Food-niche differentiation in sympatric Hen Circus cyaneus and Montagu's Harriers Circus pygargus

Hen Harriers Circus cyaneus and Montagu's Harriers Circus pygargus are medium-sized raptors that differ in size (Hen Harrier being slightly bigger than Montagu's Harrier) and breeding system (Montagu's Harriers are semi-colonial and Hen Harriers defend nesting–hunting territories). In contrast, the diets of the two species when in sympatry are very similar. We evaluated food-niche differentiation among these coexisting raptor species and how between-species differences in body size and social system influence interspecific relationships. We present data from a study conducted in 1997 and 1998 in northeastern Madrid province (central Spain). Diet of the two species largely overlapped (55–95%) during the breeding season, but Hen Harriers preyed more often on larger species. This segregation was observed both in the average size of the primary prey (lagomorphs) and in the alternative prey (birds for Hen Harriers vs. insects for Montagu's Harriers), and was particularly apparent late in the season. Accordingly, feeding frequency of Montagu's Harriers, but not of Hen Harriers, increased later in the season. Size differences between species in prey brought to the nest were apparent for both males and females. Foraging behaviour also differed, as Hen Harriers spent more time hunting close to the nest than did Montagu's Harriers. This implies that segregation in foraging areas may also exist. Observed niche partitioning may relax the potential for competition between these species.     

Feeding behaviour and diet composition of Marsh Harriers Circus aeruginosus in agricultural landscapes

Capsule Diet composition and feeding behaviour of breeding birds differed between areas of high and low agricultural intensification.

Aims To analyze diet composition and feeding behaviour of Marsh Harriers Circus aeruginosus in agricultural landscapes with different degrees of agricultural intensification.

Methods Diet composition and feeding behaviour was assessed by focal observations at nests during breeding seasons in 2001–03 in two agricultural areas within the eastern Ebro Basin (on the northern Iberian Peninsula) which differed in their degree of agricultural intensification: a traditional non-irrigated and non-intensively managed farmland area; and an irrigated and intensively managed farmland area.

Results Marsh Harriers in the intensively managed area took higher percentages of small mammals than harriers in the traditional area managed at low-intensity, where they also had a more diverse diet. Male delivery rates increased throughout the breeding season in the intensively managed area. On the contrary, no significant trend in delivery rates was observed in the traditional area, where harriers turned to larger prey late in the season.

Conclusion Marsh Harriers probably took advantage of different prey types in intensively managed and traditional farmland according to prey availability. Differences in diet composition seem to be related to different foraging strategies adopted by harriers in traditional and intensively managed farmlands. In the traditionally managed farmlands, harriers responded to increases in the energy requirements of the brood over the course of the breeding season by taking larger prey, whereas in the intensively managed farmlands harriers increased feeding rates.     

Diversionary feeding and nestling diet of Hen Harriers Circus cyaneus

Capsule: Diversionary feeding reduced Hen Harrier Circus cyaneus nestlings’ natural food intake by half. Red Grouse Lagopus lagopus scotica chicks constituted 0–4% of all nestling food items. Annually, this reduced annual grouse chick production by 0–6%.

Aim: To quantify proportions of diversionary and natural food (including grouse) delivered to Hen Harrier nestlings in relation to brood size, male status and natural prey abundance.

Methods: We recorded diversionary food provisioned to 25 Hen Harrier broods (2008–15) and studied the diet of 15 broods using observations from hides, nest cameras and regurgitated pellet analysis. Variation in nestling diet was analysed using compositional analysis.

Results: Hen Harriers took 76% of diversionary food provided. Depending on assessment method, average nestling diet was 44–53% diversionary food, 39–55% natural prey (including 24–45% passerines, 4–15% small mammals, 0–4% grouse chicks) and 0–9% unknown items. The amount of diversionary food consumed was not influenced by male status, brood size or natural prey abundance. The number of Red Grouse chicks delivered annually was 34–100% lower than expected under unfed conditions, however, the confidence intervals associated with these estimates were large.

Conclusion: Diversionary food influenced Hen Harrier nestling diet and reduced the number of Red Grouse chicks taken relative to modelled predictions. It may help reduce conflict between Hen Harrier conservation and Red Grouse shooting, but only if overall grouse productivity is thereby maintained or increased.     

Mismatches between breeding success and habitat preferences in Hen Harriers Circus cyaneus breeding in forested landscapes

During the past century, the upland breeding areas of Hen Harriers in Ireland have been extensively afforested. There is no evidence that this species avoids breeding in heavily forested landscapes and, indeed, young commercial forests in their second rotation are often selected as nest‐sites. However, Hen Harriers have coexisted with these forested areas for only a few decades and it is possible that such landscapes are suboptimal. We examined the relationship between breeding success and habitat using a dataset spanning three years and four study areas in the south and west of Ireland. We assessed whether nest success and fledged brood size were related to habitat type, both at the nest‐site and in the surrounding landscape. Neither measure of breeding productivity was related to total forest cover or to percentage cover of closed canopy forest in the landscape. However, in a subset of areas, high cover of second‐rotation pre‐thicket (young forests planted on land from which a first rotation has already been harvested) in the surrounding landscape was associated with low levels of breeding success. This may be due to factors related to predation, disturbance or prey availability. The fact that second‐rotation pre‐thicket is a preferred habitat for nesting in Ireland suggests that Hen Harriers may be making suboptimal decisions in the landscapes available to them.     

Habitat selection and foraging success of Hen Harriers Circus cyaneus in west Scotland

The foraging behaviour of Hen Harriers breeding in west Scotland was studied in a variety of afforested and moorland habitats. Time budget information from sample areas, stratified by habitat type and controlled for nest distance, was used to investigate habitat selection. Preferences were then related to measures of foraging performance in each habitat. Hen Harriers foraged preferentially over young first rotation coniferous forests, and selected heathland and grassland habitats ahead of closed canopy woodland. Harrier preference for the various habitats was positively correlated with the frequency with which they struck at, and captured, prey. Prey strike success was much lower than that reported for moorland managed for Red Grouse Lagopus lagopus scoticus. The results explain many of the reported changes in local Hen Harrier populations. The extent of young first rotation forestry is currently in decline. It is predicted that this will lead to a reduction in numbers of Hen Harriers breeding in west Scotland, and an increase in the proportion of the British harrier population occupying moorland managed for Red Grouse.     

Diet of Montagu's Harrier Circus pygargus in central Spain: analysis of temporal and geographic variation

The diet of Montagu's Harrier Circus pygargus was studied for 4 years near Madrid, Spain, with the use of pellets and prey remains collected during the breeding season. The diet of the study birds included a wide variety of prey categories, the most important one by weight being lagomorphs (mainly Hares Lepus granatensis ). The diet varied seasonally, with Hares predominating early in the breeding season, while birds increased in importance later. The shift was partly a result of variation in availability (with fledgling birds being more abundant late in the season), but statistical analysis suggested that lagomorphs and then birds were actively chosen in the prelaying and nestling periods, respectively. Montagu's Harrier seemed to be an opportunistic species, hunting a wider range of prey types in more southern latitudes, but specialized in each region in the prey species that was most profitable. This was the only study on Montagu's Harriers to date which found lagomorphs to be the most important part of the diet throughout the season.     

The direct and indirect effects of predation by Hen Harriers Circus cyaneus on trends in breeding birds on a Scottish grouse moor

Two phases of an experimental reduction in management at Langholm Moor in southwest Scotland provided an insight into factors that determined trends in the abundance of moorland birds. In 1992 the historical control of breeding Hen Harriers Circus cyaneus stopped and in 2000 grouse moor management was discontinued. Golden Plover Pluvialis apricaria , Lapwing Vanellus vanellus , Curlew Numenius arquata and Red Grouse Lagopus lagopus scoticus declined, whilst Carrion Crow Corvus corone and Snipe Gallinago gallinago increased. Hen Harriers increased from two to 20 breeding females, then declined back to two. Lapwing abundance was positively associated with that of Hen Harriers, and moorland passerines (Skylark Alauda arvensis , Meadow Pipit Anthus pratensis and Stonechat Saxicola torquata ) were negatively associated. Golden Plover, Lapwing, Curlew, Red Grouse, Skylark and Hen Harrier were more abundant when the moor was managed for grouse, whilst Carrion Crow, a common predator of clutches of ground-nesting birds, increased during the second half when management ceased. Increased Crow numbers, together with an increase in Red Foxes Vulpes vulpes , probably contributed to the observed bird declines.     

Global dispersion of nesting Hen Harriers Circus cyaneus; implications for grouse moors in the U.K.

In the U.K., a full recovery of Hen Harrier Circus cyaneus breeding numbers is prevented by illegal culling by some gamekeepers who fear the species threatens the future of grouse moors. This study's main purpose was to estimate how many more Hen Harriers there would be in the U.K. if this culling were to cease. A review of 33 studies of the Hen Harrier in nine countries showed that the densities of nesting females decreased strongly as the size of the areas studied increased and also that such information could not be used to estimate the potential density over a large area, such as the U.K. Correcting for the size of the areas studied showed that the potential density of nesting Hen Harriers on grouse moors could be predicted from studies overseas. Estimates of the densities of Hen Harriers in occupied areas globally were therefore used to estimate the density appropriate for the U.K. If all potential habitats were occupied, present numbers could more than double, to an estimated 1660 nesting females. This estimate represents an average of one nesting female per 2 5 km² of habitat, a density which would cause little or no significant economic damage on grouse moors. However, because Hen Harriers tend to aggregate, they would not spread out evenly but would nest in relatively high densities on a number of moors. The economic impact on Red Grouse Lagopus lagopus would not be a function of overall numbers, rather it would arise from the uneven dispersion of nesting Hen Harriers.     

Regional and temporal variation in diet and provisioning rates suggest weather limits prey availability for an endangered raptor

Understanding variation in food requirements of wild animals is of central importance in population ecology and conservation, as it helps to identify where and when food may be limiting. Studies on diet variation or prey provisioning rates may give useful insights when direct information on prey availability is lacking. We assess spatial and temporal variation in the diet of an endangered predator, the Black Harrier Circus maurus. This raptor is endemic to southern Africa and specializes on small mammals but also feeds on birds and reptiles as alternative prey. Using data on 1679 prey identified in 953 pellets collected in inland and coastal regions from 2006 to 2015, we show that diet composition changed little throughout the breeding season in the coastal region, whereas there was a marked seasonal decline in the occurrence of small mammal prey in the inland region, with a concomitant increase in alternative prey. The proportion of small mammals in the diet declined with increasing maximum temperature, the latter being highest at the inland region late in the breeding season. Using camera recordings at nests in 2014, we further analysed daily patterns of prey provisioning to nestlings. A marked reduction in small mammal provisioning rates occurred during the middle of the day in the hotter inland region but not in the cooler coastal region. Reduced availability of the primary prey, small mammals, in hotter conditions, through a reduction in activity or overall abundance, could explain these patterns. Finally, we show a positive relationship between winter rainfall and interannual differences in the proportion of small mammals in the diet of Black Harriers breeding in the coastal region, suggesting relationships between diet and prey abundance that are mediated through rainfall. We discuss the need to consider spatial variation in food availability in conservation strategies.     

DISPERSION, BREEDING AND PREY OF THE HEN HARRIER CIRCUS CYANEUS IN GLEN DYE, KINCARDINESHIRE

Hen Harriers were studied on a grouse moor in northeastern Scotland from 1970-74. Numbers in spring were fairly stable, and the 24 nesting territories occupied in 1974 were regularly dispersed with a mean nearest neighbour distance of 1.52 ± s. e. 0.09 km. Many pairs apparently failed to breed and left the area. The mean clutch size of those remaining was 4.70 ± s. e. 0.24 (n= 27) and the mean number of young fledged per successful nest was 3.11 ± s. e. 0.26 (n= 19). Young males were more likely to disperse far from the study area than young females. The variety of prey species observed from hides at five nests was very limited, and comprised mainly pipits, grouse chicks and lagomorphs. Estimates by weight suggest that grouse and lagomorphs accounted for 89% of all prey. Males brought more small items and fewer large items to nests than females, and on average provided 72% of all prey items seen during watches from the hides. On the basis of studies of prey at three nests in 1974, and counts of grouse in spring and late summer, harrier predation was estimated to have reduced the number of grouse which might otherwise have survived to late July that year by a maximum of 7.4%.     

Brood adoption and deceit among African Marsh Harriers Circus ranivorus

Complete adoption of unrelated broods by foster parents is surprisingly common among birds and the behaviour appears to be maladaptive in several species. In a small, marked, southern African population of African Marsh Harriers Circus ranivorus, studied over 4 years, a young male took over the complete parenting (provisioning and defence) of an unrelated brood, and several times fed the brood in preference to the soliciting maternal parent. This female then deserted and deceitfully courted (out of season) a third male, whose provisioned food she took to feed to her own young. Since the adopting male retained the territory and bred successfully for at least two further years, he suffered no expected loss in fitness and hence this case of fostering was neither maladaptive nor apparently altruistic.     

Food habits of genetically identified pine marten (<Emphasis Type="Italic">Martes martes</Emphasis>) expanding in agricultural lowlands (NW Italy)

We assessed the diet of pine marten (Martes martes) expanding in the heavily human-altered agricultural plain of the River Po, northern Italy. Between February 2008 and November 2009, surveys were carried out twice a week during seasonal sampling sessions of three to four consecutive weeks. To distinguish the faecal samples from those of sympatric carnivores, a polymerase chain reaction-restriction fragment length polymorphism method was applied. The availability of small mammals was assessed by the analysis of 59 barn owl (Tyto alba) pellets. A total of 109 pine marten faeces were analysed. Its diet consisted of fruit, rodents, lagomorphs and birds. Seasonal variation occurred for fruit, which prevailed in summer, and rodents, which were more preyed upon in autumn. In winter, the diet of the pine marten was almost totally based on vertebrates, and lagomorphs were the main source of protein in summer. Use of small mammals differed significantly from their availability, voles, particularly bank vole Myodes glareolus, being preferred to mice (Apodemus sp.) and rats (Rattus sp.). Medium-size mammals formed about 18% of pine marten diet, a value generally reported for high-latitude habitats. In agricultural areas, the pine marten proved to be an opportunistic predator, able to face the reduced availability of small mammals by preying upon medium-size prey and fruit.     

The importance of pre-thicket conifer plantations for nesting Hen Harriers Circus cyaneus in Ireland

Hen Harriers Circus cyaneus are threatened across much of their range and their conservation requires appropriate habitat management. The locations of 148 Hen Harrier nests found in the Republic of Ireland during national breeding surveys in 2000 and 2005 were used to assess nest-site selection. The distribution of these nests was compared to distributions of randomly located points to investigate selection at the scale of the nest-site and landscape. The main nesting habitats selected were pre-thicket stage of first and, particularly, second rotation plantations, mostly of exotic conifers. There was no evidence that the area of post-closure plantations negatively affected Hen Harrier nest distribution. There was a positive correlation across study areas between changes in numbers of Hen Harrier nests between 2000 and 2005 and changes in the area of pre-thicket second rotation plantations over the same period. The overall effect of plantation forests on breeding Hen Harriers in Ireland therefore appears to be positive. However, this study did not consider the effects of plantation habitats on breeding success. Improved grassland was strongly avoided as a nesting habitat. Furthermore, after controlling for the influence of nesting habitat on nest location, landscapes with a high percentage cover of improved grassland were also avoided. Further agricultural intensification of grassland in areas where Hen Harriers breed is likely to have a negative impact on this species. These results are required for the development of management strategies for the conservation of this species.     

A comparison of the feeding ecology of wintering Hen Harriers Circus cyaneus centred on two heathland areas in England

Small passerines were the principal prey in pellets from Hen Harrier Circus cyaneus communal winter roosts in Breckland and the New Forest. Skylark Alauda arvensis remains predominate in the pellets from Breckland. Dunnock Prunella modularis and Greenfinch Carduelis chloris were also important. The contents of New Forest pellets reflected the position of each roost. In pellets from a roost on the northern edge of New Forest heathland and close to farmland, Skylark was by far the predominant prey. Wren Troglodytes troglodytes and Linnet Carduelis cannabina were the next most important passerines. Pellets from the more extensive heather heathland in the south of the New Forest showed greater dependence on the heathland community of insectivorous passerines, with Wren, Dartford Warbler Sylvia undata and Meadow Pipit Anthus pratensis occurring as prey in that order of frequency. Pellets from a roost in northwest New Forest heathland with less heather and closer to farmland contained mainly a mixture of farmland-and heathland-feeding passerines. The directions of arrival and departure at roosts indicated that the Breckland-roosting harriers hunted mainly in arable fenland and the New Forest edge-roosting harriers in arable downland. Counts of the principal passerine prey species in roost catchment areas indicated that Greenfinch and Linnet were most abundant in arable fenland and arable downland squares, respectively, and the Wren was most abundant in the New Forest catchment area as a whole. The influence of the distribution of weed species on the species of seed-eating passerines in prey was investigated. Proportions of passerines in pellets from the Breckland and New Forest edge roosts were positively correlated with the proportions of “grey” males. Proportions of finches in both the Breckland and the New Forest edge-roost pellets declined sharply after January. There was a shift toward lagomorphs in the diet after January and a suggestion that this was positively correlated with the proportions of ringtails at the Breckland roost. Factors affecting prey composition and selection of foraging habitat, the effect of harrier predation on Dartford Warbler numbers and the implications for differential distribution of the harrier sexes in winter are discussed.     

Comparative breeding ecology and hybridization of Eastern and Western Marsh Harriers Circus spilonotus and C. aeruginosusin the Baikal region of Eastern Siberia

New information on the breeding ecology and inter-breeding of Western Circus aeruginosus and Eastern C. spilonotus Marsh Harriers in the Baikal region of Eastern Siberia is presented. Hybridization is relatively common (one of the parents was not 'pure' C. spilonotus in six out of 28 nesting cases) in the Irkutsk area, where nests of a mixed pair were found with chicks in 1999 and 2000. Eastern Marsh Harriers are more similar to Western in size, biology and coloration of young birds, but to the Australasian C. approximans in their displays and mating system, and intermediate between C. aeruginosus and C. approximans in the plumage coloration of females. The systematic status (species' or 'subspecies') of the Western and Eastern Marsh Harriers depends on the systematic approach taken, and existing data seem to be inadequate to resolve this. The Siberian Mongolian zone of contact of these marsh harriers is unique as other marsh harriers of the world are allopatric. More research on ecology, behaviour and molecular systematics is necessary to resolve the situation.     

Lead poisoning in wild populations of Marsh Harriers Circus aeruginosus in the Camargue and Charente-Maritime, France

Lead poisoning of waterfowl, through the ingestion of spent gunshot, has been recognized as a mortality factor for over a century. However, in Europe relatively little attention has been paid to raptors that may ingest shot embedded in the flesh of prey. The present study examines the incidence of lead poisoning in wild Marsh Harriers Circus aeruginosus trapped at two sites in France, the Camargue and Charente-Maritime.
Eight captive Marsh Harriers (controls) had blood lead (PbB) concentrations of 5.3–10.8 μg per dl. Of 94 wild birds trapped during the winters of 1990/1991 and 1991/1992 either in baited clap traps or in mist nets at night roosts, 29 (31%) had elevated (>30 μg/dl) PbB concentrations and 13 (14%) had concentrations indicative of clinical poisoning (> 60 μg/dl). Similar percentages of birds caught using the two trapping methods had elevated (>30 μg/dl) PbB concentrations. However, clap netting appeared to select for more heavily contaminated birds and for juvenile birds, although the two are not necessarily correlated. In the Camargue, significantly more females than males trapped in clap nets had elevated PbB concentrations.
The incidence of shot in regurgitated Marsh Harrier pellets increased significantly between October and December, indicating increased exposure to lead as the hunting season progressed. There appeared to be a parallel increase in PbB concentrations throughout the hunting season, although this could not be clearly demonstrated as sex ratios were different during different sampling periods.
Elevated PbB concentrations of harriers in the Camargue are likely to result primarily from the ingestion of shot in the flesh of crippled or unretrieved waterfowl and in Charente-Maritime, from eating crippled or dead mammals.
Other raptors at risk from lead poisoning and solutions to this problem are discussed.     

Diagnosing the environmental causes of the decline in Grey Partridge Perdix perdix survival in France

We studied Grey Partridge Perdix perdix mortality during breeding to identify the environmental causes of a long‐term decline in adult survival. We radiotagged and monitored daily from mid‐March to mid‐September 1009 females on ten contrasting study sites in 1995‐97. Simultaneously, we recorded habitat features and estimated the abundance of Hen and Marsh Harriers Circus cyaneus and C. aeruginosus Red Fox Vulpes vulpes and mustelids. We experimentally tested whether scavenging could have biased predation rates. We also examined, through the necropsy of 80 carcasses of Grey Partridge, whether disease, parasites or poisoning could have been ultimate causes of high predation rates. The survival rate of radiotagged females during spring and summer ranged from 0.25 to 0.65 across study areas. Mortality peaked in May, June and July when females were laying and incubating. The direct negative impact of farming practices was low (6%). Predation was the main proximate cause of female mortality during breeding (73%) and determined the survival rate, suggesting no compensation by other causes of mortality. Ground carnivores were responsible for 64% of predation cases, and raptors for 29%, but this proportion varied across study sites. Disease and poisoning did not appear to favour predation, and scavenging was not likely to have substantially overestimated predation rates. The predation rate on breeding females was positively correlated with the abundance of Hen and Marsh Harriers, suggesting an additional mortality in areas where harriers were abundant. The proportion of raptor predation was linearly related to harrier abundance. The predation rate was not correlated with the abundance of the Red Fox and mustelids. A potential density‐dependent effect on the predation rate was confounded by the abundance of harriers. We found no convincing relationship between the predation rate and habitat features, but we observed a positive relationship between the abundance of Hen and Marsh Harriers and the mean field size. This suggested that habitat characteristics may contribute to high predation rates through predator abundance or habitat‐dependent predation.     

Interspecific and intraspecific differences in habitat use and their conservation implications for Palaearctic harriers on Sahelian wintering grounds

The floodplains of the West‐African Sahel region have experienced extensive habitat transformation during the past four decades, coinciding with an impoverishment of raptor populations. We investigated foraging patterns of Palaearctic migratory Eurasian Marsh Harriers Circus aeruginosus, Pallid Harriers C. macrourus and Montagu’s Harriers C. pygargus on a floodplain system in northern Cameroon to assess species, sex‐ and age‐related habitat preferences. Sex and age have rarely been incorporated into general studies of raptor habitat associations, despite clear evidence of intrasexual and age‐related differences in foraging strategies and diet composition, potentially carrying strong conservation implications. We found evidence of sexual differences in foraging preference related to land use, particularly in the most sexually dimorphic Pallid Harrier, and evidence that juveniles used different habitats to adults. This constitutes the first quantitative documentation of such differentiation by Palaearctic raptors on African wintering grounds, indicating that general patterns of habitat use in wintering raptors may obscure sex‐ and age‐specific preferences. Contrary to expectations, we found limited evidence for interspecific foraging segregation. Food partitioning by prey mass was related to harrier body mass and facilitated by a diverse availability of prey on human‐transformed floodplains. Anticipated further large‐scale conversion of floodplain habitat into predominantly desiccated grasslands raises concerns about the survival of wintering harriers.     

Factors influencing diet variation in a generalist predator across its range distribution

Factors linked with intraspecific variation in trophic diversity are still poorly understood in generalist species like the Montagu’s harrier (Circus pygargus) but may have important implications for conservation management at a wide scale. We described geographic patterns of Montagu’s harrier diet across Eurasia, gathering diet data from 30 studies in 41 areas from 11 countries. We grouped prey as invertebrates, reptiles, small mammals, large mammals, eggs, small birds and large birds, and calculated the contribution of each prey type to the diet (as % biomass) and Shannon’s Diversity Index for each study site. We analysed qualitative estimates of prey abundance in relation to latitude and longitude, then diet composition in relation to habitat of the study area and prey abundance estimates. Diet diversity of Montagu’s harriers increased from north to south, while abundance of all prey groups other than small mammals showed the opposite trends. Agricultural areas in northern latitudes seemed to hold high densities of small mammals, but low densities of alternative prey. Overall, birds were the main prey in most of Montagu’s harrier’s distribution range, although the relative importance of each prey type in the diet was significantly explained by its local abundance and habitat, confirming the opportunistic foraging strategy of this raptor species. Consumption of mammals was an exception to this trend, being negatively associated with the abundance of alternative prey, suggesting that this prey is not preferred. Trophic diversity in this species could be influenced by land-use changes through variations in the abundance and availability of prey, which could impact its population dynamics. This may be particularly important for northern populations of Montagu’s harriers breeding in agricultural habitats, where trophic diversity is already low.     

Biogeographical patterns in the diet of an opportunistic predator: the red fox Vulpes vulpes in the Iberian Peninsula

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