卫星跟踪猎隼失联案例分析

猎隼(Falco cherrug)是迅速濒危中的物种。对近10年卫星跟踪的67只猎隼进行国籍、性别、信号时长、失联月份统计,分析其死亡原因、受威胁因素以及年返回率。就其中2016年来自俄罗斯和蒙古的10只猎隼进行更详细的死因分析,当发射器不再返回信息时,前往GPS最后位点,在一定区域内进行拉网式搜索,并分析周围的环境和动物痕迹,将找到的尸体进行解剖,查找死亡原因。结果显示,这67只猎隼中雌鸟有37只,占总数的55.2%,雄鸟有29只,另1只性别未知。其平均信号时长为(201±129.94)d(n=64),小于200 d的占60.9%,小于400 d的占92.2%。年返回率为31.4%;在秋季和初冬季(8～12月)失联的占67.2%。而2016年跟踪的10只猎隼中,获取4只失联猎隼的信息,1只在蒙古国被猎杀,1只死因不详,1只在中国因电击死亡,1只在中国因擦碰和饥饿死亡。卫星跟踪数据显示,猎隼的年返回率明显降低,表明猎隼的处境不容乐观,这引发了猛禽研究专家的担忧。     

Trapping of Saker Falcon Falco cherrug and Peregrine Falcon Falco peregrinus in Saudi Arabia: Implications for biodiversity conservation

The numbers of Falco cherrug and Falco peregrinus trapped during their migration over the Kingdom of Saudi Arabia (KSA) were investigated from published reports and through interviews with well-known trappers and dealers over several years (1989–2013). The number of trapped individuals increased for both species over a 23 year period, which is probably related to an enhanced trapping effort. Time series analysis suggests that the number of Saker Falcons being trapped is likely to be stable with annual fluctuations in the coming ten-year period, whereas the number of trapped Peregrine Falcons will probably decline with a small fluctuation initially. Using the population viability analysis suggests a high extinction rate for the Saker Falcon population migrating through KSA during the coming 10 and 20 years; whereas Peregrine Falcons probably take more than 100 years to reach the extinction threshold. However, the increase in the trapping period, especially in the spring, that has been observed during the last five years could increase the number of falcons trapped in the future. As both falcon species are migratory, implementing conservation actions across all range states is important to ensure a favourable conservation status for the Saker and Peregrine Falcons. Both species will benefit through the implementation of the Global Action Plan (GAP), developed by the Saker Falcon Task Force.     

The use of DNA fingerprinting to estimate annual survival rates in the Saker Falcon(Falco cherrug)

Summary DNA fingerprinting of nestlings ofFalco cherrug was used to determine indirectly the survival of the corresponding adult parent birds, which are difficult to catch in sufficient numbers. This approach is possible because Saker falcons show a high degree of site and mate tenacity. DNA profiles of nestlings from the same territory but from different years were compared. Three patterns of band-sharing coefficients between broods from the same territory were found: if band-sharing coefficients within and between broods from consecutive years were similar but significantly different from those of unrelated birds, it indicated that all young were full sibs and that neither adult was replaced between years. If band-sharing coefficients between broods at the same site indicated no relatedness across years and were equal to those of unrelated birds, then both breeding partners apparently had changed. If the band-sharing coefficients between broods of the same territory and consecutive years were significantly lower than those of full sibs, but higher than those of unrelated birds, the loss of one adult bird was indicated. The analysis of 32 broods (years 1993 to 1997) provided a minimal estimate for annual adult survival of 82% for a wild population of Saker Falcons in Kazakhstan.

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Abschätzung der jährlichen Überlebensraten des Sakerfalken(Falco cherrug) mittels DNA-Fingerprinting

Zusammenfassung Um die Gefährdung und Populationsdynamik des Sakerfalken (Falco cherrug) beurteilen zu können, benötigen wir genaue Angaben zu Mortalität und Überlebensraten. Während es bei dieser Art relativ einfach ist, Nestlinge zu fangen, ist es nahezu unmöglich, eine ausreichend große Zahl an Altvögeln zu markieren, um durch Wiederfang oder Ringfundmeldungen die jährliche Überlebensrate zu ermitteln. Durch DNA-Fingerprinting von Jungfalken haben wir versucht, die minimale Überlebensrate von Altfalken indirekt zu bestimmen. Dieser Forschungsansatz wird dadurch möglich, daß die Sakerfalken eine hohe Philopatrie aufweisen und jedes Jahr im selben Revier brüten. Wenn man mehrere Jahre lang Blutproben der Jungvögel aus denselben Revieren sammelt, so kann man mittels DNA Fingerprinting indirekt ermitteln, ob die jeweiligen Altvögel identisch waren oder gewechselt haben: Vergleicht man die Band-Sharing-Koeffizienten (BSK) von Jungvögeln von zwei oder mehr Jahren aus demselben Revier, so ergeben sich drei Muster: Wenn die BSK-Werte innerhalb der Bruten und zwischen den Bruten identisch aber signifikant verschieden von denen nicht verwandter Vögel sind, so handelt es sich bei den Jungvögeln um Vollgeschwister; demnach sind die Altvögel identisch geblieben, d. h. sie haben von einem Jahr zum nächsten überlebt. Wenn die BSK-Werte zwischen zwei Bruten aus demselben Revier einen Wert annehmen, wie man ihn für unverwandte Tiere ermittelt, so müssen die Eltern gewechselt haben. Liegen die Werte zwischen zwei Bruten signifikant höher als die von nicht verwandten Tieren, aber niedriger als diejenigen von Vollgeschwistern, so ist vermutlich 1 Altvogel gewechselt worden. Die Analyse von 32 Bruten des Sakerfalken aus Kasachstan zeigt, daß die minimale jährliche Adultüberlebensrate bei 82% liegt.

     

Climatic niche of the Saker Falcon Falco cherrug: predicted new areas to direct population surveys in Central Asia

Accurate species distribution data across remote and extensive geographical areas are difficult to obtain. Here, we use bioclimatic envelope models to determine climatic constraints on the distribution of the migratory Saker Falcon Falco cherrug to identify areas in data-deficient regions that may contain unidentified populations. Sakers live at low densities across large ranges in remote regions, making distribution status difficult to assess. Using presence-background data and eight bioclimatic variables within a species distribution modelling framework, we applied MaxEnt to construct models for both breeding and wintering ranges. Occurrence data were spatially filtered and climatic variables tested for multicollinearity before selecting best fit models using the Akaike information criterion by tuning MaxEnt parameters. Model predictive performance tested using the continuous Boyce index (B) was high for both breeding (B_TEST = 0.921) and wintering models (B_TEST = 0.735), with low omission rates and minimal overfitting. The Saker climatic niche was defined by precipitation in the warmest quarter in the breeding range model, and mean temperature in the wettest quarter in the wintering range model. Our models accurately predicted areas of highest climate suitability and defined the climatic constraints on a wide-ranging rare species, suggesting that climate is a key determinant of Saker distribution across macro-scales. We recommend targeted population surveys for the Saker based on model predictions to areas of highest climatic suitability in key regions with distribution knowledge gaps, in particular the Qinghai-Tibet plateau in western China. Further applications of our models could identify protected areas and reintroduction sites, inform development conflicts, and assess the impact of climate change on distributions.     

Morphological variation of the saker (Falco cherrug) and the implications for conservation

The saker (Falco cherrug) is a traditional and valued bird of Arabic falconry. Harvesting birds from the wild raises questions of its conservation status and sustainability. The external morphology of this species is highly variable and the Arabic system of classification identifies over 11 saker types, some of which are more favoured among Arab falconers than others. Our study assessed morphological variation of birds chosen for known natal origin. Although most variation was between individuals-within-regions, the remaining variation in size and plumage characters may be appropriately described as a gradual cline between small dorsally uniform brown sakers from western lowland regions (e.g. lowland Kazakstan, and south-west Russia) to large, dorsally barred russet, dark brown and grey coloured sakers from eastern highland regions (e.g. south-east Russia, Mongolia, and China). Between these extremes exists a plethora of highly variable and contiguous populations. Although preferred falconry types may originate from any of these populations, the preferred jarudi and 'sinjari sakers from northeastern Altai, Tuva, and Sayan populations, and the large sakers from eastern populations may be over exploited through trapping. The demand for wild sakers, however, may be substituted by an increasing number of captive bred falcons used for Arab falconry.     

Turnover and recruitment in a tundra population of Peregrine Falcons Falco peregrinus

The population dynamics of tundra-nesting Peregrine Falcons Falco peregrinus tundrius was studied over 7 years on a 450 km² study area in the Keewatin District of Canada's Northwest Territories. Peregrines showed strong fidelity to nest sites; none of 25 males ringed changed territories, while five of 38 females ringed were recorded changing territories. Such changes usually occurred after nesting failure. Annual turnover of territorial adults was estimated to be 22% (15% for males and 26% for females). Annual mortality of adults was estimated to be 17% (15% for males and 19% for females). If we assumed that territory vacancies, in addition to mate replacements, were indicative of mortality, then maximum annual mortality was estimated at 24% for each sex. Territories were held only by adult Peregrines. The oldest male on territory was at least 7 years old, the youngest was 2. The oldest female on territory was at least 7 years old, the youngest was 3. Territories were held by individuals of each sex for at least 6 years. One pair remained together for at least 4 years. Less than 4% of all young Peregrines produced on the study area in the first 5 years of the study were recruited into the breeding population. More male than female young were recruited despite an even sex ratio among nestlings. Peregrines did poorly in their first breeding attempts. The single young female recruited into the study population dispersed more than three times the median dispersal distance of six recruited males, suggesting that other females probably dispersed beyond the boundaries of the study area.     

Demographic consequences of nestbox use for Red‐footed Falcons Falco vespertinus in Central Asia

Nestbox programmes are frequently implemented for the conservation of cavity‐nesting birds, but their effectiveness is rarely evaluated in comparison with birds not using nestboxes. In the European Palaearctic, Red‐footed Falcon Falco vespertinus populations are both of high conservation concern and are strongly associated with nestbox programmes in heavily managed landscapes. We used a 21‐year monitoring dataset collected on 753 nesting attempts by Red‐footed Falcons in unmanaged natural or semi‐natural habitats to provide basic information on this poorly known species; to evaluate long‐term demographic trends within this population; and to evaluate response of demographic parameters of Red‐footed Falcons to environmental factors including use of nestboxes. We observed significant differences among years in laying date, offspring loss and numbers of fledglings produced, but not in egg production. Of these four parameters, offspring loss and, to a lesser extent, number of fledglings exhibited directional trends over time. Variation in laying date and in numbers of eggs were not well explained by any one model of environmental factors, but instead by combinations of models, each with informative terms for nest type. Nevertheless, laying in nestboxes occurred 2.10 ± 0.70 days earlier than in natural nests. In contrast, variation in both offspring loss and numbers of fledglings produced were fairly well explained by a single model including terms for nest type, nest location and an interaction between the two parameters (65 and 81% model weights, respectively), with highest offspring loss in nestboxes on forest edges. Because, for other species, earlier laying dates are associated with more fit individuals, this interaction highlighted a possible ecological trap, whereby birds using nestboxes on forest edges lay eggs earlier but suffer greater offspring loss and produce lower numbers of fledglings than do those in other nesting settings. If nestboxes increase offspring loss for Red‐footed Falcons in heavily managed landscapes where populations are at greater risk, or for the many other species of rare or endangered birds supported by nestbox programmes, these processes could have important demographic and conservation consequences.     

Nesting ecology of solitary‐nesting Amur Falcons (Falco amurensis) in central Mongolia

Amur Falcons (Falco amurensis) are a migratory species that face a variety of threats across their range, but little is known about their breeding ecology. These falcons breed in forest habitats in Eastern and Central Asia using nests constructed by corvids, including Eurasian Magpies (Pica pica). We monitored nests of 21 pairs of Amur Falcons at Hustai National Park in central Mongolia in 2017. Our objectives were to describe their basic nesting ecology, estimate nest survival by modeling the daily survival rate (DSR), examine nest selection by modeling it as a function of nest and site covariates, and use a spatial simulation to test hypotheses concerning intra‐ and interspecific avoidance. Clutch sizes averaged 4.1 eggs (N = 21 nests), and incubation and nestling periods averaged 25.7 and 26.1 d, respectively. The daily survival rate was 0.98, with young in 12 nests surviving to fledging. Nest structures were more likely to be selected as percent cover of nest bowls increased, usually in the form of a dome of sticks with multiple side entrances. Closed nests likely provide increased protection from predators. In contrast to congeneric Red‐footed Falcons (F. vespertinus) that nest in large colonies, Amur Falcons nested no farther from or closer to nests of either conspecifics or congeners than expected by chance. One factor likely contributing to this difference is that Red‐footed Falcons often use the nests of colonial‐nesting Rooks (Corvus frugilegus), whereas Amur Falcons typically use the nests of non‐colonial Eurasian Magpies. The ongoing loss of deciduous trees like white birch (Betula platyphylla) across the breeding range of Amur Falcons, probably due to climate change and increased grazing pressure, is likely to reduce the availability of nesting habitat for Eurasian Magpies which, in turn, will likely reduce availability of nests for Amur Falcons and other small falcons.     

Island accessibility and distance from beach influence nesting success of Sooty Falcons Falco concolor in Oman

Colonial island‐breeding birds can be particularly vulnerable to anthropogenic disturbance, which can adversely affect their nesting success. We studied Sooty Falcons Falco concolor breeding on 10 ground‐predator‐free islands in the Sea of Oman during 2007–2014 and evaluated spatio‐temporal trends in the number of breeding pairs occurring on the islands and the factors influencing nesting success. The number of breeding pairs on the islands declined during the study, due mostly to the decline on accessible islands; the rate of decline on islands accessible to humans was double that on inaccessible ones. The number of nests with one or more eggs declined during the study period, and the percentage of nests with eggs that produced one or more chicks showed an increasing trend over time. Sooty Falcon nests located farther away from beaches experienced a significantly higher probability of nesting success than those located closer to beaches. Our results suggest that the number of breeding Sooty Falcons on the islands of northern Oman is declining and that human disturbance may be a contributing factor; this probably mirrors the situation in other parts of the breeding range of this species.     

Territory occupancy and breeding success of Peregrine Falcons Falco peregrinus at various stages of population recovery

Organochlorine pesticides disrupted reproduction and killed many raptorial birds, and contributed to population declines during the 1940s to 1970s. We sought to discern whether and to what extent territory occupancy and breeding success changed from the pesticide era to recent years in a resident population of Peregrine Falcons Falco peregrinus in southern Scotland using long‐term (1964–2015) field data and multi‐state, multi‐season occupancy models. Peregrine territories that were occupied with successful reproduction in one year were much more likely to be occupied and experience reproductive success in the following year, compared with those that were unoccupied or occupied by unsuccessful breeders in the previous year. Probability of territory occupancy differed between territories in the eastern and western parts of the study area, and varied over time. The probability of occupancy of territories that were unoccupied and those that were occupied with successful reproduction during the previous breeding season generally increased over time, whereas the probability of occupancy of territories that were occupied after failed reproduction decreased. The probability of reproductive success (conditional on occupancy) in territories that were occupied during the previous breeding season increased over time. Specifically, for territories that had been successful in the previous year, the probability of occupancy as well as reproductive success increased steadily over time; these probabilities were substantially higher in recent years than earlier, when the population was still exposed to direct or residual effects of organochlorine pesticides. These results are consistent with the hypothesis that progressive reduction, followed by a complete ban, in the use of organochlorine pesticides improved reproductive success of Peregrines in southern Scotland. Differences in the temporal pattern of probability of reproductive success between south‐eastern and south‐western Scotland suggest that the effect of organochlorine pesticides on Peregrine reproductive success and/or the recovery from pesticide effects varied geographically and was possibly affected by other factors such as persecution.     

The breeding status of Peregrine Falcons Falco peregrinus in the UK and Isle of Man in 2002

Capsule Numbers of breeding Peregrines are now higher than ever recorded and the recovery of the species within the UK has continued since 1991.

Aims To estimate the current Peregrine breeding population and its changes over time by recording breeding evidence and nesting range occupancy at known and potential Peregrine eyries.

Methods Attempts were made to check all potentially suitable Peregrine nesting locations for signs of Peregrine occupation. Causes of nest failure were documented where possible. Estimates of occupancy of nesting ranges that could not be visited were based on the proportion of ranges occupied from visited nesting ranges in the same region.

Results In 2002, 1530 Peregrine nesting ranges were estimated to be occupied, comprising 1456 pairs or single birds confirmed as in occupation, and 74 estimated extras (5% of total occupied nesting ranges) that were not visited. Including non‐breeding singletons holding nesting ranges, the UK and Isle of Man figure of ‘Ratcliffe pairs’ was 1437. This showed a 12% increase from 1991 and a 64% increase from the 1930s. However, there were geographical differences in change, with rapid expansion in the south of England tempered by declines in northern Scotland.

Conclusions The primary reasons for the population increase is primarily because of range expansion driven by a plentiful food supply and perhaps an increased tolerance of human disturbance. In areas where the species has declined, various factors (separately or in combination) including reduced food supply and persecution are likely to be involved.     

Autumn migration and wintering areas of Peregrine Falcons Falco peregrinus nesting on the Kola Peninsula, northern Russia

Four female Peregrine Falcons Falco peregrinus breeding on the Kola Peninsula, Russia, were fitted with satellite-received transmitters in 1994. Their breeding home ranges averaged 1175 (sd = ±714) km², and overlapped considerably. All left their breeding grounds in September and migrated generally south-west along the Baltic Sea. The mean travel rate for three falcons was 190 km/day. Two Falcons wintered on the coasts of France and in southern Spain, which were, respectively, 2909 and 4262 km from their breeding sites. Data on migration routes suggested that Falcons took a near-direct route to the wintering areas. No prolonged stopovers were apparent. The 90% minimum convex polygon winter range of a bird that migrated to Spain encompassed 213 km² ( n  = 54). The area of the 50% minimum convex polygon was 21.5 km² ( n  = 29). Data from this study agree with others from North America that show that Falcons breeding in a single area do not necessarily follow the same migratory path southward and do not necessarily use the same wintering grounds.     

Turnover and post‐bottleneck genetic structure in a recovering population of Peregrine Falcons Falco peregrinus

Dispersal is a process that increases genetic diversity and genetic connectivity of populations. We studied the turnover rate of breeding adults and genetic population structure to estimate dispersal in Peregrine Falcons in Finland. We used relatedness estimates among Finnish Peregrine Falcons over a 5‐year period, genotyping over 500 nestlings with 10 microsatellite loci to reveal the rate of turnover. Our results reveal a high turnover rate (21.7%) that does not seem to be correlated with the breeding success of the previous year. The extent of population genetic structure and diversity, and possible signs of the population crash during the 1970s, was assessed with a reduced dataset, excluding relatives. We found genetic diversity to be similar to previously studied falcon populations (expected heterozygosity of 0.581) and no population genetic structuring among our sampled populations. We did not find a genetic imprint of the past population bottleneck that the Finnish Peregrine population experienced. We conclude that high dispersal rates are likely to have contributed to maintaining genetic diversity across the landscape, by mixing individuals within the species’ distribution in Finland and thus preventing genetic structuring and negative effects associated with the population decline in the 1970s.     

Effects of satellite transmitters fitted to breeding Cory's shearwaters

We studied the effects of a new technique to fix satellite transmitters on the feeding behavior and chick body growth of the Cory's shearwater (Calonectris diomedea). Transmitters did not significantly affect the breeding bird's body mass or diving performance. However, on average, pairs with ≥1 adult equipped with a transmitter brought a lighter meal (54.8 ± 18.9 g) to nestlings than did control birds or the same pair with the transmitter removed (77.4 ± 8.3 g) leading to a decreased chick body growth. However at fledging, chicks in control and treatment groups had similar body mass whether parents were equipped with a PTT or not. Our method of transmitter attachment could therefore be an appropriate alternative for studying the ecology of procellariiforms weighing <1 kg given that birds are fitted with transmitters for <2 weeks and restricted to 1 parent only. © 2011 The Wildlife Society.     

Phylogeography and population structure of the saker falcon (Falco cherrug) and the influence of hybridization: mitochondrial and microsatellite data

Microsatellite as well as sequence analysis of the mitochondrial control region were applied to infer phylogeography and population genetic structure of the saker falcon (Falco cherrug). Furthermore, we compared the patterns of mitochondrial haplotypes with the variation of microsatellite alleles among the species of the hierofalcon complex (F. cherrug, Falco rusticolus, Falco biarmicus, Falco jugger) to test hypotheses on population history. Historical samples from museum specimens of F. cherrug were analysed together with samples from contemporary populations to investigate possible influences of hybrid falcons escaped from falconry on the genetic composition. In the mitochondrial DNA analysis, none of the four species represents a monophyletic group. Moreover, there are no clearly defined groups of haplotypes corresponding to taxonomic entities. In the microsatellite analysis most of the variation is shared between species and no clear differentiation by private alleles is found. Yet, with a Bayesian clustering method based on allele frequencies, a differentiation of F. cherrug, F. rusticolus and two geographic groups of F. biarmicus was detected. Results from both nuclear and mitochondrial markers are compatible with the previously postulated 'Out of Africa' hypothesis assuming an African origin of the hierofalcons. From an ancestral African population, F. cherrug, F. rusticolus and F. jugger split off in separate waves of immigration into Eurasia and South Asia. A combination of evolutionary processes, including incomplete lineage sorting as well as hybridization, may be responsible for the currently observed genetic patterns in hierofalcons.     

Evidence for abundant slightly deleterious polymorphisms in bacterial populations

The nearly neutral theory of molecular evolution predicts that slightly deleterious mutations subject to purifying selection are widespread in natural populations, particularly those of large effective population size. To test this hypothesis, the standardized difference between pairwise nucleotide difference and number of segregation sites (corrected for number of sequences) was estimated for 149 population data sets from 84 species of bacteria. This quantity (Tajima's D-statistic) was estimated separately for synonymous (D(syn)) and nonsynonymous (D(non)) polymorphisms. D(syn) was positive in 70% of data sets, and the overall median D(syn) (0.873) was positive. By contrast D(non) was negative in 68% of data sets, and the overall median D(non) (-0.656) was negative. The preponderance of negative values of D(non) is evidence that there are widespread rare nonsynonymous polymorphisms in the process of being eliminated by purifying selection, as predicted to occur in populations with large effective size by the nearly neutral theory. The major exceptions to this trend were seen among surface proteins, particularly those of bacteria parasitic on vertebrates, which included a number of cases of polymorphisms apparently maintained by balancing selection.     

Evidence for hitchhiking of deleterious mutations within the human genome

Deleterious mutations present a significant obstacle to adaptive evolution. Deleterious mutations can inhibit the spread of linked adaptive mutations through a population; conversely, adaptive substitutions can increase the frequency of linked deleterious mutations and even result in their fixation. To assess the impact of adaptive mutations on linked deleterious mutations, we examined the distribution of deleterious and neutral amino acid polymorphism in the human genome. Within genomic regions that show evidence of recent hitchhiking, we find fewer neutral but a similar number of deleterious SNPs compared to other genomic regions. The higher ratio of deleterious to neutral SNPs is consistent with simulated hitchhiking events and implies that positive selection eliminates some deleterious alleles and increases the frequency of others. The distribution of disease-associated alleles is also altered in hitchhiking regions. Disease alleles within hitchhiking regions have been associated with auto-immune disorders, metabolic diseases, cancers, and mental disorders. Our results suggest that positive selection has had a significant impact on deleterious polymorphism and may be partly responsible for the high frequency of certain human disease alleles.     

Effects of implanted satellite transmitters on behavior and survival of female common eiders

Surgically implanted satellite transmitters have been widely used in studies of avian ecology, yet little is known about their potential impacts on birds. We implanted satellite transmitters with percutaneous antennae (approx. 50 g) in 17 female common eiders (Somateria mollissima) at a breeding colony in Arctic Canada. Among females implanted during incubation, 11 of 12 nests were abandoned within 1 week of being radioed. We observed no differences in the proportion of time that implanted female eiders allocated to basic behaviors. Radioed birds were more likely to pick or preen their abdominal (site of surgical incision) and posterior–dorsal (site of antenna exit) regions than unmarked females, although these behaviors were rare (approx. 0.3% of total time budget). Three of 10 females re-observed had a pronounced limp following surgery, but we observed no walking difficulties among these females in subsequent seasons, and we observed some implanted eiders nesting in subsequent years. Mark–resighting models suggest eiders with transmitters had lower apparent survival the year after implantation (67.0%; 85% CI: 47.8–81.9%) than did color-banded eiders (87.5%, 85% CI: 82.5–91.2%), but there was no model support for a survival difference in subsequent years. We conclude that transmitter implantation in common eiders leads to short-term changes in behavior and a decline in first year survival. We encourage researchers to collect similar data on their study subjects where possible and use it to determine the degree to which data are representative of the greater population. © 2011 The Wildlife Society.