Is the denser contour feather structure in pale grey than in pheomelanic brown tawny owls Strix aluco an adaptation to cold environments?

In colour polymorphic species morphs are considered to be adaptations to different environments, where they have evolved and are maintained because of their differential sensitivity to the environment. In cold environments the plumage insulation capacity is essential for survival and it has been proposed that plumage colour is associated with feather structure and thereby the insulation capacity of the plumage. We studied the structure of contour feathers in the colour polymorphic tawny owl Strix aluco. A previous study of tawny owls in the same population has found strong selection against the brown morph in cold and snowy winters whereas this selection pressure is absent in mild winters. We predicted that grey morphs have a denser and more insulative plumage, enabling them to survive better in cold climate compared to brown ones. The insulative plumulaceous part of the dorsal contour feathers was larger and the fine structure of the plumulaceous part of the feather was denser in grey tawny owls than in brown ones. In the ventral contour feathers the plumulaceous part of the feather was denser in females than in males and in older birds without any differences between morphs. Our study suggests that insulative microscopical feather structures differ between colour morphs and we propose that feather structure may be a trait associated with morph‐specific survival in cold environments.     

Exposure affects the risk of an owl being mobbed – experimental evidence

Mobbing is a widespread anti-predator strategy in birds, and predators are generally expected to avoid mobbing. For example, observational studies suggest that the cryptic roosting behaviour of nocturnal predators, such as many owls, may be a strategy to limit mobbing. In this paper, we present the results of the first experimental study investigating to what degree roost exposure influences the risk of being mobbed, and the intensity of a mobbing incidence once initiated. To determine these factors, we used an experimental setup with taxidermic mounts of tawny owls Strix aluco in Grib Skov forest, Denmark. The risk of an owl being mobbed during a 50 min morning survey period increased with the exposure of its roosting position, from 24% when hidden to 85% when openly exposed. The corresponding increase in the afternoon was from 6% to 36%. This suggests that an owl may minimize the mobbing rate by reducing the encounter rate with potential mobbers through its choice of roost. Once initiated, the duration of the mobbing (a proxy for the presumed cost of being mobbed) was independent of the roosting position of the mounted owl, but was positively correlated with the number of birds in the mob.     

Telomere length in relation to colour polymorphism across life stages in the tawny owl

Telomere erosion has been proposed to be tightly associated with senescence, environmental stressors and life history trade‐offs. How telomere dynamics vary across life stages and especially in relation to (heritable) phenotypic traits is still unclear. The tawny owl Strix aluco display a highly heritable melanin‐based colour polymorphism, a grey and a brown morph, linked to several fitness traits including morph‐specific telomere dynamics. As adults, brown tawny owls have shorter relative telomere length (RTL) and exhibit faster telomere shortening rate than grey owls. Here we test if these morph‐specific telomere dynamics emerge already during growth, or if they are induced only in adult life through differential physiological costs associated with the life history of the morphs. We analysed RTL from 287 tawny owl offspring and 81 first breeding adults to evaluate at what life stage morph‐specific patterns emerge. We found no differences in RTL between the two morphs during the nestling period nor at the first breeding attempt. Sex, brood size or size rank in the nest did not affect offspring RTL. Among first‐breeders, females had shorter telomeres than males suggesting a sampling‐time dependent difference in reproductive costs between sexes, due to the prominent sex roles in tawny owls in the early nestling period. The probability to return to breed after the first breeding attempt was not affected by RTL, sex or colour morph. The lack of morph‐specific difference in RTL among nestlings and first breeders suggests that previously observed morph‐specific differences in RTL dynamics in adults emerge at the onset of the breeding career and is likely due to different physiological profiles and life‐history strategies adopted by adults. We conclude that different telomere dynamics and senescence patterns among highly heritable phenotypes (colour morphs) are likely to be a result of differential costs of reproduction and self‐maintenance.     

Exposure affects the risk of an owl being mobbed – experimental evidence

Mobbing is a widespread anti‐predator strategy in birds, and predators are generally expected to avoid mobbing. For example, observational studies suggest that the cryptic roosting behaviour of nocturnal predators, such as many owls, may be a strategy to limit mobbing. In this paper, we present the results of the first experimental study investigating to what degree roost exposure influences the risk of being mobbed, and the intensity of a mobbing incidence once initiated. To determine these factors, we used an experimental setup with taxidermic mounts of tawny owls Strix aluco in Grib Skov forest, Denmark. The risk of an owl being mobbed during a 50 min morning survey period increased with the exposure of its roosting position, from 24% when hidden to 85% when openly exposed. The corresponding increase in the afternoon was from 6% to 36%. This suggests that an owl may minimize the mobbing rate by reducing the encounter rate with potential mobbers through its choice of roost. Once initiated, the duration of the mobbing (a proxy for the presumed cost of being mobbed) was independent of the roosting position of the mounted owl, but was positively correlated with the number of birds in the mob.     

Response of an arctic predator guild to collapsing lemming cycles

Alpine and arctic lemming populations appear to be highly sensitive to climate change, and when faced with warmer and shorter winters, their well-known high-amplitude population cycles may collapse. Being keystone species in tundra ecosystems, changed lemming dynamics may convey significant knock-on effects on trophically linked species. Here, we analyse long-term (1988–2010), community-wide monitoring data from two sites in high-arctic Greenland and document how a collapse in collared lemming cyclicity affects the population dynamics of the predator guild. Dramatic changes were observed in two highly specialized lemming predators: snowy owl and stoat. Following the lemming cycle collapse, snowy owl fledgling production declined by 98 per cent, and there was indication of a severe population decline of stoats at one site. The less specialized long-tailed skua and the generalist arctic fox were more loosely coupled to the lemming dynamics. Still, the lemming collapse had noticeable effects on their reproductive performance. Predator responses differed somewhat between sites in all species and could arise from site-specific differences in lemming dynamics, intra-guild interactions or subsidies from other resources. Nevertheless, population extinctions and community restructuring of this arctic endemic predator guild are likely if the lemming dynamics are maintained at the current non-cyclic, low-density state.     

No phylogeographic structure in the circumpolar snowy owl (Bubo scandiacus)

The snowy owl (Bubo scandiacus) is a nomadic species with a circumpolar distribution. It has recently declined in the western Palearctic and may thus be worthy of special consideration for conservation. We investigated genetic structure in three well separated geographic regions within the snowy owls’ breeding range. We sequenced two mitochondrial genes; the control region and cytochrome b, and two Z-chromosome introns; VLDLR-9 and BRM-15. We found no phylogeographic structure among the sampled regions, indicating high levels of gene flow in the recent past and possibly still today. Intra-population diversity did not vary between regions for the control region, but for Cyt b, North American birds had higher haplotype diversity than Scandinavian and eastern Siberian birds. Western Palearctic birds do not seem to be genetically deprived or inbred. Genetic diversity in the snowy owl was not lower than Scandinavian populations of three other owl species: tawny owls (Strix aluco), Tengmalm’s owls (Aegolius funereus) and eagle owls (Bubo bubo).     

Not all colors are equal: predation and color polytypism in the aposematic poison frog Oophaga pumilio

Aposematic organisms are not predicted to show high levels of warning signal diversity because they are expected to be under stabilizing selection to decrease costs of ‘educating’ predators about their unpalatability. However, systematic changes in warning signals (polytypism) can be expected if they represent adaptations to local predators. The aposematic strawberry poison frog (Oophaga pumilio) is red throughout its mainland distribution in Costa Rica and Panamá, but displays high levels of warning signal diversity in the Bocas del Toro Archipelago of Panamá. Both coloration and spot pattern vary in a polytypic sense. Sexual selection contributes to maintaining the polytypism, but little work has investigated the potential influence of predation. We used unspotted models of O. pumilio to determine if predation might help explain the color polytypism on Isla Colón in the Bocas del Toro Archipelago of Panamá. We tested whether attack rates differed among the red mainland morph, green/yellow Isla Colón morph, and the brown control. We found that frog color significantly predicted being attacked. The local green Isla Colón models were attacked more than foreign red or brown models. No difference in attack rate existed between red and brown control models. Our results suggest that the red mainland morph possesses a more effective warning signal, even when it is not the local morph. Honest signaling of unpalatability, neophobia, and the use of search images by local predators are potential explanations. Similarity of the brown model to other local poison frogs might explain the lower attack rate compared to previous work. The attack rate was lower on Isla Colón compared to mainland Costa Rica, which supports the hypothesis that less overall predation in the Bocas del Toro Archipelago may contribute to the overall warning signal diversity in O. pumilio there by relaxing selection for aposematic traits.     

Predation on rodents in Białowieza primeval forest, Poland

Autumn and winter predation on bank vole Clethrtonomys glareolus and yellow-necked mouse Apodemus flavicollis was studied in 1985/86-88/89 in an 11 2 km² area of the Bialowieza National Park Rodents regularly increased in numbers from spring to autumn and decreased throughout winter Out of 23 species of predators, the most common were tawny owl Strix aluco (43-57 adult ind 10 km⁻²) weasel Mustela ntvahs (17-27 ind), buzzard Buteo buteo (12-16) and pine marten Martes martes (5-8) Voles and mice were the staple food for two specialists the stoat Mustela erminea and the weasel, and two generahsts the tawny owl and the pine marten The generalists exploited different alternative prey when rodents were scarce tawny owl -amphibians, marten - small mammals, and the red fox Vulpes vulpes - ungulate carcasses and hares The depth of snow and abundance of voles were two major factors shaping the contribution of voles to tawny owl and marten diets No such relationships were found for mice and generalist predators The predation by 8 species (tawny owl, buzzard, marten, weasel, stoat, polecat Mustela putortus fox, and raccoon dog Nyctereutes procyonoides) from 1 October to 15 April in 1986/87, 87/88, and 88/89 was estimated to be on average 28-35 voles and 14-17 mice ha Estimates suggested that three species were responsible for 86-95% of the total predation impact tawny owl (56-71%), weasel (11-21%) and marten (10-15%) The predation impact was similar to the annual decrease in rodent numbers from autumn till spring voles 35 (SD 111) lnds ha^-1in autumn and 8 (SD 2 6) lnds ha^-1in spring, and mice 24 (SD 16 3) mds ha⁻¹ in autumn and 3 (SD 2 5) in spring Predation, therefore, was regarded as the main agent of rodent mortality throughout autumn and winter     

Diurnal exposure as a risk sensitive behaviour in tawny owls Strix aluco?

Tawny owls Strix aluco generally roost in cryptic locations during the day. To test the hypothesis that this cryptic behaviour is an effort to avoid mobbers or avian predators, we measured diurnal behaviour and cause-specific mortality of radio-tagged birds. Non-breeding adults (assumed to be well fed individuals, optimising their own survival) roosted in less exposed locations than adults with young and newly independent juveniles. Parents roosted in the most exposed sites when their young were immature and vulnerable to depredation, probably to guard offspring. Newly independent juveniles apparently selected roosting sites in exposed places to get access to food, as this behaviour was associated with lower perching heights and higher prey abundance beneath their roosting sites. They also perched in more exposed sites, closer to the ground, in summers with low prey abundance compared to summers with high prey abundance. After previous encounters with goshawks Accipiter gentilis , dependent juveniles roosted in less exposed places compared to other young. The increased risk of being mobbed was highly significant with increasing roosting exposure. Once an owl was mobbed, the intensity of the mobbing correlated positively with the mass of the mobbers, but mobbing birds never killed any owls. In contrast, diurnal raptors caused 73% of natural owl deaths (n=15) and the depredation rate by raptors was 3.8 times higher in population classes that generally roosted in more exposed locations than did non-breeding adults. We therefore suggest that depredation by diurnal raptors is the main factor shaping the diurnal behaviour of tawny owls.     

Effects of avian mobbing on roost use and diet of powerful owls,Ninox strenua

We observed the species and numbers of mobbing birds and their effects on a large, nocturnal, bird-eating predator, the powerful owl, together with the pattern of owl predation on mobbing and non-mobbing species. Owls were mobbed on 35 occasions by seven of 44 species of forest birds at a site composed of open forest (88% by area) and rainforest (12%). The majority of bouts involved individuals of a single species, although mixed groups were observed on nine occasions. Regular mobbers were between 4 and 26% of the owls’ body weight. Owls abandoned their daytime roosts during 20% of bouts and responded by calling or actively monitoring mobbers during 54% of bouts. Mobbing appeared to explain why owls roosted in rainforest significantly more often than expected by its availability, mobbing being significantly less frequent in rainforest than in open forest. Only one mobbing species regularly occupied rainforest and the canopy of roosts in rainforest was denser than that in open forest, thus reducing the chances of an owl being detected by potential mobbers. Twelve species of forest birds were within the range of prey size of the powerful owl (75–800 g): six were mobbers and six non-mobbers. The frequency of owl predation on non-mobbers was 8.75 times that on mobbers. The species in this study took a high risk by mobbing a very large predator, but benefited by greatly reducing their chances of predation.     

Dark-bellied brent geese Branta b. bernicla breeding near snowy owl Nyctea scandiaca nests lay more and larger eggs

Several studies have demonstrated that snowy owls Nyctea scandiaca defend an area around their nests against predators, hereby inadvertently creating safe havens for breeding dark-bellied brent geese Branta b. bernicla . However, studies investigating brent goose breeding ecology within the predator-exclusion zones of the snowy owls are absent. In 1999 and 2005, years of high lemming abundance Lemmus sibiricus and Dicrostonyx torquatus , brent geese were primarily breeding in association with snowy owls in the Medusa river catchment on western Taimyr, Russia. Goose nest failure, either as a result of nest abandonment by the adult birds or of nest depredation, increased with increasing distance from the owl nests. Within the brent goose colonies, clutch size as well as egg size increased with decreasing distance from the snowy owl nest, indicating an increasing adult quality closer to owl nests. However, as a result of the abandonment of eggs and goslings, the increasing clutch size did not result in a higher nest success during this study. Apparently brent geese compete for breeding sites close to owl nests, but details of this process remain unknown.     

Mechanisms determining the spatial distribution of microtine predators on the Arctic tundra

1. We studied the spatial distribution of avian microtine predators using data from 19 study areas on the tundra of northern Siberia.
2. Numbers of snowy owls, and long-tailed skuas and pomarine skuas depended strongly on lemming density. However, a significant relationship between lemming density and number of rough-legged buzzards appeared first after removal of the effect of snowy owl abundance on the distribution of rough-legged buzzards.
3. We applied a recently developed method (Manly 1995) to examine co-occurrences of species and found that rough-legged buzzards and snowy owls did not co-occur while snowy owls, long-tailed skuas and pomarine skuas did.
4. There are large differences in nest construction and chick behaviour between rough-legged buzzards and the three other species. Moreover, the snow owl is a polyphagous predator preying also on large birds including raptor chicks. Therefore, we propose that reduced risk of nest predation favours rough-legged buzzards nesting away from snowy owls.
5. Variations in abundance of the two lemming species did not seem to influence the distributions of snowy owls and rough-legged buzzards. Neither was it likely that latitudinally related factors such as breeding season length affected the distribution of rough-legged buzzards.     

Brown tawny owls moult more flight feathers than grey ones

The mechanisms by which melanin‐based colour polymorphism can evolve and be maintained in wild populations are poorly known. Theory predicts that colour morphs have differential sensitivity to environmental conditions. Recently it has been proposed that colour polymorphism covaries genetically with intrinsic and behavioural properties. Plumage moult is a costly and crucial somatic maintenance function in birds. We used a long‐term data set consisting of 761 observations on 307 individuals captured between 1985 and 2010 to examine differences in partial flight feather moult between grey (pale) and brown (pheomelanic dark) colour morphs of the tawny owl. We find that the brown morph consistently moult more primary flight feathers than the grey morph whereas there is no clear difference between colour morphs in the moulting of secondary feathers. Contrary to expectations, the difference in the number of moulted flight feathers between the morphs was independent of environmental conditions, as quantified by the abundance of prey. We discuss the potential physiological and behavioural causes for and costs of the observed difference in maintenance functions between colour morphs.     

Spatial analyses of two color polymorphisms in an alpine grasshopper reveal a role of small‐scale heterogeneity

Discrete color polymorphisms represent a fascinating aspect of intraspecific diversity. Color morph ratios often vary clinally, but in some cases, there are no marked clines and mixes of different morphs occur at appreciable frequencies in most populations. This poses the questions of how polymorphisms are maintained. We here study the spatial and temporal distribution of a very conspicuous color polymorphism in the club‐legged grasshopper Gomphocerus sibiricus. The species occurs in a green and a nongreen (predominately brown) morph, a green–brown polymorphism that is common among Orthopteran insects. We sampled color morph ratios at 42 sites across the alpine range of the species and related color morph ratios to local habitat parameters and climatic conditions. Green morphs occurred in both sexes, and their morph ratios were highly correlated among sites, suggesting shared control of the polymorphism in females and males. We found that in at least 40 of 42 sites green and brown morphs co‐occurred with proportions of green ranging from 0% to 70% with significant spatial heterogeneity. The proportion of green individuals tended to increase with decreasing summer and winter precipitations. Nongreen individuals can be further distinguished into brown and pied individuals, and again, this polymorphism is shared with other grasshopper species. We found pied individuals at all sites with proportions ranging from 3% to 75%, with slight, but significant variation between years. Pied morphs show a clinal increase in frequency from east to west and decreased with altitude and lower temperatures and were more common on grazed sites. The results suggest that both small‐scale and large‐scale spatial heterogeneity affects color morph ratios. The almost universal co‐occurrence of all three color morphs argues against strong effects of genetic drift. Instead, the data suggest that small‐scale migration–selection balance and/or local balancing selection maintain populations polymorphic.     

Interspecific offspring killing in owls

In Baranya County (Southern Hungary), tawny owls (Strix aluco) and barn owls (Tyto alba) sequentially use the same nest boxes in a significant number of cases. A total of 460 broods were observed between 1996 and 2003 and, in 12 cases, whole broods of dead tawny owl chicks were registered that had apparently been killed. On investigating the reproductive life characteristics, population sizes, and frequency of killing in these two species, it was concluded that: (1) with growing barn owl population, the number of sequential broods increases but changes in tawny owl population size have no effect on the frequency of sequential broods; (2) the number of killings depends on the number of sequential broods; and (3) with growing barn owl population, the number of killings also grows and this change is unaffected by the size of the tawny owl population. However, no killing occurs as long as 50–60% of the nest boxes are unoccupied. There is no killing either until the percentage of nest boxes occupied by barn owls reaches 40%, although a threshold value like this cannot be shown for the tawny owl. In the cases when a barn owl breeding follows the tawny owl's, the percentage of killing is significantly higher compared to that when barn owls do not breed in the same box. These results indicate that barn owls kill the offspring of tawny owls. By these means, they obtain a breeding place earlier than without killing the chicks of the other species, and this results in higher reproductive success. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 488–494.     

Positive frequency-dependent selection on warning color in Alpine leaf beetles

Müller's theory of warning color and mimicry, despite forming a textbook example of frequency-dependent selection, has rarely been demonstrated in the wild. This may be largely due to the practical and statistical difficulties of measuring natural selection on mobile prey species. Here we demonstrate that this selection acts in alpine beetle communities by using tethered beetles exposed to natural predators. Oreina gloriosa leaf beetles (Coleoptera: Chrysomelidae) possess chemical defense in the form of cardenolides, accompanied by what appears to be warning color in bright metallic blues and greens. Individuals that match the locally predominant color morph have increased survival, with odds of week-long survival increased by a factor of 1.67 over those that do not match. This corresponds to selection of 13% against foreign morphs. Such selection, acting in concert with variation in community composition, could be responsible for geographic variation in warning color. However, in the face of this purifying selection, the within-population polymorphism seen in many Oreina species remains paradoxical.     

Wintering space use and site fidelity in a nomadic species,the snowy owl

Migratory species can exploit many habitats over vast geographic areas and adopt various patterns of space and habitat use throughout their annual cycle. In nomadic species, determinants of habitat use during the non‐breeding season are poorly known due to the unpredictability of their movement patterns. Here, we analysed variability in wintering space and habitat use by a highly nomadic species, the snowy owl, in eastern North America. Using 21 females tracked by satellite telemetry between 2007 and 2016, we 1) assessed how space use patterns in winter varied according to the type of environment (marine vs terrestrial), latitudinal zone (Arctic vs temperate), local snow conditions and lemming densities and 2) investigated winter habitat and site fidelity. Our results confirmed a high inter‐individual variation in patterns of habitat use by wintering snowy owls. Highly‐used areas were concentrated in the Arctic and in the marine and coastal environments. Owls wintering in the marine environment travelled over longer distances during the winter, had larger home ranges and these were divided in more smaller zones than individuals in terrestrial environments. Wintering home range sizes decreased with high winter lemming densities, use of the marine environment increased following high summer lemming densities, and a thick snow cover in autumn led to later settlement on the wintering ground. Contrary to expectations, snowy owls tended to make greater use of the marine environment when snow cover was thin. Snowy owls were highly consistent in their use of a given wintering environment and a specific latitudinal zone between years, but demonstrated flexibility in their space use and a modest site fidelity. The snowy owls’ consistency in wintering habitat use may provide them with advantages in terms of experience but their mobility and flexibility may help them to cope with changing environmental conditions at fine spatial scale.     

The Adaptive Significance of Avian Mobbing V. An Experimental Test of the ‘Move On’ Hypothesis

This paper presents the first experimental study of the effects of mobbing on predators. Two captive little owls Athene noctua and a tawny owl Strix aluco, having stabilized activity patterns and roosting sites, were singly confronted with one or two mobbing blackbirds Turdus merula. The little owls showed marked stress (gular fluttering, panting and panic flights), abandoned chosen roosts and remained unsettled for several days. A permanent alternative roost was chosen after one mobbing encounter, other encounters resulted in several days of ‘floating’ before returning to the original sites. One little owl used ‘camouflage’ by ‘freezing’ during repeated harassments, the other tended to retaliate if mobbing was close and weak. The tawny owl seemed to ignore the rather weak blackbird mobbing, but abandoned its roost for up to 36 h when mobbed by larger numbers of woodland birds at close range. It is concluded that mobbing causes profound distress, especially in small, vulnerable predators like the little owl and forces it to change its daytime roost. If abandoning the area is not possible, time and energy are expended in increased vigilance as a protection against further mobbing attacks. Thus there is strong support for the ‘Move on’ hypothesis. Results also support the ‘Perception advertisement’ hypothesis, implying that the detected hunter deliberately forfeits the surprise element. Which effect prevails depends on predator evaluation of the signals involved.     

The role of selection in the colour polymorphism of Littorina rudis Maton and Littorina arcana Hannaford-Ellis (Prosobranchia: Littorinidae)

Forty-one mixed samples of winkles containing the closely related species, Littorina rudis and L. arcana , were collected from different parts of the British Isles. Littorina rudis was the more ubiquitous species, with L. arcana being more-or-less confined to vertical cliffs and rocks. The frequencies of different shell colour patterns were determined for both species in each sample. Several colour morphs were diagnostic of one or other of the species over large areas, which confirms that these are separate species. The frequencies, in the two species, of the two commonest morphs, brown and fawn, were strongly correlated; in several other morphs their presence and absence in the two species was significantly associated; the levels of phenotypic diversity in the two species were also correlated. These patterns could not have arisen if the variation between shores was the result of genetic drift or founder effects, so some form of selection is implicated. In the brown morph there is some evidence for frequency-dependent selection, and it is suggested that the polymorphism may be maintained by visual predators through apostatic selection.