Nocturnal flights by Bearded Vultures Gypaetus barbatus detected for the first-time using GPS and accelerometer data

ABSTRACT

Capsule: We document previously undescribed nocturnal flight behaviour by Bearded Vultures Gypaetus barbatus using a combination of accelerometer and global positioning system (GPS) information.

Aims: To study the nocturnal flight activity of the Bearded Vulture and determine whether nocturnal flights could be linked to foraging behaviour.

Methods: We used both accelerometer and GPS location data of 11 Bearded Vultures in the Spanish Pyrenees along with 88 carcasses monitored with camera traps.

Results: Over half (55%, n?=?11) of the individuals tracked were recorded flying between 0.7 and 6.1?km on at least 19 different nights, including 37% that occurred when less than 20% of the moon was illuminated. Bearded Vultures displayed feeding activity in only 8.2% of the 146 feeding events existing during the hour after dawn and the hour before dusk.

Conclusions: Our findings suggest that foraging benefits do not explain the nocturnal flights. Disturbances or adverse weather conditions may result in the abandonment of an overnight roosting site. This could also explain why individuals recovered in the field showed impact injuries.     

Breeding ecology of a suburban population of Woodpigeons Columba palumbus in northwest England

Capsule Between 1981 and 2008 population size was stable, but there were negative trends in breeding parameters.

Aims To determine the current status and long‐term population trend of an isolated breeding population of Bearded Vultures Gypaetus barbatus (Corsica, Mediterranean).

Methods The total Bearded Vulture population was monitored between 1981 and 2008.

Results The current effective breeding population size of Bearded Vultures in Corsica is ten pairs/trios with a slight increase of one to two pairs since 1983. The population is currently estimated at 25 individuals. Breeding parameters (laying rate, breeding success and productivity) have decreased significantly over the full 28‐year study period, although the decrease was not significant when the data set was restricted to 1988–2008. A mean of 60.3% (n = 204) of pairs have laid, but this proportion is highly variable between years. Productivity has been very low (0.16 young/pair/year, n = 233). Breeding parameters of the Corsican population of Bearded Vultures are very low compared with those of other western European populations in the Pyrenees.

Conclusions This isolated insular population is of small size (eight to ten pairs/trios) but shows a stability of distribution and numbers, but low (and decreasing) breeding rates, making this insular population one the most threatened in Europe.     

Cosmetic plumage coloration by iron oxides does not confer protection against feather wear

The active acquisition of plumage cosmetic coloration by iron oxides has been described in several bird species (notably the Bearded Vulture Gypaetus barbatus, but also in cranes, pelicans and other vulture species) but its function remains controversial. We tested the hypothesis that iron oxide acquired through mud bathing protects feathers from wear. We experimentally stained 13 breast feathers collected from two White Storks Ciconia ciconia with mud from a ferruginous pond regularly used by Bearded Vultures, while another 13 feathers (controls) were treated with water. All feathers were exposed to a controlled physical abrasion treatment. We found no differences in wear between control and stained feathers, which is counter to predictions of the wear protection hypothesis for the acquisition of this cosmetic plumage coloration.     

Polygyny does not explain the superior competitive ability of dominant ant associates in the African ant‐plant,Acacia (Vachellia) drepanolobium

The Acacia drepanolobium (also known as Vachellia drepanolobium) ant‐plant symbiosis is considered a classic case of species coexistence, in which four species of tree‐defending ants compete for nesting space in a single host tree species. Coexistence in this system has been explained by trade‐offs in the ability of the ant associates to compete with each other for occupied trees versus the ability to colonize unoccupied trees. We seek to understand the proximal reasons for how and why the ant species vary in competitive or colonizing abilities, which are largely unknown. In this study, we use RADseq‐derived SNPs to identify relatedness of workers in colonies to test the hypothesis that competitively dominant ants reach large colony sizes due to polygyny, that is, the presence of multiple egg‐laying queens in a single colony. We find that variation in polygyny is not associated with competitive ability; in fact, the most dominant species, unexpectedly, showed little evidence of polygyny. We also use these markers to investigate variation in mating behavior among the ant species and find that different species vary in the number of males fathering the offspring of each colony. Finally, we show that the nature of polygyny varies between the two commonly polygynous species, Crematogaster mimosae and Tetraponera penzigi: in C. mimosae, queens in the same colony are often related, while this is not the case for T. penzigi. These results shed light on factors influencing the evolution of species coexistence in an ant‐plant mutualism, as well as demonstrating the effectiveness of RADseq‐derived SNPs for parentage analysis.     

Orchids with scarce occurrence records: The case of the endangered Botany Bay Bearded Greenhood

Poorly studied species are potentially under-prioritised by conservation programmes due to knowledge gaps presenting barriers to informing effective management strategies. The Botany Bay Bearded Greenhood, Pterostylis sp. Botany Bay, is an example of a poorly studied plant that is listed as endangered under both Commonwealth and New South Wales legislation. This study reports on archival surveys from 1998 to 2005 conducted at up to eight sites known to contain the Botany Bay Bearded Greenhood and follow-up surveys in 2022. The archival surveys found that the total population count ranged from 83 to 341 individuals. Mean numbers (± standard error) of seedlings, flowering individuals and individuals exhibiting capsule development recorded in the population were 43 ± 14, 33 ± 9 and 2 ± 1 respectively. The 2022 surveys did not detect the species. An area of potential habitat was derived from records in biodiversity databases, which determined a total historical extent of 1.38 ha restricted to the Kurnell Peninsula in Sydney, New South Wales. The priority next step is to gather contemporary data to confirm whether the Botany Bay Bearded Greenhood remains extant, which can be done by applied researchers, students, landholders and land managers, botanical practitioners and/or volunteers. For optimal species detection, we recommend undertaking formal surveys or opportunistic searches in historically known sites and replicating detection efforts across July–September periods to exploit seasonal flowering.     

Reduced genetic diversity in Bearded Vultures Gypaetus barbatus in Southern Africa

The Bearded Vulture Gypaetus barbatus occurs throughout its range in small and dwindling population fragments with limited genetic differentiation between populations, suggesting that the species might be managed as a single entity. The numbers of East and Southern African Bearded Vultures included in previous studies were small, so we determine the genetic variation within, evolutionary placement of and connectivity among sub‐Saharan African populations. Mitochondrial DNA fragment analyses detected little or no differentiation between populations in Ethiopia and Southern Africa, with reduced haplotype diversity in Southern Africa compared with populations in the Northern Hemisphere. The results inform conservation management of this species globally and locally, and offer guidelines for translocations should populations continue to decline.     

Infestation by the mite Harpirhynchus nidulans in the Bearded Tit Panurus biarmicus

Capsule Mites sometimes induced voluminous subcutaneous cysts in featherless parts.

Aims To describe the first reported infestation by the skin-dweller mite Harpirhynchus nidulans in Bearded Tits Panurus biarmicus and for the Timaliidae family, to detect possible fitness costs for the host and to determine the distribution of the parasite within the distribution range of the host.

Methods Parasites were identified using a microscope. Wing-length and body mass were recorded on both uninfected and infected birds captured at different times during the year. We also considered historical data, and contact was made with 32 European ringing stations to identify the distribution range of the parasite.

Results Subcutaneous reproduction of the mite Harpirhynchus nidulans induced the development of voluminous dermal nodules in Panurus biarmicus. There were no differences in body mass or wing-length with respect to infestation. In the south of France, prevalence changed from 10.6% in spring to 4.7% in autumn. Both sexes are equally parasitized. Occurrence of dermal cysts is reported from several southern European populations of Bearded Tits, whereas it seems to be absent from northern latitudes.

Conclusion The occurrence of a Harpirhynchus mite in wild bird populations is reported for the first time. We consider aspects of its biology, host–parasite system, host-specificity, co-adaptation of the mite reproductive cycle to the social dynamics of its host and metapopulational host–parasite dynamics.     

Female defense polygyny in the territorial triggerfish <Emphasis Type="Italic">Sufflamen chrysopterum</Emphasis>

The occurrence of polygyny requires specific environmental conditions such as female aggregation or patchy resource distribution. However, it is difficult to determine the factors responsible for polygyny in species in which the territories of both sexes overlap. To overcome this, we performed female removal experiments in the polygynous triggerfish Sufflamen chrysopterum (Balistidae) in the Okinawa coral reef. Both sexes defended their territories exclusively against consexuals of the same species, and female aggregation was absent. Each male territory included 1–3 female territories, and nonterritorial males were significantly smaller than territorial males. Further, the body size of territorial males was positively correlated with that of the largest female in their territories, and larger males tended to mate with more females. The results of the female removal experiments (n = 10 females) indicated that females competed for better territories rather than larger mates. In contrast, males abandoned the territories once the females emigrated. These results strongly suggest that males defend females rather than sites and compete for larger and a greater numbers of females. Thus, in S. chrysopterum, female defense polygyny occurs in the absence of female aggregation.     

MINUTES OF THE ANNUAL GENERAL MEETING OF THE SOUTH AFRICAN ORNITHOLOGICAL SOCIETY,HELD AT CAPE TOWN, 26 JUNE 1959

Brown, C. J. 1989. Plumages and measurements of the Bearded Vulture in southern Africa. Ostrich 60: 165–171.

Four different age classes of the southern African Bearded Vulture Gypaetus barbatus are recognized and their plumages described: juvenile (3–24 months old), immature (24–45 months), subadult (45–60 months) and adult (60+ months). There was no significant difference in size between adult male and female birds. Adults were larger than juvenile birds in bill width, beard length, wingspan and mass, and had a higher aspect ratio and wing loading, while juvenile birds were larger than adults in the length of their outer rectrices, tail area, wing breadth and wing area. These features are considered to be adaptive to young birds inexperienced in flying. Immature and subadult birds were intermediate in size between juveniles and adults. Bearded Vultures differ from other large raptors in two sets of physical characteristics, (a) those adapted to cold, mountainous habitat, e.g. feathered head and face, unusually long wings, a high aspect ratio and a particularly long tail, and (b) those adapted to their diet of mainly bones, e.g. wide gape, beard and relatively long talons for carrying food.     

Ice seals as sentinels for algal toxin presence in the Pacific Arctic and subarctic marine ecosystems

Domoic acid (DA) and saxitoxin (STX)-producing algae are present in Alaskan seas, presenting exposure risks to marine mammals that may be increasing due to climate change. To investigate potential increases in exposure risks to four pagophilic ice seal species (Erignathus barbatus, bearded seals; Pusa hispida, ringed seals; Phoca largha, spotted seals; and Histriophoca fasciata, ribbon seals), this study analyzed samples from 998 seals harvested for subsistence purposes in western and northern Alaska during 2005–2019 for DA and STX. Both toxins were detected in bearded, ringed, and spotted seals, though no clinical signs of acute neurotoxicity were reported in harvested seals. Bearded seals had the highest prevalence of each toxin, followed by ringed seals. Bearded seal stomach content samples from the Bering Sea showed a significant increase in DA prevalence with time (logistic regression, p = .004). These findings are consistent with predicted northward expansion of DA-producing algae. A comparison of paired samples taken from the stomachs and colons of 15 seals found that colon content consistently had higher concentrations of both toxins. Collectively, these results suggest that ice seals, particularly bearded seals (benthic foraging specialists), are suitable sentinels for monitoring HAB prevalence in the Pacific Arctic and subarctic.     

THE INFLUENCE OF MATING SYSTEM AND OVERLAPPING GENERATIONS ON EFFECTIVE POPULATION SIZE

The effective population size (N_e) depends strongly on mating system and generation time. These two factors interact such that, under many circumstances, N_e is close to N/2, where N is the number of adults. This is shown to be the case for both simple and highly polygynous mating systems. The random union of gametes (RUG) and monogamy are two simple systems previously used in estimating N_e, and here a third, lottery polygyny, is added. Lottery polygyny, in which all males compete equally for females, results in a lower N_e than either RUG or monogamy! Given nonoverlapping generations the reduction is 33% for autosomal loci and 25% for sex-linked loci. The highly polygynous mating systems, harem polygyny and dominance polygyny, can give very low values of N_e/N when the generation time (T) is short. However, as T is lengthened, N_e approaches N/2. The influence of a biased sex ratio depends on the mating system and, in general, is not symmetrical. Biases can occur because of sex differences in either survival or recruitment of adults, and the potential for a sex-ratio bias to change N_e is much reduced given a survival bias. The number of juveniles present also has some influence: as the maturation time is lengthened, N_e increases.     

Facultative polygyny in Ectatomma tuberculatum (Formicidae, Ectatomminae)

Summary. Polygyny, the presence of several mated queens within the same colony, is widespread in insect societies. This phenomenon is commonly associated with ecological constraints such as limited nest sites. In habitats where solitary nest foundation is risky, monogynous colonies can reintegrate young daughter queens (secondary polygyny). We studied the reproductive structure (i.e. queen number) of the ectatommine ant Ectatomma tuberculatum from Bahia State, Brazil. This species was found to present facultative polygyny: out of a total of 130 colonies collected, 39.2% were monogynous, while 43.8% were polygynous. Polygynous colonies had significantly more workers than monogynous ones. Queen number in polygynous colonies ranged from 2 to 26, with an average of 4 ± 4 queens per colony. All nestmate queens were egg-layers with no apparent dominance hierarchy or agonistic behavior. Non-nestmate queens were adopted by monogynous colonies suggesting that polygyny is secondary, originating through queen adoption. This species is characterized by an open recognition system, which probably allows a switch from monogynous to polygynous colonies. The behavioral acts of queens showed that resident queens remained frequently immobile on or near the brood, contrarily to alien or adopted queens and gynes. In addition, monogynous queens showed no behavioral or physiological (i.e. by ovarian status) differences in comparison with polygynous ones. Secondary or facultative polygyny, probably associated with queen adoption, may have been favored in particular environmental conditions. Indeed, by increasing colony productivity (i.e. number of workers) and territory size (by budding and polydomy), polygyny could uphold E. tuberculatum as a dominant species in the mosaic of arboreal ants in Neotropical habitats.Received 7 April 2004; revised 11 November 2004; accepted 15 November 2004.     

Entwicklung und Mauser der Spinalflur bei der Bartmeise(Panurus biarmicus)

Zusammenfassung Die Spinalflur nestjunger Bartmeisen(Panurus biarmicus) wird etwa rhombenförmig angelegt. Ihre lateralen Federn werden jedoch schwach ausgebildet und erinnern an Plumae; sie werden sehr bald abgeworfen, so daß flügge Vögel eine schmale, bandförmige Rückenflur haben, die seitlich von nackter Haut (auf der manchmal letzte Federchen sitzen) begrenzt wird. Dieser laterale Teil der Spinalflur kann dann makroskopisch von einem Rain nicht unterschieden werden.Vor Beginn der Jugendvollmauser werden in diesem lateralen Rückenflurbereich durch die Haut Federanlagen sichtbar; die Mauser der Rückenflur (und sehr oft die Mauser überhaupt) wird bei 60% der untersuchten Vögel durch gleichzeitig einsetzendes Wachstum dieser Federn eingeleitet. Die Rückenflur der vermauserten Vögel ist wieder rhombisch wie beim Nestling und bleibt dies fortan.Altvögel werfen lange vor Beginn der Mauser die meisten Plumae ab; die Unterdrückung der lateralen Rückenflur bei Jungvögeln ist wohl eine Analogie dazu.Die Mauser der Rückenflur setzt großflächig ein und unterscheidet sich dadurch von der der Schulter- und Unterflur, wo sie reihenweise, von der Flurmitte ausgehend, erfolgt.Die gesamte Mauser der Bartmeise dauert etwa 50–60 Tage.

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Summary In Bearded Tits(Panurus biarmicus) which are a few days old, the dorsal feather tract is of rhombic shape. The feathers in the lateral part of this tract are feebly developed and are shed soon after the birds leave the nest. Thus the dorsal tract of these birds is in the form of a narrow stripe, flanked by naked skin on both sides. Before the onset of the complete post-juvenile moult, papillae of feathers begin to be visible again under the skin in this area. In 60% of the investigated birds, the growth of dorsal feathers began here, and in many cases this represented the start of the growth of adult plumage as well. The dorsal tract in adult birds is of rhombic shape again and remains so for the rest of their life.Adult Bearded Tits shed their down long before moult. It can, therefore, be concluded that this and the suppression of dorsolateral feathers in juvenile plumage are analogous phenomena.In the dorsal tract of moulting Bearded Tits, feather growth begins in one to three centers simultaneously, whereas moult in the ventral and shoulder tracts starts with the inner rows and proceeds towards the outer ones.Whole moult lasts about 50–60 days in Bearded Tits.

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Herrn Prof. Dr. Rainer Schubert-Soldern zum 70. Geburtstag herzlichst gewidmet.     

Polygyny in thief ants responds to competition and nest limitation but not food resources

Colonies of ants often house multiple queens, and variation in polygyny often tracks environmental conditions. Three hypotheses have been proposed to describe how environmental variation may account for the degree of polygyny: competition, food limitation and nest limitation. Here I evaluate these hypotheses with studies on litter-nesting thief ants (Solenopsis spp.) throughout a lowland tropical rain forest in Costa Rica. In one component, I measured how polygyny varied across a broad environmental gradient demonstrating substantial variation in resources and competition. In a second component, I manipulated the abundance of food, the spatial presentation of food and the availability of nesting space to assess the effects on queen number. The degree of polygyny increased with nest limitation and competition, but there was no indication that colonies produce queens to capitalize on food availability. The increase in queen number in response to the density of competitors suggests that an increase in queen number enhances exploitative abilities.     

Extreme life history plasticity and the evolution of invasive characteristics in a native ant

Disturbance resulting from urbanization is a leading cause of biotic homogenization worldwide. Native species are replaced with widespread non-native species and ants are among the world’s most notorious invaders. To date, all documented cases of ant invasions involve exotic introduced species that are spread around the world by human-mediated dispersal. I investigated the effect of urbanization on the evolution of invasive characteristics in a native ant species, the odorous house ant, Tapinoma sessile (Say). Colony social structure, life history traits, and the spatial pattern of nest distribution were compared by sampling T. sessile across a gradient of three distinct habitats: natural, semi-natural, and urban. Results demonstrate a remarkable transition in colony social and spatial structure and life history traits between natural and urban environments. In natural habitats, T. sessile colonies are comprised of small, monogyne (single queen), and monodomous (single nest) colonies. In urban areas, T. sessile often exhibit extreme polygyny and polydomy, form large supercolonies, and become a dominant pest. Results also suggest that urban T. sessile colonies may have a negative impact on native ant abundance and diversity. In the natural environment T. sessile coexisted with a wide array of other ant species, while very few ant species were present in the urban environment invaded by T. sessile. Habitat degradation and urbanization can lead to extreme changes in social and spatial colony structure and life history traits in a native ant species and can promote the evolution of invasive characteristics such as polygyny, polydomy, and supercolonial colony structure.     

On the song resumption, polyterritorial behaviour and their population context in the Sedge Warbler Acrocephalus schoenobaenus

Mating tactics in species with facultative polygyny seem to be very flexible and to depend on local environmental conditions. We analysed the habitat and population contexts of territorial behaviour, associated with polygyny, in a population of Sedge Warblers Acrocephalus schoenobaenus inhabiting natural wetlands. Nearly one-third of all breeding males (46 individuals) resumed song after completing their mating with the first female, in order to attract another one. Resuming males were usually the earliest arrivals. There was a continuity in the territorial behaviour between resuming song on first territory and polyterritorial behaviour. Fifty-nine percent of resuming males set up second territories, clearly separated from the first. The second territory was usually located close to the primary female activity area. The quality of the first and second territories were correlated, with second territories being significantly inferior. However, their quality was not significantly different from the territories of non-breeding males. The quality of the second territories was also negatively related to the distance from the first territories. Polyterritorialism was influenced by population numbers: the higher the number of territorial males, the lower the number of second territories settled and the lower their average quality. Although the polygyny frequency in the studied population was very low, polyterritorial males were significantly more likely to be polygynous. We concluded that polygyny frequency can be significantly influenced by population numbers, which might be one of the main factors responsible for the variability in the mating system in this species.     

Nesting patterns,ecological correlates of polygyny and social organization in the neotropical arboreal ant <Emphasis Type="Italic">Odontomachus hastatus</Emphasis> (Formicidae,Ponerinae)

Queen number varies in the population of O. hastatus in SE Brazil. Here, we evaluate how nesting ecology and colony structure are associated in this species, and investigate how reproduction is shared among nestmate queens. Queen number per colony is positively correlated with nesting space (root cluster of epiphytic bromeliads), and larger nest sites host larger ant colonies. Plant samplings revealed that suitable nest sites are limited and that nesting space at ant-occupied bromeliads differs in size and height from the general bromeliad community. Dissections revealed that queens in polygynous colonies are inseminated, have developed ovaries, and produce eggs. Behavioral observations showed that reproduction in polygynous colonies is mediated by queen–queen agonistic interactions that include egg cannibalism. Dominant queens usually produced more eggs. Field observations indicate that colonies can be initiated through haplometrosis. Polygyny in O. hastatus may result either from groups of cofounding queens (pleometrosis) or from adoption of newly mated queens by established colonies (secondary polygyny). Clumping of bromeliads increases nest space and probably adds stability through a strong root system, which may promote microhabitat selection by queens and favor pleometrosis. Rainstorms that frequently knock down bromeliads can be a source of colony break-up and may promote polygyny. Bromeliads are limited nest sites and may represent a risk for young queens leaving a suitable nest, thus favoring secondary polygyny. Although proximate mechanisms mediating queen number are poorly understood, this study suggests that heterogeneous microhabitat conditions probably contribute to the coexistence of variable forms of social structure in O. hastatus.     

Sexual size dimorphism and sexual selection in primates

1. In most animals, females are larger than males. Paradoxically, sexual size dimorphism is biased towards males in most mammalian species. An accepted explanation is that sexual dimorphism in mammals evolved by intramale sexual selection. I tested this hypothesis in primates, by relating sexual size dimorphism to seven proxies of sexual selection intensity: operational sex ratio, mating system, intermale competition, group sex ratio, group size, maximum mating percentage (percentage of observed copulations involving the most successful male), and total paternity (a genetic estimate of the percentage of young sired by the most successful male).
2. I fitted phylogenetic generalised least squares models using sexual size dimorphism as the dependent variable and each of the seven measures of intensity of sexual selection as independent variables. I conducted this comparative analysis with data from 50 extant species of primates, including Homo sapiens, Pan troglodytes, and Gorilla spp.
3. Sexual dimorphism was positively related to the four measures of female monopolisation (operational sex ratio, mating system, intermale competition, and group sex ratio) and in some cases to group size, but was not associated with maximum mating percentage or total paternity. Additional regression analyses indicated that maximum mating percentage and total paternity were negatively associated with group size.
4. These results are predicted by reproductive skew theory: in large groups, males can lose control of the sexual behaviour of the other members of the group or can concede reproductive opportunities to others. The results are also consistent with the evolution of sexual size dimorphism before polygyny, due to the effects of natural, rather than sexual, selection. In birds, the study of molecular paternity showed that variance in male reproductive success is much higher than expected by behaviour. In mammals, recent studies have begun to show the opposite trend, i.e. that intensity of sexual selection is lower than expected by polygyny.
5. Results of this comparative analysis of sexual size dimorphism and sexual selection intensity in primates suggest that the use of intramale sexual selection theory to explain the evolution of polygyny and sexual dimorphism in mammals should be reviewed, and that natural selection should be considered alongside sexual selection as an evolutionary driver of sexual size dimorphism and polygyny in mammals.

     

Diet and food preferences of the endangered Bearded Vulture Gypaetus barbatus: a basis for their conservation

The Bearded Vulture Gypaetus barbatus population in the Pyrenees is managed using feeding stations to increase breeding success and reduce mortality in the pre-adult population. Nevertheless, very little quantitative and qualitative information has been published on such basic aspects of the species' ecology as feeding habits and dietary preferences. This study investigated both aspects through direct and unbiased observation of breeding Bearded Vultures during the chick-rearing period. Bearded Vulture diet comprises mammals (93%), birds (6%) and reptiles (1%), with medium-sized ungulates (mainly sheep/goats) the most important species in the diet (61%, n  = 677). Prey items were not selected in proportion to their availability, with the remains of larger species (cows and horses) being avoided, probably due to the variable cost/benefit ratios in handling efficiency, ingestion process and transport. There is no relationship between the proportion of sheep limbs in the diet and the proximity of feeding stations, suggesting that these sites are probably less important for breeding adults than for the pre-adult population. On the other hand, diet specificity seems related to productivity, with territories with greater trophic breadth being those with higher fecundity. Bearded Vultures prefer to eat limbs, although meat remains (provided principally by small mammals) can play an important role in guaranteeing breeding success during the first few weeks after hatching. The management of carrion provided by animals that die naturally in extensive livestock practices and the remains of wild ungulates which have died naturally or by human hunting, are important conservation tools for the Bearded Vulture and other carrion-eating species.