The effect of nest temperature on growth and survival in juvenile Great Tits Parus major

1. For birds, maintaining an optimal nest temperature is critical for early‐life growth and development. Temperatures deviating from this optimum can affect nestling growth and fledging success with potential consequences on survival and lifetime reproductive success. It is therefore particularly important to understand these effects in relation to projected temperature changes associated with climate change.
2. Targets set by the 2015 Paris Agreement aim to limit temperature increases to 2°C, and, with this in mind, we carried out an experiment in 2017 and 2018 where we applied a treatment that increased Great Tit Parus major nest temperature by approximately this magnitude (achieving an increase of 1.6°C, relative to the control) during the period from hatching to fledging to estimate how small temperature differences might affect nestling body size and weight at fledging and fledging success.
3. We recorded hatching and fledging success and measured skeletal size (tarsus length) and body mass at days 5, 7, 10, and 15 posthatch in nestlings from two groups of nest boxes: control and heated (+1.6°C).
4. Our results show that nestlings in heated nest boxes were 1.6% smaller in skeletal size at fledging than those in the cooler control nests, indicating lower growth rates in heated boxes, and that their weight was, in addition, 3.3% lower.
5. These results suggest that even fairly small changes in temperature can influence phenotype and postfledging survival in cavity‐nesting birds. This has the potential to affect the population dynamics of these birds in the face of ongoing climatic change, as individuals of reduced size in colder winters may suffer from decreased fitness.

     

Experimental manipulation of temperature reduce ectoparasites in nests of blue tits Cyanistes caeruleus

Several models predict changes in the distributions and incidences of diseases associated with climate change. However, studies that investigate how microclimatic changes may affect host–parasite relationships are scarce. Here, we experimentally increased the temperature in blue tit Cyanistes caeruleus nest boxes during their breeding season to determine its effects on the parasitic abundance (i.e. of nest‐dwelling ectoparasites, blood‐sucking flying insects and hemoparasites) in nests and the host condition of nestlings and adults. The temperature was increased using heat mats placed underneath the nest material, which resulted in an average temperature increase of 3ºC and a reduction in relative humidity of about six units. The abundance of mites Dermanyssus gallinoides and blowfly pupae Protocalliphora azurea was significantly reduced in heated nest boxes. Although not statistically significant, a lower prevalence of flea larvae Ceratophyllus gallinae was also found in heated nests. However, heat treatment did not affect hemoparasite infection of adult blue tits or the body condition of adult and nestling blue tits. In conclusion, heat treatment in blue tit nests reduced nest‐dwelling ectoparasites yet without any apparent benefit for the host.     

Avoidance,tolerance, and resistance to ectoparasites in nestling and adult tree swallows

We examined avoidance, tolerance, and resistance strategies of nestling and adult tree swallows Tachycineta bicolor in response to ectoparasitic blowflies Protocalliphora sialia. Tree swallows avoided settling in north‐facing nest boxes early in the breeding season. These boxes were more likely to be parasitized later in the season, suggesting that box selection may facilitate blowfly avoidance. After experimentally manipulating blowfly intensity, we found that nestlings were generally tolerant of parasitism. Parasites significantly reduced nestling blood hemoglobin but had no effect on nestling body mass, primary feather growth, age at fledging, or fledging success. Parents of parasitized nestlings did not increase their food provisioning rate to promote nestling tolerance. Adult female tree swallows demonstrated both tolerance and resistance: blowfly parasitism had no effect on adult hemoglobin and body mass, and those with higher P. sialia‐binding antibody levels had fewer blowfly larvae in their nests. Nestling antibodies were unrelated to blowfly intensity. Despite considerable variation among years, our results suggest that the costs of blowfly parasitism to nestling and adult tree swallows are modest, and limited to blood loss in nestlings. Future work should examine the effects of reduced blood hemoglobin on fledgling survival and the importance of parasite‐specific antibodies.     

Behavioural responses to ectoparasites in pied flycatchers Ficedula hypoleuca: an experimental study

Nests of cavity‐nesting birds usually harbor some species of haematophagous ectoparasites that feed on the incubating adults and nestlings. Given the negative impact of ectoparasites on nestlings there will be selection on hosts to reduce parasite infestations through behavioural means. We have experimentally reduced the abundance of all ectoparasites in nests of pied flycatchers Ficedula hypoleuca to explore both whether there are changes in the frequency and duration of putative anti‐parasite behaviours by tending adults, as well as whether such anti‐parasite behaviours are able to compensate for the deleterious effects that parasites may have on nestlings. Heat treatment of nests substantially decreased the density of ectoparasites, and thereby positively affected nestling growth. The frequency and intensity of female grooming and nest sanitation behaviours during the incubation and nestling periods decreased as a consequence of the experimental reduction of ectoparasite infestation. Although nestlings begged more intensely in infested nests, the experiment had no significant effect on parental provisioning effort. Reduction of parasites resulted in larger nestlings shortly before fledging and increased fledging success. This study shows a clear effect of a complete natural nest ectoparasite fauna on parental behaviour at the nest and nestling growth in a cavity‐nesting bird. Although ectoparasites induce anti‐parasite behaviours in females, these behaviours are not able to fully remove parasite's deleterious effects on nestling growth and survival.     

The importance of microclimate variation in determining size,growth and survival of avian offspring: experimental evidence from a cavity nesting passerine

Organisms are expected to balance energy allocation in such a way that fitness is maximized. While much research has focussed on allocation strategies of reproducing parents, in particular birds, relatively little attention has been paid to how nestlings allocate energy while in the nest. Nestling birds are faced with a trade-off between devoting energy to growth or to thermoregulation, and in altricial species it is likely that the thermal environment of the nest site influences the nature of this trade-off. Here, we experimentally investigate how altering the microclimate of nests affects the growth, size and survival, as well as cell-mediated immune (CMI) response, of nestling tree swallows (Tachycineta bicolor) in a temperate environment. We place air-activated heating pads in nests of swallows when young were between 4 days and 16 days of age, and compared performance of offspring to control nests. Our manipulation raised temperatures of heated nests by approximately 5°C compared to control nests. Offspring in heated nests had enhanced survival while in the nest, and we also found that they were heavier and had longer ninth primary feathers at 16 days of age. In addition, heating nest boxes resulted in significantly faster growth of primaries, and there was a trend for growth rates of mass to also be higher in heated nests. There were no significant differences between heated and control nests in growth rate or size of tarsus at age 16 days, and we speculate that this lack of response to elevated nest temperatures may be due to growth of skeletal structures being limited by other factors such as calcium availability. We also found no difference between heated and control nests in CMI response. Nonetheless, our results show overall that increasing temperatures of nests has significant benefits that enhance the fitness of offspring. As provisioning rates to offspring did not differ between heated and control nests, we suspect that the beneficial effects of heating were not the consequence of changes in parental behaviour. Our results provide insight into factors, other than food supply, that have important consequences in determining reproductive success of birds breeding in temperate environments.     

Incubation temperature impacts nestling growth and survival in an open‐cup nesting passerine

For oviparous species such as birds, conditions experienced while in the egg can have long‐lasting effects on the individual. The impact of subtle changes in incubation temperature on nestling development, however, remains poorly understood, especially for open‐cup nesting species with altricial young. To investigate how incubation temperature affects nestling development and survival in such species, we artificially incubated American robin (Turdus migratorius) eggs at 36.1°C (“Low” treatment) and 37.8°C (“High” treatment). Chicks were fostered to same‐age nests upon hatching, and we measured mass, tarsus, and wing length of experimental nestlings and one randomly selected, naturally incubated (“Natural”), foster nest‐mate on days 7 and 10 posthatch. We found significant effects of incubation temperature on incubation duration, growth, and survival, in which experimentally incubated nestlings had shorter incubation periods (10.22, 11.50, and 11.95 days for High, Low, and Natural eggs, respectively), and nestlings from the Low treatment were smaller and had reduced survival compared to High and Natural nestlings. These results highlight the importance of incubation conditions during embryonic development for incubation duration, somatic development, and survival. Moreover, these findings indicate that differences in incubation temperature within the natural range of variation can have important carryover effects on growth and survival in species with altricial young.     

Effects of temperature and nest heat exposure on nestling growth,dehydration and survival in a Mediterranean hole‐nesting passerine

Variable environments impose constraints on adaptation by modifying selection gradients unpredictably. Optimal bird development requires an adequate thermal range, outside which temperatures can alter nestling physiology, condition and survival. We studied the effect of temperature and nest heat exposure on the reproductive success of a population of double‐brooded Spotless Starlings Sturnus unicolor breeding in a nestbox colony in central Spain with a marked intra‐seasonal variation in temperature. We assessed whether the effect of temperature differed between first and second broods, thus constraining optimal nest‐site choice. Ambient temperature changed greatly during the chick‐rearing period and had a strong influence on nestling mass and all body size measures we recorded, although patterns of clutch size or nestling mortality were not influenced. This effect differed between first and second broods: nestlings were found to have longer wings and bills with increasing temperature in first broods, whereas the effect was the opposite in second broods. Ambient temperature was not related to nestling body mass or tarsus‐length in first broods, but in second broods, nestlings were lighter and had smaller tarsi with higher ambient temperatures. The exposure of nestboxes to heat influenced nestling morphology: heat exposure index was negatively related to nestling body mass and wing‐length in second broods, but not in first broods. Furthermore, there was a positive relationship between nest heat exposure and nestling dehydration. Our results suggest that optimal nest choice is constrained by varying environmental conditions in birds breeding over prolonged periods, and that there should be selection for parents to switch from sun‐exposed to sun‐protected nest‐sites as the season progresses. However, nest‐site availability and competition for sites are likely to impose constraints on this choice.     

Substrate and structure of ground nests have fitness consequences for an alpine songbird

Songbird nests are an important life‐history component with multiple functions, including the creation of a suitable microclimate for offspring development. Thus, functional nest characteristics may influence fitness correlates, such as nestling size traits, and may co‐vary with prevailing environmental conditions. We investigated among‐ and within‐female variation in nest substrate, lining and decoration structures with associated fitness consequences (hatching success, nestling size traits, nest survival) across two breeding seasons for an alpine population of Horned Lark Eremophila alpestris. We combined these observations with explicit measures of nest temperature to address the influence of nest characteristics on microclimate. Nests in heather substrate had the coldest microclimates compared with grass and bare‐ground substrate, but also the greatest nest survival rates (68% versus 37–44% in other substrates), indicating the potential for substrate use decisions to reflect a trade‐off between microclimate and nest survival in response to prevailing weather and predation risk conditions. Furthermore, nest lining and nest decoration patterns indicated some support for a thermoregulatory function. Nests that were lined with willow (Salix sp.) seed‐down were associated with larger, heavier nestlings and the use of down lining decreased in frequency as the season warmed up. Nest decoration placed in front of the nest (e.g. stones or dirt clumps varying in mass from 5.3 to 186.6 g) was positively associated with warmer nest microclimates. Females demonstrated high phenotypic flexibility, as 61–94% of the observed variance in nest characteristics was explained by within‐female rather than among‐female differences. Such flexible nesting behaviour suggests the capacity to adjust to changing environmental conditions to maintain vital fitness correlates such as nest survival and nestling size development.     

Experimental food supplementation affects the physical development,behaviour and survival of Little Owl Athene noctua nestlings

In birds, energy supply during growth is a major predictor of the fledglings' physical condition and survival prospects. Differential quantity and quality of fledglings produced under varying nestling food supplies are likely to affect the number of offspring that recruit into the breeding population. However, the underlying mechanisms and associated consequences are still poorly known. Using a partial cross‐fostering and food supplementation experiment, we estimated the effect of variation in food supply during growth on nestling survival and fledgling phenotypic traits of Little Owls Athene noctua. Survival to fledging was much higher in food‐supplemented nestlings (98.6%) than in control nestlings (82.4%). Furthermore, supplemented nestlings were on average 8.9 g heavier and were more likely to develop subcutaneous fat deposits (99.4 vs. 73.7% of treatment and control nestlings, respectively). Supplemented nestlings also had on average longer wings than control nestlings, but tarsi and culmen did not differ significantly. Furthermore, experimentally supplemented fledglings struggled more when handled and emerged sooner from tonic immobility than control fledglings. The irises of supplemented fledglings were less intensely coloured. The experimentally induced changes in nestling development probably affect individual performance beyond fledging. Nestlings from orchard‐dominated habitats were larger than those from habitats dominated by arable land. As nestling food supply is largely determined by natural food availability, we conclude that habitat quality affects Little Owl productivity and offspring quality, and ultimately, population dynamics.     

Philornis infection in blue‐black grassquits: impact on nestlings and risk factors involved

Parasitic botfly larvae (Philornis ssp., Diptera: Muscidae) are found in nests of several bird taxa, although prevalence and impact on nestling survival vary considerably among species. Here we describe patterns of botfly infestation in blue‐black grassquit Volatinia jacarina nestlings. We identified the most typically affected nestling body parts and assessed parasite prevalence, impact on nestling survival, and changes in nestling body shape. Additionally, we tested whether climatic conditions, nest morphology and habitat characteristics are associated with larvae abundance. Blue‐black grassquits had low breeding success (16% of eggs/nestlings survived to fledged; 19% of the nests fledged at least one), but most failures resulted from predation by vertebrate predators. We estimated that 1% of nestlings died due to botfly infestation, and the number of subcutaneous larvae (range 1–18) in a nestling's body did not predict fledging success. Infected chicks exhibited higher tarsus asymmetry. Thus, we argue that although botflies had a small impact on offspring survival, they may reduce fitness in adulthood. There was no evidence that environmental conditions and nest morphology are linked to the number of larvae on nestlings. Nesting areas with higher food supply had lower infestation rates. Possibly, food‐rich habitats allow parents to invest more time in offspring care (brooding nestlings), thus protecting them from fly attacks. Alternatively, vegetation composition could influence local invertebrate diversity, which could provide a natural trophic buffer against adult Philornis. The present study brings to light new perspectives concerning bird–botfly interaction.     

How does an increase in minimum daily temperatures during incubation influence reproduction in the great tit Parus major?

Temperature variation affects all life stages of organisms, especially early development, and considering global warming, it is urgent to understand precisely its consequences. In egg‐laying species, incubation behaviour can buffer embryo developmental temperature variation and influence offspring development. We experimentally investigated the effect of an increase in minimum daily nest temperature during incubation in the great tit Parus major, by placing a hand warming pad under the nest in the evenings. As compared to controls, the experimental treatment increased nest temperature at night by an average of 4°C, and this increase carried over to the following day. We measured the consequences of this mainly nocturnal temperature increase during incubation on 1) parental behaviour (incubation and nestling feeding), 2) parental health (quantified by body condition, immune status, physiological and oxidative stress) and 3) reproductive success (nestling body condition, growth, i.e. mass gain, hatching and fledging success, and nestling immune status, physiological and oxidative stress). This study yielded three major results. First, we found that heating the nest did not change the duration of incubation as compared to controls. Second, increasing nest temperature during incubation decreased nestling feeding behaviour but did not affect parental health in terms of body condition, immune status, physiological and oxidative stress. Third, nestling mass at hatching was greater but nestling mass gain was slower in heated nests than in control nests, resulting in similar fledging mass. The present study demonstrates that increased environmental temperatures during incubation influenced nestling development in the great tit and especially hatchling mass, which might produce long‐term life history consequences.     

Comparison of digestive efficiency in the parasitic great spotted cuckoo and its magpie host nestlings

Altricial nestlings are under strong selection pressures to optimize digestive efficiency because this is one of the main factors affecting nestling growth and survival. Bird species vary in their ability to assimilate different nutrients and current theory predicts that nestlings should also be able to adjust their nutritional physiology to feeding frequency. Variation in parental provisioning to nestlings would select for flexibility in nestling digestive physiology, which would allow maximization of nutrient assimilation. In the present study, by making use of a brood parasite–host study system in which great spotted cuckoo nestlings (Clamator glandarius) are reared by magpie (Pica pica) host foster parents when sharing the nest with host nestlings, we tested several predictions of the adaptive digestive efficiency paradigm. A hand‐feeding experiment was employed in which we fed both great spotted cuckoo and magpie nestlings with exactly the same diet simulating one food abundance period and one food deprivation period. The results obtained show that cuckoo nestlings ingested more food, gained significantly more weight during the abundance period, and assimilated a higher proportion of the ingested food than magpie nestlings. These results demonstrate for the first time that cuckoo nestlings enjoy digestive adaptations that favour a rapid processing of the ingested food, thereby maximizing their intake rate but without decreasing digestive efficiency. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 280–289.     

Age‐related prenatal maternal effects and postnatal breeding experience have different influences on nestling development in an altricial passerine

Reproductive success and nestling performance are related to the age of parents across several vertebrate taxa. However, because breeding experience and prenatal maternal investment in reproduction often covary, the source of these age‐related differences can be difficult to determine. In this study, we evaluated the influence of prenatal maternal effects and postnatal breeding experience on the performance of nestling tree swallows Tachycineta bicolor by conducting a carefully controlled partial cross‐fostering experiment. We swapped half‐broods of nestlings between the nest of a young first‐time breeding female and the nest of a female known to have previously raised and fledged young. Our manipulation did not influence the within‐brood nestling hierarchies, and controlled for the effects of egg laying order. We found that nestlings of older females were heavier just prior to fledging regardless of the breeding experience of the attending female. In addition, fledglings raised by experienced females grew their flight feathers faster, and had greater probability of fledging. Our study demonstrates that prenatal investment in reproduction by older females can have long‐term consequences on nestling mass, and suggests limited potential for compensatory mass gains prior to fledging. Because our analyses controlled for feeding rates, our results also suggest that foraging quantity and quality are not the only benefits nestlings gain by being raised by an experienced female.     

Ectoparasitism and the role of green nesting material in the European starling

Summary The use of green nesting material is widespred among birds. Recent evidence suggests that birds use secondary chemicals contained in green plants to control ectoparasites. We manipulated green nesting material and ectoparasites of European starlings (Sturnus vulgaris) to test two hypotheses: (1) ectoparasites adversely affect prefledging survival and morphometrics or postfledging survival, and (2) green nesting material ameliorates the effects of ectoparasites. We recorded fat score, numbers of scabs, tarsal length, body mass, and hematocrit level on each nestling 17 days after hatching. We also fitted each nestling with unique patagial tags and resighted the starlings for 6–8 weeks after fledging to estimate survival and sighting rates. Nests devoid of green nesting material and dusted with the insecticide, carbaryl, had fewer high ectoparasite infestations, and nestlings had significantly lower scab scores, and significantly higher body masses than nestlings in undusted boxes. However, there was no difference in postfledging survival between birds from carbaryl-treated and undusted nests. There also was no difference in prefledging survival and morphometrics or postfledging survival between nestlings from boxes with and without green nesting material. These results do not support the hypothesis that starlings use green nesting material to control nest ectoparasites. We suggest an alternative hypothesis; green nesting material is used for mate selection or pairbonding in the starling.     

Discrepancy between factors affecting nestling growth and survival and maternal success in Common Grackles

ABSTRACT Multiple factors potentially affect nestling survival and maternal reproductive success. However, little is known about the relative importance of different factors when operating simultaneously or whether the same factors are important for nestlings and their mothers. We determined the effect of hatching asynchrony, individual egg size, mean egg size, nestling sex, and clutch initiation date on the survival of individual nestlings and on maternal reproductive success in Common Grackles (Quiscalus quiscula) from 2004 to 2006 in central Illinois. Factors most important to maternal success differed from those important for individual nestling growth and survival. Hatching asynchrony had the greatest within‐nest influence on the fate of nestlings; the earlier a nestling hatched relative to siblings, the greater its mass and likelihood of fledging. Clutch size had the greatest influence on maternal reproductive success, with females with larger clutches fledging more young. Thus, both nestling survival and maternal success were largely determined by a single, albeit different, factor. A possible explanation for the apparent unimportance of most factors we measured in determining maternal success is that we did not consider variation among females. Individual variation in maternal attributes such as condition, size, age, experience, or mate quality may result in females tailoring clutch attributes (i.e., egg size, sex, and degree of hatching asynchrony) in ways that allow them to maximize their reproductive success. The discordance between factors that benefited mothers versus their offspring illustrates the importance of considering the maternal consequences of any factor that appears to affect offspring survival. Factors that increase the mass and survival of some offspring may not result in increased maternal reproductive success.     

Fitness-related consequences of egg mass in nestling house wrens

The fitness-related consequences of egg mass, independent of confounding influences associated with parental quality, remain poorly understood for wild birds in general and for passerines in particular. We performed cross-fostering experiments to test the hypothesis that egg mass, independent of parental quality, is the primary determinant of fitness-related traits in nestling house wrens (Troglodytes aedon), an insectivorous passerine. Nestling mass was significantly correlated with the mass of the eggs from which nestlings hatched early but not late in the nestling period in early-season broods. In contrast, in late-season broods, nestling mass was correlated with egg mass until nestlings achieved asymptotic mass. Neither nestling growth nor survival to nest leaving was related to egg mass in either early- or late-season broods; however, nestlings in late-season broods grew more slowly than did nestlings in early-season broods. We propose that nestling mass and egg mass remained correlated throughout the nestling period in late-season broods because decreased arthropod food resources late in the breeding season constrain parents'' ability to provision nestlings. We conclude that female house wrens in this population trade-off clutch size for greater egg mass to maximise reproductive success in late-season broods.     

The effects of force‐fledging and premature fledging on the survival of nestling songbirds

Despite the broad consensus that force‐fledging of nestling songbirds lowers their probability of survival and therefore should be generally avoided by researchers, that presumption has not been tested. We used radiotelemetry to monitor the survival of fledglings of Ovenbirds Seiurus aurocapilla and Golden‐winged Warblers Vermivora chrysoptera that we unintentionally force‐fledged (i.e. nestlings left the nest in response to our research activities at typical fledging age), that fledged prematurely (i.e. nestlings left the nest earlier than typical fledging age), and that fledged independently of our activities. Force‐fledged Ovenbirds experienced significantly higher survival than those that fledged independent of our activities, and prematurely fledged Ovenbirds had a similarly high survival to those that force‐fledged at typical fledging age. We observed a similar, though not statistically significant, pattern in Golden‐winged Warbler fledgling survival. Our results suggest that investigator‐induced force‐fledging of nestlings, even when deemed premature, does not necessarily result in reduced fledgling survival in these species. Instead, our results suggest that a propensity or ability to fledge in response to disturbance may be a predictor of a higher probability of fledgling survival.     

Cross‐fostering reveals sources of variation in innate immunity and hematocrit in nestling tree swallows Tachycineta bicolor

Innate immunity plays an important role in protecting birds early in development. The strength of innate immunity in nestlings appears to be affected by heritable and early maternal effects, as well as later environmental effects. To explore factors affecting innate immunity, we conducted a partial cross‐fostering experiment in nestling tree swallows Tachycineta bicolor. The ability of nestling blood plasma to kill Escherichia coli in vitro was influenced by heritable or early maternal effects and was unrelated to body condition of nestlings or brood size of the foster nest. The relative importance of heritable and early maternal effects versus environmental effects on nestling hematocrit (packed volume of red blood cells) and body condition were also examined. Hematocrit was significantly influenced by the nest in which nestlings were reared, though it was not related to nestling body condition. Body condition was significantly influenced by the nest in which nestlings were reared, though it was also weakly influenced by heritable or early maternal effects. In summary, heritable and early maternal effects are stronger predictors than rearing conditions of innate immunity in nestling tree swallows. Furthermore, hematocrit is largely environmentally determined but is not related to body condition or innate immunity, as measured in this study.     

European starlings: nestling condition, parasites and green nest material during the breeding season

Male European starlings (Sturnus vulgaris) intermingle fresh herbs, preferably species rich in volatile compounds, into their dry nest material. In a field study, we investigated whether these herbs affect the mite and bacteria load of the nests and the condition of the nestlings either directly or via parasite control. We examined the amount of herbs and the number of plant species males carried into their nests, the variation of volatile compounds in the headspace air of the nest boxes and mite/bacteria load of the nests throughout the season. The amount of herb material and the number of plant species, the number of substances emanated by these plants and the infestation of the nests with bacteria and mites (Dermanyssus gallinae) increased with season. In a field experiment, we exchanged natural starling nests with experimental nests with or without herbs. We found that the herbs had no effect on the mites but fewer bacteria were sampled in nests with herbs than in nests without herbs. The body mass of the fledging was not related to the season or the mite/bacteria load of the nests. However, nestlings from nests with herbs fledged with higher body mass than nestlings from nests without herbs. Both bacteria and mite load were related to nestling mortality. In nests containing no herbs, the numbers of fledglings declined significantly with the increasing mite load while the mites had no effect on the number of fledglings in nests with herbs. Thus, the nest herbs counteracted the effect of the mites. In conclusion, it seems that volatile herbs can reduce bacterial but not mite infestation of the starling nests. The positive influence of herbs on nestling growth indicates that herbs either directly (perhaps as immunostimulants) improve the condition of the nestlings and help them cope with the harmful effects of mites, or they provide a nest environment beneficial for the nestlings‘ development by the reduction of germs.