Parental preference for red mouth of chicks in a songbird

Parental preferences during feeding and care-giving may select for ornamental traits in young, such as bright coloration. For chicks of coots, there is experimental evidence for this idea. We examined the hypothesis that bright yellow, orange and red mouths of chicks of songbirds have been favoured by feeding preferences in parents. In a field experiment, the orange–yellow mouths of great tit nestlings were dyed brightly red, and the feeding response of parents recorded. In nest boxes with extra daylight through a window, experimental chicks were on average given twice as much food (biomass) as control chicks (sham dyed). In normal nest boxes, the tendency was similar, but not significant. Thus, at least in good light, great tit parents prefer to feed young with red mouths, a preference for colourfulness that helps explain the evolution of bright gapes in chicks of songbirds (passerine birds).     

Contrast versus colour in aposematic signals

Conspicuousness is an important feature of warning coloration. One hypothesis for its function is that it increases signal efficacy by facilitating avoidance learning. An alternative, based on the handicap hypothesis, suggests that the degree of conspicuousness holds information directly about the quality of the prey, and that predators associate and learn about the conspicuousness of the coloration, and not the actual colour pattern. We studied the relative importance of signal contrast and the colours of signals for predator attention during discrimination. We used young chicks, Gallus gallus domesticus, as predators and small blue or red paper cones on either matching or contrasting paper backgrounds as stimuli associated with palatable or unpalatable chick crumbs. In four treatment groups, birds could use either cone and/or background colour, cone colour only, background colour only or cone-to-background contrast as cues for discrimination. Only birds in the contrast treatment failed to learn their discrimination task. Birds that had a choice between cone and background colour as cues used the cone colour and they learned the task faster than did birds that had to use background colour as a cue. The results suggest that birds primarily attend to the colours of signals and disregard contrast in discrimination tasks; they thus fail to support a handicap function of conspicuous aposematic coloration. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Do colours that deter birds affect cereal bait acceptance by possums (

Poisonous baits used for pest control in New Zealand commonly contain green dye and cinnamon oil to make them less attractive to birds. Research aimed at reducing the impact of poison based pest control on birds has shown that some birds are initially deterred from feeding on blue or, to a lesser extent, green coloured food and are attracted to yellow or red food. We determined whether colours that deter or attract birds affected the acceptance of non-toxic and toxic cereal baits by captive brushtail possums (Trichosurus vulpecula). Individual possums were offered, daily, a choice between a standard green dyed non-toxic cereal bait and either a blue dyed(17 possums) or yellow dyed non-toxic bait (16 possums) for 10 days. Following this, for the first group of 17 possums, 1080 toxin was added to either the green bait (9 possums) or blue bait (8 possums) and possums were offered the green versus blue choice again. Two additional groups that had not previously been fed cereal baits were also given a choice between blue and green baits, one of which was toxic. All possums offered non-toxic bait ate less on the first day of presentation than on subsequent days. There was no difference in acceptance of either blue or yellow coloured non-toxic bait compared to the standard green non- toxic bait on any days. Bait colour appeared to be unimportant in cereal bait choice and did not deter possums from eating any of the baits. The addition of toxin to baits did not significantly alter bait choice in any groups, although some individuals which had no previous experience with baits ate more toxic than non-toxic bait. These data suggest that adding a stronger bird deterring colour (i.e., blue) to poisonous baits is unlikely to adversely affect the acceptance of baits by possums.     

Red oilseed rape? The potential for manipulation of petal colour in control strategies for the pollen beetle (Meligethes aeneus)

The pollen beetle (Meligethes aeneus) is a major pest of oilseed rape (Brassica napus) at the inflorescence stage and is well known to prefer colours called yellow by human observers over many other colours. While commercial cultivars of oilseed rape have yellow flowers, little is known about the potential to manipulate host plant location and reduce subsequent infestation by this pest through variation in flower colour. We investigated the responses of pollen beetles to flowers of a white-petalled oilseed rape variety that had been dyed different colours in semi-field arena and field experiments. Flowers dyed blue or red were less heavily infested than those dyed yellow or the white flowers, indicating that blue and red flowers were less attractive than yellow and white ones. This response was most likely due to differences in petal colour because olfactometer studies showed that beetle responses to the odours of the coloured treatments did not differ. The comparatively high infestation of untreated white flowers is interpreted as a consequence of their high UV reflectance; the presence of a UV receptor in M. aeneus is suggested, and its role in visually guided insect–plant interactions in this species described. The potential for manipulation of petal colour in control strategies for the pollen beetle is discussed.     

Colour Vision Impairment in Young Alcohol Consumers

Alcohol consumption among young adults is widely accepted in modern society and may be the starting point for abusive use of alcohol at later stages of life. Chronic alcohol exposure can lead to visual function impairment. In the present study, we investigated the spatial luminance contrast sensitivity, colour arrangement ability, and colour discrimination thresholds on young adults that weekly consume alcoholic beverages without clinical concerns. Twenty-four young adults were evaluated by an ophthalmologist and performed three psychophysical tests to evaluate their vision functions. We estimated the spatial luminance contrast sensitivity function at 11 spatial frequencies ranging from 0.1 to 30 cycles/degree. No difference in contrast sensitivity was observed comparing alcohol consumers and control subjects. For the evaluation of colour vision, we used the Farnsworth-Munsell 100 hue test (FM 100 test) to test subject’s ability to perform a colour arrangement task and the Mollon-Reffin test (MR test) to measure subject’s colour discrimination thresholds. Alcohol consumers made more mistakes than controls in the FM100 test, and their mistakes were diffusely distributed in the FM colour space without any colour axis preference. Alcohol consumers also performed worse than controls in the MR test and had higher colour discrimination thresholds compared to controls around three different reference points of a perceptually homogeneous colour space, the CIE 1976 chromaticity diagram. There was no colour axis preference in the threshold elevation observed among alcoholic subjects. Young adult weekly alcohol consumers showed subclinical colour vision losses with preservation of spatial luminance contrast sensitivity. Adolescence and young adult age are periods of important neurological development and alcohol exposure during this period of life might be responsible for deficits in visual functions, especially colour vision that is very sensitive to neurotoxicants.     

How hollow melanosomes affect iridescent colour production in birds

Developmental constraints and trade-offs can limit diversity, but organisms have repeatedly evolved morphological innovations that overcome these limits by expanding the range and functionality of traits. Iridescent colours in birds are commonly produced by melanin-containing organelles (melanosomes) organized into nanostructured arrays within feather barbules. Variation in array type (e.g. multilayers and photonic crystals, PCs) is known to have remarkable effects on plumage colour, but the optical consequences of variation in melanosome shape remain poorly understood. Here, we used a combination of spectrophotometric, experimental and theoretical methods to test how melanosome hollowness—a morphological innovation largely restricted to birds—affects feather colour. Optical analyses of hexagonal close-packed arrays of hollow melanosomes in two species, wild turkeys (Meleagris gallopavo) and violet-backed starlings (Cinnyricinclus leucogaster), indicated that they function as two-dimensional PCs. Incorporation of a larger dataset and optical modelling showed that, compared with solid melanosomes, hollow melanosomes allow birds to produce distinct colours with the same energetically favourable, close-packed configurations. These data suggest that a morphological novelty has, at least in part, allowed birds to achieve their vast morphological and colour diversity.     

Looking like mother makes mallard ducklings dominant over their siblings

Colour variation in time and space among animals may affect social relationships such as pairing and dominance interactions. For instance, some birds are naturally sensitive to leg colour, with some colours being more visible or attractive than others. The colour of the leg-rings used to mark birds may thus be related to behavioural and reproductive variables. Most studies have investigated this effect for adults during reproduction, but leg-ring colour may also affect the behaviour of young birds. We tested the potential effect of leg-ring colours on the within-brood dominance hierarchy of mallard (Anas platyrhynchos) ducklings while each brood formed a stable and exclusive family unit with its mother. Ducklings did not acquire a within-brood dominance rank according to the colour of their own ring. This result suggests that mallards may not have a sensory bias for a given colouration. However, ducklings wearing a ring of the same colour as one of the two rings of their mother were dominant over their siblings. We discuss the potential behavioural and methodological implications of this result.     

Extremely reduced dispersal and gene flow in an island bird

The Réunion grey white-eye, Zosterops borbonicus, a passerine bird endemic to Réunion Island in the Mascarene archipelago, represents an extreme case of microgeographical plumage colour variation in birds, with four distinct colour forms occupying different parts of this small island (2512 km²). To understand whether such population differentiation may reflect low levels of dispersal and gene flow at a very small spatial scale, we examined population structure and gene flow by analysing variation at 11 microsatellite loci among four geographically close localities (<26 km apart) sampled within the distribution range of one of the colour forms, the brown-headed brown form. Our results revealed levels of genetic differentiation that are exceptionally high for birds at such a small spatial scale. This strong population structure appears to reflect low levels of historical and contemporary gene flow among populations, unless very close geographically (<10 km). Thus, we suggest that the Réunion grey white-eye shows an extremely reduced propensity to disperse, which is likely to be related to behavioural processes.     

Impacts of Tree Rows on Grassland Birds and Potential Nest Predators: A Removal Experiment

Globally, grasslands and the wildlife that inhabit them are widely imperiled. Encroachment by shrubs and trees has widely impacted grasslands in the past 150 years. In North America, most grassland birds avoid nesting near woody vegetation. Because woody vegetation fragments grasslands and potential nest predator diversity and abundance is often greater along wooded edge and grassland transitions, we measured the impacts of removing rows of trees and shrubs that intersected grasslands on potential nest predators and the three most abundant grassland bird species (Henslow’s sparrow [Ammodramus henslowii], Eastern meadowlark [Sturnella magna], and bobolink [Dolichonyx oryzivorus]) at sites in Wisconsin, U.S.A. We monitored 3 control and 3 treatment sites, for 1 yr prior to and 3 yr after tree row removal at the treatment sites. Grassland bird densities increased (2–4 times for bobolink and Henslow’s sparrow) and nesting densities increased (all 3 species) in the removal areas compared to control areas. After removals, Henslow’s sparrows nested within ≤50 m of the treatment area, where they did not occur when tree rows were present. Most dramatically, activity by woodland-associated predators nearly ceased (nine-fold decrease for raccoon [Procyon lotor]) at the removals and grassland predators increased (up to 27 times activity for thirteen-lined ground squirrel [Ictidomys tridecemlineatus]). Nest success did not increase, likely reflecting the increase in grassland predators. However, more nests were attempted by all 3 species (175 versus 116) and the number of successful nests for bobolinks and Henslow’s sparrows increased. Because of gains in habitat, increased use by birds, greater production of young, and the effective removal of woodland-associated predators, tree row removal, where appropriate based on the predator community, can be a beneficial management action for conserving grassland birds and improving fragmented and degraded grassland ecosystems.     

Broodcare helpers in the cichlid fish Lamprologus brichardi: Their costs and benefits

‘Helping’ in birds and mammals involves seemingly altruistic behaviour. In the cichlid fish Lamprologus birchardi helpers are usually young of former broods staying in their parents' territories and participating in all kinds of parental duties (Broodcare, territory maintenance and defence). The discovery of helpers in fish offered the chance of attempting an extensive analysis of potential costs and benefits influencing the evolution of helpers in a vertebrate. Three factors proved to be of major importance in the cost-benefit analysis of helping as opposed to leaving for family-independent nonreproductive aggregations. Due to investment and to their rank within a family's hierarchy, helpers grow at a slower rate than non-helpers. This cost is compensated for by (i) a lower mortality risk to helpers caused by their access to a defended shelter and by protection afforded by bigger family members, and (ii) a positive contribution by helpers to the future reproductive success of their parents: females with helpers produce bigger clutches and consequently more free-swimming fry (=siblings). Other variables, such as the helpers' influence on the relative breeding success of their parents, broodcare experience through helping, the chances of territory take-over, parasitism of parents' reproduction and cannibalism are of minor importance. Similar social organizations in other fish are discussed with respect to their ecology and are compared with cooperatively breeding birds and mammals.     

Does avian malaria reduce fledging success: an experimental test of the selection hypothesis

Like many parasites, avian haematozoa are often found at lower infection intensities in older birds than young birds. One explanation, known as the “selection” hypothesis, is that infected young birds die before reaching adulthood, thus removing the highest infection intensities from the host population. We tested this hypothesis in the field by experimentally infecting nestling rock pigeons (Columba livia) with the malaria parasite Haemoproteus columbae. We compared the condition and fledging success of infected nestlings to that of uninfected controls. There was no significant difference in the body mass, fledging success, age at fledging, or post-fledging survival of experimental versus control birds. These results were unexpected, given that long-term studies of older pigeons have demonstrated chronic effects of H. columbae. We conclude that H. columbae has little impact on nestling pigeons, even when they are directly infected with the parasite. Our study provides no support for the selection hypothesis that older birds have lower parasite loads because parasites are removed from the population by infected nestlings dying. To our knowledge, this is the first study to test the impact of avian malaria using experimental inoculations under natural conditions.     

Mechanisms of spacing in groups of juncos: Measurement of behavioural tendencies in social situations

To investigate behavioural mechanisms that regulate the spacing of individuals, we studied aggression, approach, and avoidance of captive male juncos (Junco hyemalis) in relation to the distance between individuals. We measured behavioural tendencies by analysing the terminations of social states so as to determine the instantaneous probability rates of terminal actions. These rates permit accurate comparisons of behavioural tendencies in different social situations. An individual's tendencies to supplant, approach, or withdraw from an opponent depended on the dominance relationship of the two birds and on the distance between them. Interactions did not vary with the magnitude of the difference between opponents' ranks. The dependence of aggression, approach, and avoidance on the distance between opponents controls the spacing of individuals.     

Effects of long-acting testosterone treatment on freeliving pied flycatchers, Ficedula hypoleuca, during the breeding period

Breeding behaviour and spermatogenesis of all males in a population of pied flycatchers, Ficedula hypoleuca, were experimentally prolonged by long-acting testosterone. The behaviour of these birds was compared with that of birds from an untreated population and a population where the males were given the vehicle only. The testosterone treatment prolonged territorial behaviour, and a higher percentage of nests without a feeding male during the nestling period was found in the experimental area. The proportion of polygamous males did not however differ between the experimental and control areas. The feeding activity of those testosterone-treated males returning to their home territories was markedly depressed. The female of such a pair did not compensate for the male's low feeding activity, but fed the nestlings as she would have done with a normal untreated male at the nest. On the other hand, females without a male present during the nestling period compensated for their mates' absence by increasing their own feeding frequency. It was also shown that the song of the male pied flycatcher is dependent upon androgens. The testosterone-treated males produced fewer fledglings than the control males.     

Carotenoid-based plumage colouration is associated with blood parasite richness and stress protein levels in blue tits (Cyanistes caeruleus)

Carotenoids are molecules that birds are not able to synthesize and therefore, must be acquired through their diet. These pigments, besides their function of giving birds red and yellow colouration when deposited in feathers, seem to act as immune-stimulators and antioxidants in the organism. Hence, only the healthiest individuals would be able to express carotenoid-based ornaments to a larger extent without compromising the physiological functions of carotenoids. Various studies have reported that birds infected by parasites are paler than those uninfected, but, to our knowledge, none of them has assessed the possible effect of multiple infections by blood parasites on plumage colour. By comparing the yellow colour in the breast plumage of blue tits, Cyanistes caeruleus, between birds infected by different numbers of blood parasite genera, we found that those birds infected by more than one genus were paler than those parasitized just by one. In addition, we examined the potential role of carotenoid-based plumage colour of blue tits as a long-term indicator of other parameters of health status, such as body condition and immunoglobulin and heat shock protein (HSP) levels. Our results indicate that more brightly coloured birds had lower HSP70 levels than paler birds, but we did not find any significant association between colour and body condition or immunoglobulin levels. In addition, we found a positive significant association between Haemoproteus density of infection and HSP60 levels. Overall, these results support the role of carotenoid-based colours as indicators of health status in blue tits and show detrimental effects of parasitism on this character.     

Effects of age,sex, and ENSO phase on foraging and flight performance in Nazca boobies

Age‐related changes in survival and reproduction are common in seabirds; however, the underlying causes remain elusive. A lack of experience for young individuals, and a decline in foraging performance for old birds, could underlie age‐related variation in reproduction because reproductive success is connected closely to provisioning offspring. For seabirds, flapping flight during foraging trips is physiologically costly; inexperience or senescent decline in performance of this demanding activity might cap delivery of food to the nest, providing a proximate explanation for poor breeding success in young and old age, respectively. We evaluated the hypothesis that young and old Nazca boobies (Sula granti), a Galápagos seabird, demonstrate deficits in foraging outcomes and flight performance. We tagged incubating male and female adults across the life span with both accelerometer and GPS loggers during the incubation periods of two breeding seasons (years), during the 2015 El Niño and the following weak La Niña. We tested the ability of age, sex, and environment to explain variation in foraging outcomes (e.g., mass gained) and flight variables (e.g., wingbeat frequency). Consistent with senescence, old birds gained less mass while foraging than middle‐aged individuals, a marginal effect, and achieved a slower airspeed late in a foraging trip. Contrary to expectations, young birds showed no deficit in foraging outcomes or flight performance, except for airspeed (contingent on environment). Young birds flew slower than middle‐aged birds in 2015, but faster than middle‐aged birds in 2016. Wingbeat frequency, flap–glide ratio, and body displacement (approximating wingbeat strength) failed to predict airspeed and were unaffected by age. Sex influenced nearly all aspects of performance. Environment affected flight performance and foraging outcomes. Boobies'' foraging outcomes were better during the extreme 2015 El Niño than during the 2016 weak La Niña, a surprising result given the negative effects tropical seabirds often experience during extreme El Niños.     

Effect of Campylobacter-Specific Maternal Antibodies on Campylobacter jejuni Colonization in Young Chickens

Using laboratory challenge experiments, we examined whether Campylobacter-specific maternal antibody (MAB) plays a protective role in young chickens, which are usually free of Campylobacter under natural production conditions. Kinetics of C. jejuni colonization were compared by infecting 3-day-old broiler chicks, which were naturally positive for Campylobacter-specific MAB, and 21-day-old broilers, which were negative for Campylobacter-specific MAB. The onset of colonization occurred much sooner in birds challenged at the age of 21 days than it did in the birds inoculated at 3 days of age, which suggested a possible involvement of specific MAB in the delay of colonization. To further examine this possibility, specific-pathogen-free layer chickens were raised under laboratory conditions with or without Campylobacter infection, and their 3-day-old progenies with (MAB⁺) or without (MAB⁻) Campylobacter-specific MAB were orally challenged with C. jejuni. Significant decreases in the percentage of colonized chickens were observed in the MAB⁺ group during the first week compared with the MAB⁻ group. These results indicate that Campylobacter-specific MAB plays a partial role in protecting young chickens against colonization by C. jejuni. Presence of MAB in young chickens did not seem to affect the development of systemic immune response following infection with C. jejuni. However, active immune responses to Campylobacter occurred earlier and more strongly in birds infected at 21 days of age than those infected at 3 days of age. Clearance of Campylobacter infection was also observed in chickens infected at 21 days of age. Taken together, these findings (i) indicate that anti-Campylobacter MAB contributes to the lack of Campylobacter infection in young broiler chickens in natural environments and (ii) provide further evidence supporting the feasibility of development of immunization-based approaches for control of Campylobacter infection in poultry.     

Extensive hybridization reveals multiple coloration genes underlying a complex plumage phenotype

Coloration is an important target of both natural and sexual selection. Discovering the genetic basis of colour differences can help us to understand how this visually striking phenotype evolves. Hybridizing taxa with both clear colour differences and shallow genomic divergences are unusually tractable for associating coloration phenotypes with their causal genotypes. Here, we leverage the extensive admixture between two common North American woodpeckers—yellow-shafted and red-shafted flickers—to identify the genomic bases of six distinct plumage patches involving both melanin and carotenoid pigments. Comparisons between flickers across approximately 7.25 million genome-wide SNPs show that these two forms differ at only a small proportion of the genome (mean F_ST = 0.008). Within the few highly differentiated genomic regions, we identify 368 SNPs significantly associated with four of the six plumage patches. These SNPs are linked to multiple genes known to be involved in melanin and carotenoid pigmentation. For example, a gene (CYP2J19) known to cause yellow to red colour transitions in other birds is strongly associated with the yellow versus red differences in the wing and tail feathers of these flickers. Additionally, our analyses suggest novel links between known melanin genes and carotenoid coloration. Our finding of patch-specific control of plumage coloration adds to the growing body of literature suggesting colour diversity in animals could be created through selection acting on novel combinations of coloration genes.     

Factors Affecting the Duration of Nestling Period and Fledging Order in Tengmalm’s Owl (Aegolius funereus): Effect of Wing Length and Hatching Sequence

In altricial birds, the nestling period is an important part of the breeding phase because the juveniles may spend quite a long time in the nest, with associated high energy costs for the parents. The length of the nestling period can be variable and its duration may be influenced by both biotic and abiotic factors; however, studies of this have mostly been undertaken on passerine birds. We studied individual duration of nestling period of 98 Tengmalm’s owl chicks (Aegolius funereus) at 27 nests during five breeding seasons using a camera and chip system and radio-telemetry. We found the nestlings stayed in the nest box for 27 – 38 days from hatching (mean ± SD, 32.4 ± 2.2 days). The individual duration of nestling period was negatively related to wing length, but no formally significant effect was found for body weight, sex, prey availability and/or weather conditions. The fledging sequence of individual nestlings was primarily related to hatching order; no relationship with wing length and/or other factors was found in this case. We suggest the length of wing is the most important measure of body condition and individual quality in Tengmalm’s owl young determining the duration of the nestling period. Other differences from passerines (e.g., the lack of effect of weather or prey availability on nestling period) are considered likely to be due to different life-history traits, in particular different food habits and nesting sites and greater risk of nest predation among passerines.     

Delayed plumage maturation increases overwinter survival in North Island robins

Many bird species show delayed plumage maturation (DPM), retaining sub-adult plumage until after their first breeding season. Most explanations assume that DPM increases fitness over the breeding season. However, unless birds undergo a full moult before breeding, DPM could also be an adaptation to increase survival over the previous winter. The winter adaptation hypothesis has never been tested owing to the difficulty of measuring overwinter survival. We experimentally tested this hypothesis in North Island robins (Petroica longipes) using a closed island population where we could accurately estimate survival. The experiment involved dyeing 41 juveniles to mimic adult males, and comparing their survival with 41 control juveniles treated with the same peroxide base minus the pigment. The population was monitored with a series of resighting surveys, and mark-recapture analysis used to estimate overwinter survival. Survival probability was estimated to be 10% for dyed birds versus 61% for control birds in 2001, and 29% for dyed birds versus 40% for control birds in the winter of 2002, supporting the winter adaptation hypothesis for DPM. Access to suitable habitat is the key factor limiting juvenile survival in this population, and the locations where dyed juveniles were sighted suggest that they were often excluded from suitable areas.     

A bee’s eye view of remarkable floral colour patterns in the south-west Australian biodiversity hotspot revealed by false colour photography

Background and AimsColour pattern is a key cue of bee attraction selectively driving the appeal of pollinators. It comprises the main colour of the flower with extra fine patterns, indicating a reward focal point such as nectar, nectaries, pollen, stamens and floral guides. Such advertising of floral traits guides visitation by the insects, ensuring precision in pollen gathering and deposition. The study, focused in the Southwest Australian Floristic Region, aimed to spot bee colour patterns that are usual and unusual, missing, accomplished by mimicry of pollen and anthers, and overlapping between mimic-model species in floral mimicry cases.MethodsFloral colour patterns were examined by false colour photography in 55 flower species of multiple highly diverse natural plant communities in south-west Australia. False colour photography is a method to transform a UV photograph and a colour photograph into a false colour photograph based on the trichromatic vision of bees. This method is particularly effective for rapid screening of large numbers of flowers for the presence of fine-scale bee-sensitive structures and surface roughness that are not detectable using standard spectrophotometry.Key ResultsBee- and bird-pollinated flowers showed the expected but also some remarkable and unusual previously undetected floral colour pattern syndromes. Typical colour patterns include cases of pollen and flower mimicry and UV-absorbing targets. Among the atypical floral colour patterns are unusual white and UV-reflecting flowers of bee-pollinated plants, bicoloured floral guides, consistently occurring in Fabaceae spp., and flowers displaying a selective attractiveness to birds only. In the orchid genera (Diuris and Thelymitra) that employ floral mimicry of model species, we revealed a surprising mimicry phenomenon of anthers mimicked in turn by model species.ConclusionThe study demonstrates the applicability of ‘bee view’ colour imaging for deciphering pollinator cues in a biodiverse flora with potential to be applied to other eco regions. The technique provides an exciting opportunity for indexing floral traits on a biome scale to establish pollination drivers of ecological and evolutionary relevance.