The effect of food quality on breeding strategy in Griffon vultures (Gyps spp.)

Griffon vultures feed from the soft tissues of large mammal carcasses: a diet which contains very little calcium. Comparatively large amounts of calcium are required in order to breed, first for eggshell formation and then for bone mineralization of the chick. These requirements are estimated. Adults were not found to supply the chick with a mineral-rich crop secretion during rearing, but they must feed the young with calcium-rich food items, such as bone fragments. These are not abundant, and it is suggested that the mineral requirements for breeding have determined certain features of the breeding strategy in these birds—such as laying a single egg and having a prolonged rearing period.     

Modelling the energy budget of a colonial bird of prey, the Ruppell's griffon vulture, and consequences for its breeding ecology

Colonial nesting is rare in birds of prey. In this study we develop further Pennycuick's (1979 ) model of energy balance to consider the implications of colonial nesting for the breeding ecology of Ruppell's griffon vultures. To achieve a realistic foraging range, and remain in energy balance, the birds need to do more than fill their crop once on each foraging trip. They must remain in the feeding area and digest some of this food and refill the crop to obtain sufficient energy to pay for the flight costs and have sufficient energy to satisfy their own requirements and that of the chick. Given the known distances that the birds have to travel to forage, it would be impossible for them to rear more than one chick. The low growth rate of griffon vulture chicks may be an adaptation to the low rate at which energy can be delivered by the parents. The optimal time for a bird to be away from the nest changes with the distance they have to travel. Assuming that one parent remains on the nest at all times to guard the chick, it is optimal for both parents to take turns to forage on the same day if the distance to a feeding area is under 150 km, but to switch to each parent being away for a whole day when the distance is greater than this. Soaring flight is essential for such a scavenger, because of the low energy expenditure. If a vulture relied on the more energetically demanding flapping flight its maximum foraging range would be under 40 km. Griffon vultures are known to be able to depress their basal metabolic rate, and this has major implications for their foraging range, which then becomes constrained by the flight speed rather than by the amount of food they need to obtain. Griffon vultures minimize energy expenditure on all activities, because even small increases in their energy demands have a large impact on the foraging range that the bird can use.     

A change in the breeding season of Rüppell's Griffon Vultures Gyps rueppellii in the Serengeti in response to changes in ungulate populations

Breeding by Rüppell's Griffon Vultures Gyps rueppellii is synchronized but the time of breeding of colonies in the Serengeti region of Tanzania has changed by 5 months over the period between 1969–1970 and 1985. During this interval there have been considerable changes in the ungulate populations within the feeding range of the vultures, resulting in an increase to the food supply available to the birds. It is suggested that this species may have two alternative breeding seasons, the selective value of each depending on the magnitude of the food supply.     

FACTORS AFFECTING EGG-WEIGHT, BODY-WEIGHT AND MOULT OF THE WOODPIGEON COLUMBA PALUMBUS

The heaviest clutches (2 eggs) laid by Woodpigeons Columba palumbus in a Cambridgeshire study area weighed 30% more than the lightest. Yet the variation in egg-weight within clutches was less than 1 %. Irrespective of initial weight, eggs lost weight at the same constant rate during incubation. Heavy eggs hatched more successfully than light eggs and none weighing less than 16 g hatched. There was no correlation between chicks' weight at hatching and their weight at day 6 during the July-September part of the breeding season. The ability to feed crop milk at this stage could compensate for low chick-weight, but this might not be true early in the season. Weight at day 6 was correlated with the weight at day 16 or 17. The growth pattern is discussed. Chicks in broods of one achieved a higher weight at day 17 than those in broods of two. The survival rate both in and after leaving the nest was the same in both brood-sizes. Chick-weight in artificially created broods of three was almost as high as in broods of two, but again data refer to the July-September period when abundant cereal food is available. Survival before and after fledging was lower in broods of three. Clutch- and egg-weight declined from April until September. It is suggested that this is adaptive, in that the adults produce heavier eggs when food supplies are most difficult to collect. The critical period probably occurs during the few days when the adult must produce crop milk and the young cannot be left unattended. Thus egg-weight depends on the female's capacity to acquire nutrients, and is related to the needs of embryonic development and the amount of compensation in nutrient supply which can be provided immediately after hatching. But clutch-size is more related to the bird's ability to feed and rear young to the point of fledging, thereby influencing the number of offspring which survive to leave progeny. Egg-weight and female body-weight were positively correlated in females weighing less than 480 g but not in heavier females. First-year birds did not acquire adult weight until midsummer and they would probably produce light eggs if they could breed before this month. However, their gonads do not recrudesce until July and this prevents them breeding in the spring. Seasonal changes in body-weight and fat content of adults and first-year birds are described and discussed; differences were noted between adult males and females which were considered to be adaptive. The moult is described. It begins in April and continues until November, approximately one pair of primaries being replaced per month. The moult ceases during the winter months, when it is known that food supplies become limiting. Woodpigeons lay light eggs relative to their body-weight but can achieve the extra parental care needed for the altricial chicks by producing crop milk. Because the moult is extended, the energy demands of moulting and breeding combined are relatively low and this enables the Woodpigeon to have a long breeding season and to moult coincidentally.     

The stress hormone corticosterone in a marine top predator reflects short‐term changes in food availability

In many seabird studies, single annual proxies of prey abundance have been used to explain variability in breeding performance, but much more important is probably the timing of prey availability relative to the breeding season when energy demand is at a maximum. Until now, intraseasonal variation in prey availability has been difficult to quantify in seabirds. Using a state‐of‐the‐art ocean drift model of larval cod Gadus morhua, an important constituent of the diet of common guillemots Uria aalge in the southwestern Barents Sea, we were able to show clear, short‐term correlations between food availability and measurements of the stress hormone corticosterone (CORT) in parental guillemots over a 3‐year period (2009–2011). The model allowed the extraction of abundance and size of cod larvae with very high spatial (4 km) and temporal resolutions (1 day) and showed that cod larvae from adjacent northern spawning grounds in Norway were always available near the guillemot breeding colony while those from more distant southerly spawning grounds were less frequent, but larger. The latter arrived in waves whose magnitude and timing, and thus overlap with the guillemot breeding season, varied between years. CORT levels in adult guillemots were lower in birds caught after a week with high frequencies of southern cod larvae. This pattern was restricted to the two years (2009 and 2010) in which southern larvae arrived before the end of the guillemot breeding season. Any such pattern was masked in 2011 by already exceptionally high numbers of cod larvae in the region throughout chick‐rearing period. The findings suggest that CORT levels in breeding birds increase when the arrival of southern sizable larvae does not match the period of peak energy requirements during breeding.     

The cost of reproduction in the glaucous-winged gull

Summary Experimental enlargement of brood size in the glaucous-winged gull (Larus glaucescens) resulted in increased adult foraging time, decreased adult body weight at the end of the breeding season, and decreased over-winter adult survival. The decreased survival of breeding adults was associated with reduced body condition at the end of breeding (resulting from physiological costs of reproduction). Decreased survival was not due to an increased risk of injury or predation during the breeding season. Brood size did not directly affect the fecundity of surviving birds in the subsequent year. However, brood size may have an indirect effect on subsequent fecundity because the probability of mate loss increased among birds with large broods and the reproductive performance of birds with new mates was reduced. Based on estimates of life-time fitness calculated from fecundity and survivorship, birds with two- or three-chick broods (the normal brood size) have higher fitness than birds with one- or four-chick broods. However, the decreased fitness of birds with four-chick broods was slight, and probably not a sufficient explanation for the absence of natural four-chick broods in the glaucouswinged gull.     

Seasonal sex–specific energy expenditure in breeding and non-breeding Palestine sunbirds Nectarinia osea

The timing of the chick‐rearing phase is known to have a profound effect on the reproductive success of birds. However, little is known about the energetic costs faced by the parents during different periods of the breeding season. These costs may have vital consequences for both their survival and future reproduction. In most studies, daily energy expenditure (DEE) of breeding and non‐breeding birds has been compared, without controlling for the effect of season. In the present study, we examined the energy demands of breeding compared to non‐breeding Palestine sunbirds Nectarinia osea and whether there were sex‐specific differences in DEE within and between different seasons. We predicted that DEE would be elevated when birds rear chicks, especially at cooler ambient temperatures. Time‐energy budgets were constructed for pairs of sunbirds, rearing chicks, or not breeding, in spring and summer. There were significant seasonal differences in estimates of DEE in non‐breeders that were 21% higher in spring than in summer. We attributed these to increases in non‐flight metabolic rate rather than changes in time spent on different activities. Our estimates of DEE for the birds that were rearing chicks were higher than non‐breeding adults. In females the increase in DEE when breeding, compared to when not breeding, was similar in both spring and summer, while males increased their DEE much less when breeding in spring. The differences in estimated DEE, however, were not significant between male and female birds in any season. Between seasons, female breeders had 17.1% higher DEE in spring than in summer, while male breeders showed no difference in DEE when rearing chicks in different seasons. Accordingly, our initial prediction was supported, as DEE in chick‐rearing adults was higher than in non‐breeding adults. In addition, although temperatures are lower in spring, breeding in the spring is only more costly than breeding in summer for females. Apparently, males are more flexible in reallocating their time and energy spent on different activities.     

Nonbreeder birds at colonies display qualitatively similar seasonal mass change patterns as breeders

The difficulty in studying nonbreeding birds means that little is known about them or their resource requirements, despite forming a large and significant component of a population. One way to assess food requirements is to examine changes in body mass, because it indicates the amount of food acquired. In terms of body mass changes, our expectation is that nonbreeders will either (a) be in poorer condition than the breeders which potentially explains why they do not breed or (b) remain at a stable higher mass as they are unconstrained by the physiological costs associated with rearing chicks. Here, we interrogate body mass datasets of breeding and nonbreeding birds of two penguin species to assess these predictions and determine whether differences in mass exist between these two groups throughout the breeding season. The first dataset is from a wild Adélie penguin population, where bird mass was recorded automatically and breeding status determined from a resighting program. A second population of captive gentoo penguins were weighed regularly each breeding season. We demonstrate that although there were times in each year when breeders were heavier than their nonbreeding counterparts for both populations, the mass changes showed qualitatively similar patterns throughout the breeding season irrespective of breeding status. Heavier breeders at times during the breeding season are not unexpected but the overall similar pattern of mass change irrespective of breeding status is in contrast to expectations. It appears that breeding status per se and the constraints that breeding places on birds are not the only driver of changes in mass throughout the breeding season and, although not explicitly studied here, the role of hormones in driving changes in appetite could be key to explain these results. These results present a significant step toward understanding food requirements of nonbreeders in avian populations.     

Common Guillemot Uria aalge parents adjust provisioning rates to compensate for low food quality

The quantity and quality of food available within the foraging area set important constraints for chick‐rearing birds, but responses to low quality are not well understood. This study explored the potential for parent birds to adjust quantity (feeding rate) and quality (energy content) in chick provisioning, by studying Common Guillemots Uria aalge on Stora Karlsö, Baltic Sea, predominantly utilizing Sprat Sprattus sprattus, during conditions of high food quantity but reduced food quality. Quality is central to reproductive success in this single‐prey loader. From the chick's perspective, provisioning rates should be increased to compensate for low food quality and to fulfil its growing needs with increasing age. However, the high energy cost of flying in Guillemots makes it important for parent birds to minimize commutes to feeding areas. Provisioning parameters were recorded during three dawn‐to‐dusk watches each breeding season from 2005 to 2013, when clupeids, presumably Sprat, constituted 98% of chick diet. Generalized additive mixed models showed that both feeding rate and size of clupeids (a proxy for energy content) varied between years and changed non‐linearly with chick age, but that there was no change within breeding seasons. Chick age and year explained 36% of the variation in feeding rate but only 2% of the variation in the size of clupeids in chick diets. We conclude that parent birds tried to adjust both feeding rate and prey size, but were less successful with the latter. A strong negative correlation was found between annual feeding rates and size of clupeids, evaluated as the differences relative to the baseline year, and adjusted for the effects of chick age. Although the differences between years were small, the relationship indicates a compensation mechanism that does not seem to impact adult survival, and by which increased feeding rates can partly counteract reduced chick energy intake when food quality is low.     

Hematozoa from Southern African vultures, with a description of Haemoproteus janovyi sp. n.

This study represents the first major survey of avian hematozoa from southern Africa and the only one dealing with blood parasites of vultures. Blood smears from 506 Rhodesian, Botswanan and South African vultures (Hooded, White-headed, Lappetfaced, Cape Griffon and Whitebacked Vultures) were examined for hematozoa. Haemoproteus janovyi sp. n. was observed in 35.2% of the vultures, Leucocytozoon toddi in 0.8%, Plasmodium fallax in 0.6%, Atoxoplasma sp. in 1.4% and microfilariae in 0.2%. Hematozoan prevalence increased with age of the vultures. Only 2 of 133 nestlings sampled during the dry season had patent parasitemias (L. toddi). Haemoproteid prevalence in immature vultures was depressed during the dry season, whereas it was stable throughout the year in adults. The only species which nests on cliffs (the Cape Griffon Vulture) did not harbor hematozoa whereas the other species which nest and roost in trees were infected with at least one hematozoan species.     

Who Are We Feeding? Asymmetric Individual Use of Surplus Food Resources in an Insular Population of the Endangered Egyptian Vulture Neophron percnopterus

Supplementary feeding stations, or “vulture restaurants”, are common conservation management tools. While a number of studies have investigated the consequences of surplus food on the population dynamics of scavengers, relatively little is known about the effects of such practices at the individual level. Within the long-term monitored breeding population of Canarian Egyptian vultures (Neophron percnopterus majorensis) we investigated individual bird’s patterns of use of a supplementary feeding station at Fuerteventura (Canary Islands), over the course of breeding (2001, 2002; 2004-2011) and non-breeding seasons (2000-2010). Our results show that during the breeding season the individual use of the supplementary feeding station was inversely related to the distance to the breeding territory, which suggests the existence of central-place foraging constraints. In addition, larger birds of poor body-condition and individuals that ultimately failed to fledge young were detected more frequently at the feeding station. During the non-breeding season, and because most breeding birds abandoned the breeding territories, the overall abundance of Egyptian vultures at the feeding station grew. Moreover, the only variable increasing the probability of presence of individuals was poor body condition so that birds with lower wing residual visited the feeding station more frequently. Supplementary feeding may benefit individuals who would otherwise have been subject to selective pressures. From our results it follows that this conservation strategy must be used with caution because it can have consequences on an individual level and thus potentially affect the viability of endangered populations.     

A theoretical investigation of the effect of latitude on avian life histories

Tropical birds lay smaller clutches than birds breeding in temperate regions and care for their young for longer. We develop a model in which birds choose when and how often to breed and their clutch size, depending on their foraging ability and the food availability. The food supply is density dependent. Seasonal environments necessarily have a high food peak in summer; in winter, food levels drop below those characteristic of constant environments. A bird that cannot balance its energy needs during a week dies of starvation. If adult predation is negligible, birds in low seasonal environments are constrained by low food during breeding seasons, whereas birds in high seasonal environments die during the winter. Low food seasonality selects for small clutch sizes, long parental care times, greater age at first breeding, and high juvenile survival. The inclusion of adult predation has no major effect on any life-history variables. However, increased nest predation reduces clutch size. The same trends with seasonality are also found in a version of the model that includes a condition variable. Our results show that seasonal changes in food supply are sufficient to explain the observed trends in clutch size, care times, and age at first breeding.     

SURVIVAL OF MANX SHEARWATERS PUFFINUS PUFFINUS

The breeding success of Manx Shearwaters at Skokholm Island, Pembrokeshire, was followed in study burrows, and data on survival of the young were augmented by ringing large numbers as they were about to leave the island and recapturing them in later years.
The weight of the young at fledging and the date at which they leave affect their chances of survival; overall, as many as 30% of the young birds may survive to reach breeding age, which is thought to be normally about 5–6 years. Adult mortality varies between wide limits of about 5 and 20%, but juvenile survival appears to be of about the right order to balance adult losses.
The significance of the one-egg clutch is discussed. At the normal time of laying the female may not be able to obtain sufficient food to form a second egg. A two-egg clutch, laid at a later date, would not produce more surviving young since these would fledge too late in the season to have a good chance of survival.
The advantage of leaving the island at a heavy weight is discussed, and it is concluded that the fat stores laid down give heavy young a greater chance than light young of migrating a long distance without food. It is possible that the young may even reach their winter quarters (off Brazil) without having to feed on passage if they depart with enough stored fat.     

Population dynamics of the bearded vulture Gypaetus barbatus in southern Africa

The essential features of mortality and survivorship of bearded vultures Gypaetus barbatus in southern Africa were deduced from age class plumage characteristics. The population consisted of about 204 adult pairs within a breeding range of about 35,000 sq km. Pairs bred every year and produced, on average, about 0·9 young per pair per year. Young birds made up about 37% of the population, subadults 3·5% and adults 60%. About 182 fledged young were recruited to the population each year. The proportion of young birds in the population in different areas was inversely related to the breeding density of adult birds (range 24–47%). Young bird mortality over the four years to subadult age was 87%, the survival rate of adults was 94% and the mean lifespan of birds surviving to adulthood was 21·4 years. This study demonstrates the need to understand the relationship between adult breeding density and young bird numbers in different parts of their range to accurately deduce population dynamics characteristics.     

Recruitment to food in black vultures: evidence for following from communal roosts

Experimental and observational evidence from a 5-year study of a partially marked population of black vultures, Coragyps atratus, suggests that individuals can find food by following others from overnight roosting groups. Natural food sources usually lasted long enough for recruitment to occur. Black vultures formed larger roosting groups and had enhanced second-day recruitment during winter, when their main food source was least abundant. Successful foragers returned to experimental food sources on subsequent days and generally preceded new arrivals. Arrivals at bait were clumped on days of discovery and second days, but incoming groups were larger on second days and highly skewed towards early arrivals. In general, adults arrived at baited sites earlier than young adults and juveniles. During natural roost departures, adults were as likely to depart at the head of departing groups as at the rear, while young were concentrated in the rear. Birds removed experimentally from the population long enough to be naive about the location of food followed others from the roost when reintroduced. Control adults (birds caught and handled but not restrained for long) were not concentrated in the rear of roost-departing groups, as were experimental adults. Juveniles tended to follow regardless of recent experience. These results suggest that roost attendance facilitates foraging for black vultures; these roosts probably function as information centres.     

Movements of radio-tagged Grey Herons Ardea cinerea during the breeding season in a large pond area

This paper describes the strategies of resource utilization in the course of the breeding season by five radio-tagged Grey Herons Ardea cinerea. The seasonal changes in exploitation of the environment by two breeding adults, one non-breeding adult and two non-breeding first-year birds were studied from March to August 1982, near Zonhoven in Belgium. Two adult breeding birds could be followed continuously from the end of March until the middle of June. During the first month they explored an extended area all around the colony, but each concentrated its search in a specific direction. From the end of April until the beginning of June, most probably from egg-hatching until the end of breeding activities, each bird spent a very large proportion of its time at a particular feeding site, from which other herons were actively excluded. In the first part of June they again visited different sites, each maintaining its preferred direction. From the middle of June onwards they seemed to have left the fish-pond area. The pattern of movements of the first-year birds differed markedly from that of the breeding adults. In April, although both non-breeding and breeding birds explored large areas, only the areas used by non-breeders were centred on the colony. From the end of April onwards, probably after general egg-hatching in the colony, the non-breeders very rarely revisited the colony, and from May till August their ranges became more and more restricted to very small areas at an increasing distance from the colony. They were never observed defending particular sites. The results are discussed with regard to recent speculations about the evolution of colonies as an adaptation for the exploitation of food resources. Breeding herons seem to explore a large part of the environment during incubation and defend a particular site while feeding their young. Choice of feeding site by non-breeding birds may be influenced by the site defence of the breeding birds. Non-breeding birds exploit a large area when breeding birds occupy feeding territories. Perhaps they are forced to forage in less suitable places at this time. Colonies might have evolved as a strategy to minimize effort in resource esploitation as, especially at the beginning of the breeding season, the colony could act as an assembly point in the exploration of the environment. However, its importance as an assembly point diminishes in the course of the season, as non-breeding birds no longer visit the colony and adults defend territories.     

BODY WEIGHTS IN LAYSAN ALBATROSSES DIOMEDEA IMMUTABILIS

Male Laysan Albatrosses are heavier (3300 gm.) than females (3000 gm.) upon arrival on Midway. This difference between the sexes persists except at egg-laying and during certain periods of incubation. Weights of males and females are essentially the same at egg-laying, perhaps because of the presence of the 300 gm. egg in the female, but mostly because the males have lost weight in the pre-egg phase. Losses in weight during the long, continuous incubation spans are about 18–25% of the body weight. Males show a net loss of about 10% during the entire incubation period; there is no statistically significant change in the weights of the females. Weights of both sexes decline in parallel from the end of incubation to mid-May. Thereafter, they increase as attentiveness to the young is less and the parents have more time to forage at sea. Nevertheless, the body weights of both sexes at the end of the breeding season (July) are down 10–45%. Decreased body weights of breeding birds cannot be attributed solely to the rigours of parental care; former breeders and juveniles not yet of breeding age show a similar decline in weight. It is suggested that the normal pattern of yearly breeding by adult Laysan Albatrosses may be broken by failure to add weight (fat) before the breeding season. Former breeders consistently weigh 15–20% less than current breeders. Nestlings normally show a gradual increase in body weight from hatching in late January until mid-May, at which time they may be significantly heavier than either parent. As a result of the increased energy requirements of locomotion and wing exercise and decreased food supplied by parents, nestlings lose about 30% of the May weight before they finally go to sea in Jufy A possible “mutation” for small body weight is discussed.     

The use of (GACA)4 PCR to sex Old World vultures (Aves: Accipitridae)

A PCR method was developed employing a single primer (GACA)₄ to sex Black vultures ( Aegypius monachus ), Lappet-faced vultures ( Torgos tracheliotus ), and Griffon vultures ( Gyps fulvus ). Using the (GACA)₄ primer several PCR products were generated. One or more PCR products displayed a sex-specific pattern, i.e. they were only present in females (probably corresponding to repetitive DNA on the W chromosome) but absent in males. The sex ratio of 85 Griffon vultures from Spain was almost 1.     

Body weight, gonad development and moult in the Tawny owl (Strix alum)

Gonad development, moult and seasonal changes in body weight and composition in the Tawny owl Strix aluco were studied by examining the carcasses of 369 owls (mostly road casualties) supplemented by 112 weights of live birds. In breeding females laying was preceded by the accumulation of fat and to a lesser extent protein which meant that they weighed more at this time (February/March) than at any other. Females declined in weight after laying but were still heavy during incubation. In contrast, males and non-breeding females did not increase in weight before the start of the breeding season. Juveniles reached or even exceeded adult weight well before independence due to the deposition of fat. Even after the exclusion of diseased or contaminated individuals, 9·4% of the birds examined were identified as starving; most of these were in the autumn and were probably newly-independent young wandering in search of territories. In both sexes gonad maturation was of brief duration coinciding with the period (mid-March to mid-April) in which eggs are normally laid. Ovarian growth was biphasic. In the three months prior to the breeding season ovarian condition in different birds was positively correlated with body weight and it appeared that the largest ovarian follicles of females in poor condition failed to attain the size from which rapid growth to final ovulation occurs. in males testis size in the breeding season was correlated with pectoral muscle weight (an index to protein condition) but not body weight. The majority of adults commenced wing moult in June. The average duration of primary moult was estimated to be 77 days. Healthy birds replaced the primaries of both wings at the same rate but most diseased birds moulted asymmetrically and/or out of season. First-year birds renewed their body feathers between September and November. In the Tawny owl territory establishment, breeding and moult are temporally separated.     

Poison-related mortality effects in the endangered Egyptian vulture (<Emphasis Type="Italic">Neophron percnopterus</Emphasis>) population in Spain

A total of 211 poisoning incidents registered over the period 1990–2007 and affecting 294 Egyptian vultures (Neophron percnopterus) were studied to address the impact of poison-related mortality in the Spanish population. Poison-related mortality mainly affected the birds on an individual level, with low numbers of individuals being found in each incident (mean 1.39) with 94.9% being adults. Deaths were largely recorded (81.8%) during the breeding season, with mortality peaking during May and June (52.1%). In contrast with other raptor species, a high proportion of adult individuals (74.2%) were found in the nest or its surroundings. Age-related differences in the poisoning rate are probably related with different feeding and behavioral strategies between age classes. The illegal use of poison to control predators was the main cause of mortality (93.8%), and particularly in small hunting reserves (74.9%), since the kind of food resources that adults exploit are coincident with the type of baits employed to illegally control predators and the preferred habitat coincides with areas of small game hunting. Our results suggest that poisoning is probably one of the main causes of Egyptian vulture mortality in Spain. The eradication of the illegal use of poisoning and supplementary feeding in specific territories to provide safe food seems priority for its conservation.