previous partner laid earlier and produced heavier chicks than those paired with unfamiliar partners. Excluding birds paired with familiar partners, survival and reproductive success in year x+1 was similar for males and females that did or did not switch territories. Males had a higher probability of producing chicks after switching than before, but females were more likely to lose their clutch after switching. For both sexes, birds that switched territories were as successful as the birds that replaced them on their former territories. We conclude that high site fidelity in willow ptarmigan is maintained because of the benefits of pairing with a familiar partner.     

The demands of incubation and avian clutch size

We reviewed information on the demands of incubation to examine whether these could influence the optimal clutch size of birds. The results indicate that appreciable metabolic costs of incubation commonly exist, and that the incubation of enlarged clutches can impose penalties on birds. In 23 studies on 19 species, incubation metabolic rate (IMR) was not elevated above the metabolic rate of resting non-incubating birds (RMR), but contrary to the physiological predictions of King and others, IMR was greater than RMR in 15 studies on 15 species. Across species, IMR was substantially above basal metabolic rate (BMR), averaging 1.606 × BMR. Of six studies on three species performed under thermo-neutral conditions, none found IMR to be in excess of RMR. IMRs measured exclusively within the thermo-neutral zone averaged only 1.08 × BMR contrasting with the significantly higher figure of 1.72 × BMR under wider conditions. 16 of 17 studies on procellariiforms found IMR below RMR, indicating a significant difference between this and other orders. We could find no other taxonomic, or ecological factors which had clear effects on IMR. Where clutch size was adjusted experimentally during incubation, larger clutches were associated with: significantly lower percentage hatching success in 11 of 19 studies; longer incubation periods in eight of ten studies; greater loss of adult body condition in two of five studies; and higher adult energy expenditure in eight of nine studies. Given that incubation does involve metabolic costs and given that the demands of incubation increase sufficiently with clutch size to affect breeding performance, we propose that the optimal clutch size of birds may in part by shaped by the number of eggs the parents can afford to incubate.     

Does follicle excision always result in enlargement of offspring size in lizards?

An experimental reduction of offspring number has been reported to result in enlargement of offspring size in lizards. We applied the “follicle excision” technique to a lacertid lizard (Takydromus septentrionalis) to examine whether this effect is generalisable to lizards. Of the 82 females that produced 3 successive clutches in the laboratory, 23 females underwent follicle excision after they oviposited the first clutch. Follicle excision reduced clutch size, but did not alter egg size. This result indicates that egg size is not altered during vitellogenesis in T. septentrionalis. Females undergoing follicle excision produced a third clutch (a second post-surgical clutch) as normally as did control females. Females switched from producing more but smaller eggs early in the breeding season to fewer but larger eggs later in the season. Our results indicate that female T. septentrionalis maximize reproductive success by diverting an optimal, rather than a higher, fraction of the available energy to individual offspring. This optimized allocation of the available energy to offspring production explains why follicle excision does not result in enlargement of egg size in this species. Our study provides evidence that an experimental reduction of offspring number does not always result in enlargement of offspring size in lizards.     

Schistosoma mansoni: Schistosomicidal effect of mefloquine and primaquine in vitro

We investigated the effects of the anti-malarials mefloquine and primaquine against the juvenile and adult life stages of Schistosoma mansoniin vitro. Cercariae were incubated with 0.5 μg/ml, 1 μg/ml and 2 μg/ml mefloquine or primaquine and with 1 μg/ml praziquantel for 12 h. Schistosomula, pre-adults and adults were incubated with 0.5 μg/ml, 1 μg/ml and 2 μg/ml mefloquine or primaquine and with 1 μg/ml praziquantel for 7 days. The viability status was classified as viable, damaged or dead and was checked every 3 h for cercariae and every 12 h for schistosomula, pre-adults and adults. Both, mefloquine and primaquine show time and dose-dependent schistosomicidal effects on the four life stages of S. mansoni. The promising in vitro effects on all stages of the blood fluke S. mansoni warrants further evaluation of both anti-malarials and their derivatives for their prophylactic and therapeutic values in early and late schistosomiasis in field trials.     

Yolk testosterone and estradiol variation relative to clutch size, laying order and hatching asynchrony in Common Grackles

Maternal effects are typically thought to enhance rather than reduce offspring performance, but asynchronous hatching (ASH) in birds typically produces a size hierarchy within a clutch that frequently reduces the growth and survival of nestlings from eggs that hatch later. Given that yolk steroids can significantly affect offspring phenotype and that in many species the levels of yolk steroids have been found to increase with laying order, the maternal transfer of steroids to egg yolk has been proposed as a mechanism for females to offset the deleterious effects of ASH. To test this hypothesis, we determined whether yolk steroids varied with laying order or clutch size in Common Grackles (Quiscalus quiscula). Because ASH varies with clutch size (hatching span averages 48 h in five-egg clutches, 24 h in four-egg clutches) and regularly results in the starvation of later hatched nestlings, we predicted: (1) testosterone and 17?-estradiol levels should increase with laying order in both clutch sizes to mitigate the negative effects of ASH on last-hatched nestlings, and (2) the increase should be greater in five-egg clutches due to more pronounced hatching asynchrony. Using a competitive-binding steroid radioimmunoassay, we found no systematic variation in either testosterone or estradiol levels relative to laying order or clutch size. In the absence of evidence that yolk steroids interact adaptively with ASH, research must look elsewhere for potential benefits that might compensate for the costs these steroids impose on nestlings.