Female colour polymorphism covaries with reproductive strategies in the tawny owl Strix aluco

Variation in coloration with a strong underlying genetic basis is frequently found in birds, insects, anurans, molluscs and plants. Although such a variation can be large, little is known about its functional value. Correlative data, however, can help suggest testable hypotheses about potential covariation between reproductive parameters and a colour polymorphism displayed by individuals belonging to a single population. In this context, we studied two Swiss populations of tawny owls Strix aluco , a polymorphic species that varies in coloration from reddish-brown to grey. Observations in the first population showed that although greyer females had shorter tarsi, they produced heavier offspring in two of three years. Pairing with respect to plumage coloration was not significantly disassortative, indicating that these correlations were probably not inflated by plumage coloration of the mate. In the second population, where breeding females had been monitored for 14 years, the proportion of all breeding females that were reddish-brown was greater in years when the breeding density was lower. Capture-recapture analyses show that the latter result is explained by the fact that greyish females bred less often than reddish-brown females, although their survival probability was similar. The number of greyer breeding females was greater when spring/summer temperatures were lower. When combined, the results from the two populations lend support to the hypothesis that grey females do not breed every year, but produce offspring of higher quality. Whatever the mechanism underlying the correlations reported in this study, colour polymorphism in female tawny owls appears to reflect some components of individual quality.     

Pre-hatching maternal effects inhibit nestling humoral immune response in the tawny owl Strix aluco

Although evidence is accumulating that mothers can transfer antibodies to their offspring, little is known about the consequences of such a transfer to the offspring immune system. Because maternal antibodies are effective only during a short period of time after their transfer to offspring, one hypothesis is that maternal antibodies provides a transitory antigen-specific protection to offspring, thus lessening the need for offspring to mount their own humoral immune response towards these specific antigens. In birds, this scenario predicts that offspring immune response towards a specific antigen is inhibited to a larger extent in hatchlings than in older nestlings. We tested this hypothesis in tawny owls Strix aluco by cross-fostering clutches between nests and then challenging siblings with a vaccine either two times (at 4- and 11-d-old) or only one time at 11-d-old to compare the strength of the humoral response between nestlings born from mothers with naturally high and low levels of antibodies against this vaccine. Because maternal antibodies are expected to be effective only during a short period of time after hatching, we predict that maternal antibodies should inhibit the immune response of nestlings vaccinated from the fourth day after hatching more than in nestlings vaccinated only at a later age. As expected, the inhibitory effect of maternal antibodies was stronger in nestlings vaccinated soon after hatching than in siblings injected at a later age. Therefore, in wild avian populations pre-hatching maternal effects may confer offspring with a transitory immune protection in the first days following hatching.     

Impact of habitat fragmentation on activity and hunting behavior in the tawny owl, Strix aluco

I examined the effect of woodland fragmentation on activityand hunting behavior in the tawny owl. I made three predictionson the effect of fragmentation, two of which were upheld andone rejected. Over 2 years, 24 owls were radio tracked in thenonbreeding season. Eight of these were in continuous woodlandand the rest in highly fragmented woodland. For each owl, Irecorded data on small mammal availability, diet, home rangesize, perch time, and interperch distance. In accordance withthe first two predictions, owls in fragmented woodlands hadlonger interperch distances and perch times, but contrary tothe third prediction, perch times were positively correlatedwith small mammal density. This was thought to be due to owlsaltering their hunting to other prey types when small mammalswere scarce. Within fragmented woodlands, perch times were thesame between the sexes, but males had slightly greater interperchdistances than females. Male owls in fragmented woods flew 40%further per hour than males in continuous woodland. For datathroughout the night, food availability and area of trees inhome range accounted for 65% of the variation in perch times.For data before 2200 h, area of trees alone accounted for thesame amount of variation. The data suggest that habitat fragmentation,particularly through reducing woodland area, can greatly influenceowl activity and hunting behavior.     

Contrasting patterns of mitochondrial and microsatellite genetic structure among Western European populations of tawny owls (Strix aluco)

A recent study of mitochondrial phylogeography of tawny owls (Strix aluco) in western Europe suggested that this species survived the Pleistocene glaciations in three allopatric refugia located in Iberia, Italy, and the Balkans, and the latter was likely the predominant source of postglacial colonization of northern Europe. New data from seven microsatellite loci from 184 individual owls distributed among 14 populations were used to assess the genetic congruence between nuclear and mitochondrial DNA (mtDNA) markers. Microsatellites corroborated the major phylogeographical conclusions reached on the basis of the mtDNA sequences, but also showed important differences leading to novel inferences. Microsatellites corroborated the three major refugia and supported the Balkan origin of northern populations. When corrected for differences in effective population size, microsatellites and mtDNA yielded generally congruent overall estimates of population structure (N*ST=0.12 vs. RST=0.16); however, there was substantial heterogeneity in the RST among the seven nuclear loci that was not correlated with heterozygosity. Populations representing the Balkans postglacial expansion interact with populations from the other two refugia forming two clines near the Alps and the Pyrenees. In both cases, the apparent position of the contact zones differed substantially between markers due to the genetic composition of populations sampled in northern Italy and Madrid. Microsatellite data did not corroborate the lower genetic diversity of northern, recently populated regions as was found with mtDNA; this discrepancy was taken as evidence for a recent bottleneck recovery. Finally, this study suggests that congruence among genetic markers should be more likely in cases of range expansion into new areas than when populations interact across contact zones.     

Variation in digestion of prey by the Tawny owl (Strix aluco)

Two Tawny owl chicks were collected from their nests and observations were made on their food consumption, digestion and pellet production. The female chick escaped after three months in captivity; the male continued to provide data for a further 23 months.     

Pleiotropy in the melanocortin system: expression levels of this system are associated with melanogenesis and pigmentation in the tawny owl (Strix aluco)

The adaptive function of melanin‐based coloration is a long‐standing debate. A recent genetic model suggested that pleiotropy could account for covariations between pigmentation, behaviour, morphology, physiology and life history traits. We explored whether the expression levels of genes belonging to the melanocortin system (MC1R, POMC, PC1/3, PC2 and the antagonist ASIP), which have many pleiotropic effects, are associated with melanogenesis (through variation in the expression of the genes MITF, SLC7A11, TYR, TYRP1) and in turn melanin‐based coloration. We considered the tawny owl (Strix aluco) because individuals vary continuously from light to dark reddish, and thus, colour variation is likely to stem from differences in the levels of gene expression. We measured gene expression in feather bases collected in nestlings at the time of melanin production. As expected, the melanocortin system was associated with the expression of melanogenic genes and pigmentation. Offspring of darker reddish fathers expressed PC1/3 to lower levels but tended to express PC2 to higher levels. The convertase enzyme PC1/3 cleaves the POMC prohormone to obtain ACTH, while the convertase enzyme PC2 cleaves ACTH to produce α‐melanin‐stimulating hormone (α‐MSH). ACTH regulates glucocorticoids, hormones that modulate stress responses, while α‐MSH induces eumelanogenesis. We therefore conclude that the melanocortin system, through the convertase enzymes PC1/3 and PC2, may account for part of the interindividual variation in melanin‐based coloration in nestling tawny owls. Pleiotropy may thus account for the covariation between phenotypic traits involved in social interactions (here pigmentation) and life history, morphology, behaviour and physiology.     

Breeding of Tawny Owls Strix aluco in rural and urban habitats in southern Finland

Capsule The annual average breeding frequency, clutch size, offspring production and chick survival of Tawny Owls did not differ between rural and urban nesting territories.

Aims To determine whether the general intensity of human habitation in the territory affects breeding.

Methods Clutch size, offspring production, breeding frequency and prey abundance were determined from 210 rural and 60 urban nesting territories monitored between 1994 and 2006.

Results Fluctuations in the annual average clutch size did not differ between habitats. Clutch size and offspring production paralleled each other in rural habitats but not in urban ones. Annual average clutch size followed the regional spring abundance of small mammals in rural Tawny Owls but not in urban ones. The breeding frequency was higher after mild winters in rural environments but not in urban ones.

Conclusion Over an extended time period, rural and urban habitats were largely of equal quality. In urban environments, however, owls seem to be less affected by the pronounced regional abundance fluctuations of small mammals and weather conditions of the preceding winter that largely govern the breeding of owls elsewhere.     

The importance of body reserves for successful reproduction in the Tawny owl (Strix aluco)

One-quarter of Tawny owl nests fail to hatch young, mainly because the eggs are chilled and/or deserted. In 1973–74 automatic photography was employed at four nests near Oxford to relate the incubation behaviour of females to the ration of prey supplied to them by their mates. The eggs did not hatch in two nests and young fledged from only one of the others. Females were less attentive at the nests which failed during incubation and on average received less prey, but even at successful nests there were some nights when the female was supplied with less than her estimated daily food requirement. Female Tawny owls accumulate large reserves of fat and protein before laying. These buffer against any temporary inability of the male to provide sufficient food during incubation, and enable the female to stay on the nest rather than hunt for herself and risk the eggs becoming chilled. When prey are scarce, many females do not lay at all, and the ultimate factor determining whether breeding takes place may be the female's ability to acquire body reserves sufficient to provide a chance of breeding successfully.     

Does variation of sex ratio enhance reproductive success of offspring in tawny owls (Strix aluco)

Tawny owls, Strix aluco, laid female-biased clutches on territories with more abundant prey (field voles) in June, the month that chicks fledge. This appeared to enhance the subsequent reproductive success of fledglings, as in 1995 there was a significant correlation between the number of chicks fledged by adult females and the June vole abundance in the territory on which they were reared as chicks. This relationship did not hold for males. Since tawny owls lay eggs in March, these results indicate that owls are able to predict the June vole numbers on their territory, and respond by producing more of the sex most likely to gain a long-term benefit when resources are good.     

A case study of male tawny owl (Strix aluco) vocalizations in South Korea: call feature,individuality, and the potential use for census

Vocal individuality has been used as a monitoring tool, and two criteria are a prerequisite: high variation among individuals and low variation within individuals, and vocal consistency within and across seasons. We examined individual variation in the territorial hoot calls of the tawny owl (Strix aluco) to discriminate between males and to assess a possible conservation technique that would allow for monitoring individuals within a study area. The territorial calls were recorded from five males in the Naejang Mountain National Park in South Korea during the breeding season in 2015 and 2016 and analyzed both quantitatively and qualitatively to determine the amount of variation within and between individuals. Our results showed that the territorial calls were specific to individuals within a population and that the acoustic distances between males living in the same territory during the two years were the smallest for the four nesting sites. Our results suggest that territorial calls of the tawny owls are individually identifiable over two years and that this acoustic technique can be useful for monitoring individual site fidelity.     

Parent-offspring conflict over duration of parental care and its consequences in tawny owls Strix aluco

In species defending territories, fitness of newly independent juveniles could depend on phenotypic quality or early access to vacant resources as a result of rapid dispersal. In the first case, parent-offspring conflict will arise when parents cease feeding maturing offspring while these still demand provisioning. In the latter case, the young should decide when to stop begging to search for vacant territories. I radio-tracked 72 juvenile tawny owls from independence to sexual maturity to investigate whether parents or offspring decided the timing of the onset of independence and compare effects of age versus date of independence on survival and reproductive status. Juvenile owls stopped begging when 90–123 d old. This was synchronous within broods, independent of gender or age rank. Independence age of cross-fostered young varied across foster nests but was independent of hatching nest. After independence, young roosted in their parents' territory for 18 d on average before dispersal. This suggests that juvenile owls would rather extend dependency than disperse early. As predicted from this behaviour, age of independence had a positive influence on survival and reproductive status at maturity, whereas there were no effects of date of independence.     

Combining information from range use and habitat selection: sex-specific spatial responses to habitat fragmentation in tawny owls Strix aluco

How individuals respond to habitat heterogeneity is usually measured as variation in range size and by ranking the relative importance of habitat types (habitat selection). The combined effect of how individuals incorporate different habitat types in their home ranges and allocate their time budget between them is rarely derived. Additionally, when home range size varies between individuals, habitat selection analyses might be flawed if foraging decisions are based on variation in absolute rather than proportional availability. We investigated the suitability of standard analytical approaches by measuring the spatial responses of tawny owls to habitat fragmentation. These owls inhabited woodland of various sizes, representing a fragmentation gradient from open farmland with small, isolated woodland patches, to continuous woodland within their home ranges. In 17 territories within open farmland, the available area covered by woodland increased with the square root of the area of open land embraced in the home range. The owls did not display functional response in habitat selection, but females selected woodland more strongly than males. Females utilised woodland 10 times more intensively in farmland than in continuous woods, whereas males utilised farmland woods 3.2 times more intensively. Moreover, females in farmland exploited woodland 3.2 times as intensively as males, apparently because of higher travel costs in open areas. Since the extensive variation in intensity of use as a function of total availability was not indicated from the analysis of habitat selection, we suggest that information about intensity of use be more widely used as a supplementary measure of habitat use patterns than appears to be the practice at present.     

Variation in the male territorial hoot of the Tawny Owl Strix aluco in three English populations

Little is known about owl song. We made sonagrams of the territorial calls of 50 male Tawny Owls Strix aluco from three different areas. Six temporal and four frequency measures of the calls were recorded from the sonagrams. The measures of the calls were then subjected to analysis to try to separate between individual owls and between owls from different areas. We also looked for similarities between calls of neighbouring owls and for any effect of habitat on owl hoots. Individual owls were separated on the basis of their hoots with a high degree of success (98.6% overall), and there were significant differences between areas. Differences were found between calls in woodland and farmland habitats, but these differences were not in the direction expected to increase sound transmission. Calls of neighbouring owls did not resemble each other more than calls from owls that were not in vocal contact, implying that if calls are learned by Tawny Owls, they are learned before dispersal.     

Demographic models of the northern spotted owl (Strix occidentalis caurina)

Summary Calassical demographic methods applied to life history data on the northern spotted owl yield and estimate of the annual geometric rate of increase for the population of λ=0.96±0.03, which is not significantly different from that for a stable population (λ=1.00). Sensitivity analysis indicates that adult annual survivorship has by far the largest influence on λ, followed by the probability that juveniles survive dispersal, and the adult annual fecundity. Substantial temporal fluctuations in demographic parameters have little effect on the long-run growth rate of the population because of the long adult life expectancy. A model of dispersal and territory occupancy that assumes demographic equilibrium is evaluated using data on the amount of old forest habitat remaining in the Pacific Northwest and the current occupancy of this habitat by northern spotted owls. This model is employed to predict the effect of future habitat loss and fragmentation on the population, implying that extinction will result if the old forest is reduced to less than a proportion 0.21±0.02 of the total area in a large region. The estimated minimum habitat requirement for the population is greater than that allowed in management plants by the USDA Forest Service.