The mucous covering of fecal sacs prevents birds from infection with enteric bacteria

Nestlings of many bird species produce fecal sacs, excrements encapsulated within a mucous covering. Although it facilitates parents' removal of feces from nests, which would improve hygienic conditions for developing nestlings, no functional (i.e. adaptive) explanation of fecal sac production has been previously investigated. We propose that the mucous covering would isolate enteric pathogenic bacteria, thereby preventing contamination of nestlings and parents. This antimicrobial hypothesis therefore predicts that density of bacteria would be drastically reduced from the inside to the outside of nestlings' droppings, and that the fecal sac covering would inhibit other bacterial grow. We tested these predictions by means of culturing bacteria obtained from different parts of the sac and inhibition tests. In accordance with the hypothesis, bacterial loads of the outside of fecal sacs were significantly lower than those estimated from the inside of the covering. In addition, we did not find evidence of antimicrobial activity of the covering, which suggests that the hypothesized bacterial isolation function is accomplished by a physical rather than a chemical protection. Bacterial density of the liquid that permeates out after 23 min does not differ with that estimated for the inside of the sac, suggesting short‐term effects of fecal sacs as bacterial barrier. These findings highlight the major role of bacterial infections as a selective pressure for explaining the evolution of traits that, as the covering of fecal sacs, facilitate nest sanitation in this group of animals.     

Green plants in starling nests: effects on nestlings

European starlings, Sturnus vulgaris, intermingle fresh herbs, especially species rich in volatile compounds, with their otherwise dry nest material. In this field study we investigated whether these herbs reduce ectoparasites and thereby protect nestlings (the nest protection hypothesis). We also considered whether volatile compounds in herbs improve the condition of nestlings (the drug hypothesis). As measures of condition we used body mass, haematocrit levels and immunological parameters. We replaced 148 natural starling nests with artificial ones: half contained herbs and half (controls) contained grass. The ectoparasite loads (mites, lice, fleas) in herb and control nests were indistinguishable. However, nestlings in herb nests weighed more and had higher haematocrit levels at fledging than nestlings in control nests. Fledging success was similar in herb and control nests, but more yearlings from herb nests were identified in the colony the year after hatching. The response of the immune system when challenged with phytohaemagglutinin did not differ in nestlings from herb and control nests. Nestlings from herb nests had more basophils and fewer lymphocytes in their blood than those from control nests, while the eosinophil and heterophil counts did not differ. We conclude that herbs do not reduce the number of ectoparasites, but they improve the condition of nestlings, perhaps by stimulating elements of the immune system that help them to cope better with the harmful activities of ectoparasites. Copyright 2000 The Association for the Study of Animal Behaviour.     

Experimental evidence for edge-related predation in a fragmented agricultural landscape

Abstract This study tested the hypothesis that increased predation of experimental nests occurs close to a forest edge in a fragmented agricultural landscape. Artificial nests and eggs of willie wagtails Rhipidura leucophrys and superb fairy-wrens Malurus cyaneus were used in experiments to assess the extent and nature of predation occurring throughout the known breeding seasons of these species. Predators were identified by the imprints they left in plasticine eggs, and by remote photography. Surveys of avian predators were undertaken to investigate the relationship between predation intensity and predator distribution and abundance. Avian predators accounted for almost all predation for which a predator could be identified (96%). Five of seven predator species photographed attacking wagtail nests were corvids or artamids. Fairy-wren nests suffered relatively low rates of predation (29%) compared to wagtail nests (87%). Increased predation at the habitat edge was recorded for wagtail nests only; predation was correlated with the distribution and abundance of predatory avian species. The different extent and pattern of predation on fairy-wren nests could be explained by problems in detecting predation by mammals, and by possible failure of avian predators to locate the cryptic nests.     

Sheep in wolf's clothing: host nestling vocalizations resemble their cowbird competitor's

Nestlings of many avian brood parasites are virtuosos at mimicking host nestling vocalizations, which, like egg mimicry, presumably ensures acceptance by host parents. Having been accepted, parasitic nestlings then often exaggerate the aspects of the host's display to increase parental care. Host nestlings may, in turn, exaggerate their vocalizations to keep up with the parasite, though this possibility has not been evaluated. We experimentally parasitized song sparrow (Melospiza melodia) nests with a brown-headed cowbird (Molothrus ater) chick to evaluate how host nestlings respond. Vocalizations emitted from experimentally parasitized nests were higher in frequency, and louder, than those from unparasitized nests, consistent with the cowbird exaggerating its signalling. In response, host nestlings exaggerated the frequency and amplitude of their vocalizations, such that they resembled the cowbird's while they 'scaled back' on calls per parental provisioning bout. Sparrows in parasitized nests were fed equally often as sparrows in unparasitized nests, suggesting that exaggerating some aspects of vocalization while scaling back on others can help host nestlings confronted with a cowbird. Our results support the recently proposed hypothesis that signalling in parasitized nests involves a dynamic interaction between parasitic and host nestlings, rather than a one-way process of mimicry by the parasite.     

Nest‐dwelling ectoparasites reduce begging effort in Pied Flycatcher Ficedula hypoleuca nestlings

Parasitized nestlings might be expected to increase begging effort to obtain additional resources to compensate for those sequestered by their parasites. However, begging is costly and chicks harbouring parasites may find it more difficult to attain high begging levels. Consequently, we predicted that, for the same level of nutritional need, nestlings that are parasitized will invest less in begging than those that are not parasitized. We tested this prediction by measuring begging in Pied Flycatcher Ficedula hypoleuca nestlings parasitized with haematophagous mites Dermanyssus gallinoides and Dermanyssus gallinae and blowfly larvae Protocalliphora azurea, and subjected to different levels of food deprivation in order to control for short‐term nutritional need. Nestlings from nests with ectoparasites spent less time begging than those from nests without parasites, especially when very hungry, although there was no association with latency to beg or begging intensity. Our results suggest that time invested in begging may indicate not only the level of need, but also nestling parasitism status.     

Effect of alarm calling by male Red‐winged Blackbirds on nestling begging and female provisioning behavior

ABSTRACT Nestling begging and parental provisioning can attract nest predators and reduce reproductive success, so parents and their offspring might be expected to respond adaptively by minimizing predator‐attracting cues when predators threaten nests. Male Red‐winged Blackbirds (Agelaius phoeniceus) are well known for their antipredator alarm calls that contain information about the approach of potential nest predators. We examined the begging behavior of nestlings and the provisioning behavior of females in response to antipredator alarm calls of males to test the adaptive response hypothesis. Playback experiments provided no evidence that alarm calls function to switch off vocal begging; nestlings were equally likely to beg vocally during playback and control periods. Video recordings showed that male alarm calling had no significant effect on inappropriate vocal begging (in the absence of an adult), but significantly reduced the incidence of spontaneous calling (in the absence of begging). Adult females responded to male antipredator alarm calls by delaying their provisioning visits. In addition, although having no significant effect on use of nest‐arriving calls by females, male alarm calling significantly reduced their use of nest‐leaving calls. We conclude that nestling and female Red‐winged Blackbirds respond to male alarm calls in ways that might reduce the risk of predation, but nestlings beg vocally when females arrive to feed them, regardless of male alarm calling, perhaps to avoid a competitive disadvantage with broodmates.     

Olfactory eavesdropping: The odor of feathers is detectable to mammalian predators and competitors

The role of olfactory eavesdropping in interactions between mammalian predator and prey species is well established. Bird plumage can be odorous and consequently nest predators could use odor to identify and locate avian prey, and nest competitors could use odor to assess occupancy of nest cavities by birds. However, despite extensive research on avian nest predation and competition, the costs of olfactory eavesdropping on plumage odor by nest predators or competitors remain largely unknown. We used two experiments to investigate whether feather odor is detected by marsupial species which are competitors for nest hollows and predators of eggs and nestlings of crimson rosellas, Platycercus elegans. In the first experiment, odor presentation at nest boxes utilized by ringtail possums (Pseudocheirus peregrinus) and rosellas showed that the latency of possums to enter the nest was shorter when crimson rosella odor was present compared to the controls. In the second experiment, carried out away from nest hollows, brushtail possums (Trichosurus vulpecula) discriminated odors of two predators (dingo, Canis lupus dingo, and cat, Felis catus) from crimson rosella and control odors; however, they did not discriminate between crimson rosella odor and a control. We show that marsupials may use feather odor cues to assess nest hollow status, information which could aid their detection of avian prey or their vigilance at nest hollows (for which they compete with parrots). To our knowledge, our study is the first to show that wild mammalian predators and competitors of birds respond to plumage odor at nests and suggest that odor signaling may have hitherto unrecognized costs for birds.     

Green nesting material has a function in mate attraction in the European starling

The function of fresh green nest material has long been debated. It has been suggested that it reduces the number of ectoparasites in nests and on nestlings (nest protection hypothesis), or is used by males to signal condition and paternal quality (male quality hypothesis) or is used as a sexually selected ornament to attract females (courtship hypothesis). We simultaneously tested these three hypotheses in the European starling, Sturnus vulgaris, in the field. Green material was carried by male starlings only, and mainly during nest building. It was not used to reduce ectoparasites. Males nesting in nestboxes that were experimentally contaminated with ectoparasites did not carry more green nest material than males nesting in control boxes, and experimental removal and addition of green material had no effect on the number of ectoparasites on the nestlings or on their body mass. Furthermore, the amount of green material carried into a nestbox was not associated with male body mass, paternal incubation attendance or nestling food provisioning. There is two-fold experimental evidence that males use green plant material to attract mates. First, removal of greenery resulted in a significantly lower percentage of nestboxes containing a clutch than the control or addition treatment. Second, unpaired male starlings sang more and carried more greenery into a nestbox when a caged female was positioned adjacent to the nestbox than when a caged male or an empty cage was present. Paired males, when subjected to the same experimental design, did not respond.     

Building multiple nests is associated with reduced breeding performance in a south temperate population of Grass Wrens Cistothorus platensis platensis

Grass Wrens Cistothorus platensis build two types of non-breeding nest structures: platforms and dummy nests. Platforms are rudimentary accumulations of grasses concealed between vegetation. Dummy and breeding nests are dome-shaped with a similar structural layer. We used a nest-removal experiment and observational data to evaluate several hypotheses regarding the adaptive significance of building multiple nests in a south temperate population of Grass Wrens. Building non-breeding nests was not a strategy of males to attract additional females, as most of these nests were built after pair formation and both sexes collaborated during building. Building non-breeding nests was not a post-pairing display as the presence of multiple nests did not increase female investment in the breeding attempt: clutch size and female provisioning to nestlings did not differ between experimental and control territories where no non-breeding nests were removed. Similarly, in non-manipulated territories, clutch size and female provisioning were not correlated with the number of non-breeding nests or with males’ nest-building effort. Contrary to this hypothesis, the number of non-breeding nests was associated with delayed clutch initiation and reduced hatching success. The presence of non-breeding nests did not reduce nest predation and brood parasitism, which did not differ between experimental and control territories. We did not detect differences in concealment between non-breeding and breeding nests, suggesting that non-breeding nests were not the result of abandonment before egg-laying to reduce subsequent nest predation. Dummy nests did not provide shelter; they were not used frequently for roosting over the breeding season and were not maintained during the non-breeding season. We suggest that building non-breeding nests may be an attempt by males to manipulate the decision of females to breed with a mate they might otherwise reject or to start reproduction earlier than optimal for the females.     

Nest site attributes and temporal patterns of northern flicker nest loss: effects of predation and competition

To date, most studies of nest site selection have failed to take into account more than one source of nest loss (or have combined all sources in one analysis) when examining nest site characteristics, leaving us with an incomplete understanding of the potential trade-offs that individuals may face when selecting a nest site. Our objectives were to determine whether northern flickers (Colaptes auratus) may experience a trade-off in nest site selection in response to mammalian nest predation and nest loss to a cavity nest competitor (European starling, Sturnus vulgaris). We also document within-season temporal patterns of these two sources of nest loss with the hypothesis that flickers may also be constrained in the timing of reproduction under both predatory and competitive influence. Mammalian predators frequently depredated flicker nests that were: lower to the ground, less concealed by vegetation around the cavity entrance and at the base of the nest tree, closer to coniferous forest edges and in forest clumps with a high percentage of conifer content. Proximity to coniferous edges or coniferous trees increased the probability of nest predation, but nests near conifers were less likely to be lost to starlings. Flickers may thus face a trade-off in nest site selection with respect to safety from predators or competitors. Models suggested that peaks of nest predation and nest loss to eviction occurred at the same time, although a competing model suggested that the peak of nest loss to starlings occurred 5 days earlier than the peak of mammalian predation. Differences in peaks of mammalian predation and loss to starlings may constrain any adjustment in clutch initiation date by flickers to avoid one source of nest loss.     

The nest defence by the red‐backed shrike Lanius collurio – support for the vulnerability hypothesis

The majority of altricial bird species defend their brood against predators more intensively in nestlings rather than eggs stage. Several hypotheses have been proposed to explain this difference. The majority of existing experimental studies have recorded a gradually increasing intensity of nest defence supporting the reproductive value hypothesis. We have compared nest defence in two nesting stages of the red‐backed shrike against two predators of adult birds and against two predators of nests. While the nests with nestlings were defended by parents against three out of four predators, nests with eggs were almost not defended at all. This rapid change in parent nest defence supports rather the vulnerability hypothesis, predicting that the threat to nests with nestlings increases rapidly after hatching, as they became more conspicuous due to their begging and parental provisioning. Unlike most of the species tested previously, the red‐backed shrike uses very vigorous mobbing towards predators. We suggest that the occurrence of this active mobbing (strikes, including physical contact) is a good proxy of the current threat to the nest.     

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.     

Wait Until Dark? Daily Activity Patterns and Nest Predation by Snakes

Predation involves costs and benefits, so predators should employ tactics that reduce their risk of injury or death and that increase their success at capturing prey. One potential way that predators could decrease risk and increase benefits is by attacking prey at night when risks may be reduced and prey more vulnerable. Because some snakes are facultatively nocturnal and prey on bird nests during the day and night, they are ideal for assessing the costs and benefits of diurnal vs. nocturnal predation. We used automated radiotelemetry and cameras to investigate predation on nesting birds by two species of snakes, one diurnal and the other facultatively nocturnal. We predicted that snakes preying on nests at night should experience less parental nest defence and capture more adults and nestlings. Rat snakes (Pantherophis obsoletus) were relatively inactive at night (23–36% activity) but nearly always preyed on nests after dark (80% of nest predations). Conversely, racers (Coluber constrictor) were exclusively diurnal and preyed on nests during the times of day they were most active. These results are consistent with rat snakes strategically using their capacity for facultative nocturnal activity to prey on nests at night. The likely benefit is reduced nest defence because birds defended their nests less vigourously at night. Consistent with nocturnal predation being safer, rat snake predation events lasted three times longer at night than during the day (26 vs. 8 min). Nocturnal nest predation did not make nests more profitable by increasing the likelihood of capturing adults or removing premature fledging of nestlings. The disconnect between rat snake activity and timing of nest predation seems most consistent with rat snakes locating prey during the day using visual cues but waiting until dark to prey on nests when predation is safer, although designing a direct test of this hypothesis will be challenging.     

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.     

Experimental study of alarm calls of the oriental tit (Parus minor) toward different predators and reactions they induce in nestlings

Anti-predatory strategies of birds are diverse and may include predator-specific alarm calls. For example, oriental tit (Parus minor) parents can distinguish snakes from other predators and produce snake-specific referential vocalizations ("jar" call) when a snake poses a threat to their nest. The “jar” call has a very specific function to induce fledging of nestlings close to fledging age. This reaction ensures nestlings' survival in natural encounters with snakes that are capable of entering nest cavities and kill entire broods. Sciurid rodents, like chipmunks, may pose a similar threat to cavity-nesting birds. We explored the hypothesis that parents use the fledging-inducing alarm vocalizations in this situation, because chipmunks, like snakes, can kill the brood upon entering the nest cavity. We compared alarm calls of parents toward two predators (chipmunk and snake) who pose a similar threat to the nestlings in a nest cavity, and toward an avian predator (Eurasian jay) who cannot enter nest cavities and poses no threat to the nestlings in a nest. Our results show that the vocal responses of oriental tits were different among the three predators. This suggests that the acoustic properties of vocal responses to predators are different between predators of a similar hunting strategy (nest-cavity entering). The playback of recorded vocal responses of parents to chipmunks did not trigger the fledging of old nestlings, whereas the vocalizations toward a snake did, as shown by earlier studies. Our study suggests that the vocal response of parents does not carry information about the ability of predators to enter the nest cavity and confirms the special status of alarm calls triggered by snakes.     

Calling at a cost: elevated nestling calling attracts predators to active nests

Begging by nestling birds has been used to test evolutionary models of signalling but theory has outstripped evidence. Eavesdropping predators potentially impose a cost on begging that ensures signal honesty, yet little experimental evidence exists for such a cost at active nests because the use of artificial nests, long playback bouts and absence of parents may have exaggerated costs. We broadcast short periods (1 h) of either nestling vocalizations or background noise at active white-browed scrubwren, Sericornis frontalis, nests. Nestlings called naturally during both treatments, allowing us to test whether elevated calling increases risk, a key but rarely tested assumption of evolutionary models. Predators visited nests exclusively during periods of elevated calling. Furthermore, playbacks affected neither adult visits nor nestling activity, suggesting that calling alone attracted predators. Adults gave alarm calls and nestlings usually called less when predators approached nests. Predation risk to broods is, therefore, likely to fluctuate substantially over short periods of time, depending on nestling hunger and whether adults or young have detected predators. This study confirms a present-day cost of nestling begging, demonstrates that this cost can be incurred over short periods and supports the importance of parent-offspring antipredator strategies in reducing predation risk.     

How do predators and scavengers locate resource hotspots within a tropical forest?

In many parts of the world, wildlife species congregate at ‘hotspot’ locations that offer feeding opportunities unmatched in the wider landscape. But to exploit those resource‐rich sites, animals must first locate them. In tropical Australia, predators and scavengers (especially dingos, scrub turkeys, snakes, and invasive toads) gather beneath large canopy‐emergent trees that house breeding colonies of metallic starlings (Aplonis metallica). Some wildlife species feed on fallen nestlings whereas others consume the rich insect fauna supported by bird detritus, or the other species attracted to those resources. Those congregations largely cease as soon as colony trees fall, suggesting that wildlife aggregations are responses to bird‐associated cues rather than to specific locations. To identify the proximate cues that elicit congregation of wildlife under such trees, we deployed sound cues (starling‐chatter) and two types of scent cues (soil from beneath a starling tree, and complete nests on broken branches). We recorded visitations by animals with camera‐traps. Starling‐chatter did not attract significant numbers of animals, but soil from beneath colony trees attracted many animals (mostly scrub turkeys). Complete nests attracted nest‐predators (dingos, snakes). Our experiments suggest that faunal aggregations beneath colony trees are driven by proximate responses to distinctive scent cues in the soil, especially for species that obtain their food from that bird‐fertilized substrate; but predators that feed directly on fallen nestlings key in specifically on that resource.     

Intraseasonal effects of clutch manipulation on parental provisioning and residual reproductive value of eastern phoebes (Sayornis phoebe)

Summary First clutches of double-brooded eastern phoebes Sayornis phoebe were manipulated (up two eggs, down 2 eggs or no change) to test for intraseasonal reproductive tradeoffs and to test whether size of first brood influenced food delivery rates to nestlings and nestling quality in second broods.Considering all nests from both broods, rate of feeding nestlings increased linearly with brood size but nestling mass per nest decreased with increasing brood size. High nestling weights in small broods may have resulted from parents delivering better quality food, but we did not test this.Among treatment groups in first broods, nestlings from decreased broods weighed more than those in control or increased broods. Treatment did not influence the likelihood that second nests would be attempted after successful first nests nor did it alter the interval between nests. Nestlings of parents that renested weighed more than those of parents that did not, regardless of treatment, suggesting that post-fledging care may preclude renesting. Mass of individual females did not change between broods, regardless of brood size. Clutch sizes of second attempts were not affected by manipulations of first broods but increasing first broods reduced the number of nestlings parents were able to raise to day 11 in their second broods. However, manipulation of first broods did not affect mean nestling mass per nest of nestlings that survived to day 11.In phoebes, parents of small first broods are able to raise nestlings in better condition. We predict that in harsh years, parents of small first broods would be more likely to renest. Parents of enlarged first broods sacrificed quality of offspring in second broods, which seems a reasonable strategy if nestlings from second broods have lower reproductive value.     

Diets of adult and nestling starlings (Sturnus vulgaris) in Hawke’s Bay,New Zealand

Abstract

Gizzards were examined from 334 adult and 62 nestling starlings collected in mixed farmland during 1971–72. The birds ate insects, spiders, earthworms, snails, millipedes, centipedes, seeds, and fruits. Starlings ate fewer subterranean animals (7%) than those usually living partially hidden (45%), on the ground (31%), or on vegetation (18%). About half the invertebrates eaten were 2–5 mm long and about a quarter 6–10 mm long. Nestlings tend to be fed significantly larger items than were eaten by adults. The commonest items in adult starlings were Coleoptera adults, Lepidoptera larvae, Hemiptera, and fruits; in nestlings, Coleoptera adults and Diptera adults and larvae were important. Earthworms were found in all the nestlings. More than 50% of adult gizzards contained earthworm chaetae in wetter months, but fewer in drier months. The diet of starlings, despite considerable overlap with the foods eaten by mynas, rooks, and magpies, included somewhat different components and proportions of the food supply. Fruits were found in adults only; most were probably eaten after harvest. Although predation on two major pasture pests—Costelytra zealandica and Wiseana cervinata larvae—was insubstantial, nearly 40% of the total invertebrates eaten by adult starlings were insect pest species.     

Behaviour of the haematophagous mite Ornithonyssus bursa in starling nest boxes in New Zealand

Abstract

Ornithonyssus bursa (Berlese) (Mesostigmata: Dermanyssidae) is a continuous ectoparasite of starlings, and infests most of their nests. Visits by non-resident starlings are considered to be the principal means by which mites spread to uninfested nests during the breeding season. Mites occur between the clenched toes of young nestlings, but as the nestlings' feathers grow and their preening ability develops, mites shift to between primaries and to folds of skin under the bill. After the nestlings fledge, mites migrate to the upper surfaces of the nest box, where some are able to regain contact with adult starlings which daily revisit their nests.