Vulture, culture and more: putting human-dimensions back in the saddle for conservation policy

Vulture collapse in South Asia accompanied rapid urbanisation. However, the Indian-Subcontinent’s “Action Plan for Vulture Conservation'' and the Conservation of Migratory Species of Wild Animals’ (CMS) “Multi-species Action Plan to Conserve African-Eurasian vultures” reect poorly on how their scavenging services factored in the regional social-ecological tool - a nature-based-solution. We report the ethnography of the extinction of experience concerning vultures in the tropical megacity of Delhi to contribute to wildlife restoration policies in human-use landscapes. People anthropomorphised avian scavengers while sharing perceptions that promoted ritual feeding of crows and kites. It attracted and supported enormous bird-ocks in the region, an ecological response to the rapid niche-evacuation. Stakeholders’ perceptions that offered links in vulture salience and charisma corresponded with respective socio-cultural legends, based on bird morphology, behaviour, and ecosystem services. Conating with ethnoecology, cultural legends mediated human-animal interface, based on species-specic life-history traits. The latter inextricably tied humans and vultures in their population and demographic parameters and mutual tolerance in behaviour that promoted co-existence. Therefore, wildlife restoration in urbanising landscapes is a moving target, necessitating policies sensitive to progressive loss and/or changes in associative heritage through shifting economic and cultural practices, and socio-cultural stories. In order to uphold their erstwhile functional ecology, vultures would need to behaviourally fathom new built-up spaces, interference from competing scavengers and mediatised misinformation.


Introduction
Vulture decline and its aftermath in expanding human-dominated landscapes of South Asia have been complex processes revealed by transdisciplinary research on the opportunistic obligatory avian scavenger 1 . This complexity was shaped by multifarious interactions in and around the built environment, socio-economy and local cultures that correspond to faunal responses. First, the change in quantity, quality and spatial outlay of solid waste-a characteristic aspect of tropical urban ecosystems which supports an extraordinarily high density of scavengersvultures, kites, crows, dogs, rodents, etc. 2,3 . Secondly, socio-economic transitions oversaw several agricultural movements, like the green and white revolutions, and the development of milk cooperatives 4 in the 20 th century.
These events fuelled the availability of carcasses, attracting and potentially contributing to the survival of huge ocks of avian scavengers. Third, raptor populations are typically limited by food and nest-sites 3 , wherein, relics of native vegetation, tree plantations and other man-made physical structures likely mediate accessibility to carcasses and offal (e.g. [ 5 ]). Lastly but perhaps most de ning are cultural beliefs that sustain patronising tolerance for vultures and other commensals and scavengers, despite social legends which classify them as agents of death associated with bad omens 6 .
Vultures' ecosystem services are myriad, beyond the usually discussed biophysical aspects of their niche as obligate scavengers 7,8 . For centuries, their gregarious feeding has been harnessed as a nature-based-solution 9 in the disposal of cattle carcasses, simultaneously supporting village and town-based processing of bovine leather 10,11 . Depending on the economics and stigma for urban and rural 'leatherwork' and 'leatherworkers' by speci c communities, whose settlements were segregated from the mainstream village and urban dwellers, the extent of human-vulture direct encounters was mediated by certain caste (Chamars) and religious (Muslims) status 10 .
Alongside, leather tanning contributed to small scale industries based on the bovine bone remains-e.g. bone meal fertiliser units in proximity of carcass dumping sites in Delhi 7,11 . Prior studies on opportunistic commensals offer limited insights, considering how human dimensions found infrequent mentions while reporting opportunistic commensal ecology, despite the evidence on inextricable socio-ecological associations in human and non-human animal agencies (e.g. 12,13 ). Collectively, these human entanglements with non-human species re ect on myriad aspects of ecosystem services. In addition to the scavenging, commensals provide information/knowledge about human-dominated ecosystems that have found poor appreciation in literature, e.g. the practice of dokhmenishini or sky burial by the parsi (Zoroastrians) community, using vultures for cremation in dakhma or the Tower of Silence 11,14 .
The dynamics of an erstwhile human-vulture relationship highlights similar connections between animal population densities, key anthropogenic resources and human beliefs at the fulcrum of urban well-being in tropics 15,16 .
However, except for a coarse estimation of the size of the overall Delhi's breeding raptor population, quantitative data for vultures and other raptors in this biogeographic region are extremely scarce 2,5,[17][18][19] . In Galushin's study 2 , vultures constituted 5% of ~3000 breeding pairs of raptors sampled in 150 km 2 , while Black Kites Milvus migrans were prominent urban raptors (83% or 2400 of all breeding pairs), way before the vulture-collapse in the region.
Since raptors as a taxon have relatively long lifespans and young birds evolutionarily delay rst indulgent breeding for the initial 3-5 years of life as predators 3 , the absence of information on the non-breeding population of pre-adults and post-prime adults precludes complete assessment ecological dynamics. Ecology of non-breeding population should cater higher attention in the case of opportunistic commensals, where the superabundance of food can impact breeding success and survivorship 3 . Finally, soaring avian scavengers like vultures and kites can y anywhere between 30-125 km on foraging trips 20 . In various folktales, their keen eyesight and sense of navigation in "landscapes of anthropogenically afforded foraging opportunities'' have enabled entry within the socio-cultural fabric of urban ecosystems 21 .
Unlike the attempts in Europe 22 , reintroducing captive-bred vultures from various centres in the Indian subcontinent within or near the human use landscapes would be the rst of its kind ecological restoration 23,24 . About three decades have gone by since vultures crashed from 40 million to a few thousand, given that synurbic vulture populations were attuned to opportunistically consume cattle carcasses. Carcasses that contained diclofenac, a veterinary drug which affected liver and kidneys of vultures 7,25 , caused indirect poisoning. Meanwhile, rapid increase in human population and their rapid urbanisation-wherein 65% or more people of the Indian subcontinent are currently estimated to be below the age of 35 years 26,27 -effectively subjects extinction of experience in youngsters. Considering that most people in the region have not visually encountered vultures' scavenging ecosystem-services, extinction of experience involves loss of phenotypic and ecological salience. Fortunately, however, perceptual and cultural salience 28 are frequently promoted by 'synergies of reverence', which contributes to the resilience of mutual prosocial perceptions and behaviours for innumerable animal species in the region 2,16 . Under the purview of coproduction of urban ecosystems by humans and more-than-human agencies, people's perceptions modulate nonhuman animals' access to anthropogenic resources and shape human-wildlife interactions within SES in variably urbanised landscapes [29][30][31][32][33][34][35][36][37] . Still, the literature and thereby policies guiding the practice of restoring vultures in human-use landscapes are silent about the mechanisms that erstwhile shaped ecological, aesthetic and corporeal charisma for avian scavengers that affects their salience amongst public 28,38 . Considering highly interspersed periurban, protected and human-use landscapes in the region, their association with food-subsidies are likely to increase proximity and exposure of wildlife to humans, and vice versa. It is an area of research that needs actionable implementation in the Global South 31, 39 .
While rare and endangered species get research and conservation attention, common species provide signi cantly higher ecosystem services 40 , are easily noticeable by citizens, and have been known to face rapid population declines (e.g. red kites, passenger pigeon, vulture, etc) 3 . To contribute to these overlooked aspects that concern every wildlife restoration effort within or near human-use landscapes, we focused on common urban scavenging birds that are readily encountered interacting with residents in Delhi-Black Kites Milvus migrans, crows Corvus splendens, and erstwhile common vultures (Gyps spp. and Neophron spp.). We examined the 'affective attitudes' people have for animals in South Asia, generally driven by anthropomorphic beliefs. Our rationale focused on capturing the changing role of multiple human and nonhuman agencies through semi-structured ethnographic surveys that shed light on the drivers mediating the interactions of direct and mediatised encounters of people with avian commensal scavengers. Analyses of complex relationships among biotic, physical and socio-cultural agencies on the cultural characterization of urban commensals by local citizens identi ed the merits as well as perils of anthropomorphising commensals. We discuss how the effects of urban driven homogenization extend beyond biophysical relationships, impacting human-animal relationships among multiple stakeholders.

Methods
This study is part of long-term research on the ecology and ethno-zoology of urban scavengers in Delhi, seeded to examine the ecological impacts of local vulture loss. The city has rich birdlife and the world's second-largest urban human population of 29 million inhabitants 41 , making it a melting pot of cultures that is composed of immigrants from across the Indian subcontinent 42 (details in Supplementary material 1).
For the horizon scan of ethnoecological aspects in a heterogeneously-developed city, we adapted a Delphi-like ethnographic approach to incorporate Delhi's solid waste management, and a wide range of stakeholders and their interactions with avian scavengers in the frameworks that represent a tropical megacity's human-animal interface 43,44 (Fig. 1). Since 2012, we have been longitudinally surveying commensals and people as co-producing units in the megacity of Delhi at 32 sampling plots of approximately 1 km 2 . The design systematically covers all urban settings, from semi-natural to extremely built-up sites, including all the three sanitary-land lls (for details, see 15,16 ; Supplementary material 1).
During the ongoing study, where ethnography followed a stepwise procedure, we interacted with 27.03 ± 2.31 new onlookers at our sampling units on every eld visit (see Supplementary material 1). Horizon-scans involved people who voluntarily engaged in conversations. It solicited direct inputs on vultures' local extinction and the corresponding prominence of other urban scavengers. The overall estimate for this voluntary availability of citizens for semi-structured horizon scan is 61500 (n = approximately 250 eld visits, every year, since 2012). While incorporating anonymity in all our previous reports 5, [15][16][17][18]45 , we have used iterated surveys, facilitated discussions, structured elicitation, and aggregation of individual perceptions and associations with animals to incorporate the full range of socio-cultural perspectives regarding opportunistic scavengers in the city. Through iterated interactions with new, random sets of respondents from the same stakeholder unit, we identi ed and mitigated typical individuallevel psychological biases 43 (Stage 1). This helped mitigate potential biases of people from the data, considering our respondents revered or vili ed myriad types of non-human animals, which contextualised their own professional or socio-cultural obligations. Out of the total estimated respondents, amongst 826 contributors (designated as decision makers with respect to human-animal interface), who were a liated with stakeholder groups likely to have higher encounters with avian scavengers on a regular basis, 5.7 % were a liated with academia and 5.5 % with NGOs (e.g. World Wide Fund for Nature, Wildlife Trust of India, etc.); 2.4 % were ritual feed-sellers (grains, fruits and meat-chunks); 9.3 % worked for the government (zoo, municipality, health workers) and 2.1 % for religious organizations (priests or Moulavis); and 75 % were associated with informal waste-work sector (butchers, chicken and sh market cleaners, rag pickers. municipality contract labourer, contractors). The major section of more than 60,000 contributors also included residents who immigrated to the city from multiple regions and shared their native biocultural perspectives.
In Stage 2, before every iteration step, the lists of responses were thematically organized to identify the relationships in socio-cultural rooting of the respondents and their professional background. Subsequently, further contributors to this study from randomly available respondents were chosen to balance background, expertise, and other parameters of urban social diversity. Team members, who were multilingual, helped create the necessary familiarity by initiating conversations in native dialects to build mutual trust to interview the urban immigrants from the nearby states (see below). Simultaneously, the team also scanned the immediate surroundings where the surveys were conducted to obtain a relative index for a. anthropogenic resources; b. the abundance of kites, crows, dogs, other commensals and livestock; and c. prosocial or agonistic human and non-human animal interactions in shared space (Stage 3; see 17 ).
Ultimately, before each successive step, the usefulness of horizon scanning can only be judged retrospectively 46 .
Therefore, subsequently, in Stage 4 (this study), the semi-structured and key-informant interviews we conducted to probe the social-ecological signi cance of urban opportunism by commensals were a surrogate for identifying major patterns in vulture-speci c perceptions amongst citizens, community groups (abattoir and waste-workers), NGOs, government agency o cials, and religious clerics. The interviews (n= 71) conducted by one of the authors (UG) in 2017 focused on common avian scavengers-kites, house crows and vultures through semi-structured conversations (details in Supplementary material 2). These species exploit anthropogenic food and have been studied to attain extremely high densities 2,15 . At this stage, interviewees were selected by snowball sampling, after stratifying the potential respondents into various stakeholder units. Efforts also incorporated sampling professionals involved in animal husbandry, and with other wild or feral animals in shared urban space or zoological parks 47 . All interviews for Stages 1 -4 were conducted by the authors in Hindi, other native languages (Bhojpuri, Maithili, Bengali, Assamese) or English, as deemed appropriate.
Further, we reworked nal issue descriptions and grouped them into overarching themes (details in Supplementary material 1, 2). This focused study allowed us to cross-validate these thematic groupings and links between (i) political ecology driven by individual-to-society feedback for people; and (ii) individual-to-population feedbacks that characterise urban ecology of opportunistic avian scavengers like vultures, shaping their functional ecology in human-dominated landscapes (Fig. 2).
To clarify the policy relevance of issues pertaining to ethnoecology of human-vulture interface, and potentially suggest re nement of vulture conservation breeding practices, we collated interviewees' perceptions to draw upon the issue-speci c impacts of vulture-loss for each stakeholder unit (Stage 5this study). We expected salience for avian scavengers to be driven by our subjects' socio-cultural, professional and economic backgrounds, corresponding to patch-level variations for socio-ecological systems. Inductive-opinions, embedded in folk-biology were expected to be accrued via direct encounters that shape ecological, aesthetic and corporeal charisma. Semistructured interviews enabled exibility to identify patterns in avian scavengers' salience that was contingent on the interactions expected between factors that affect social-ecological constructs-e.g., age, profession, gender, and domicile.

Ethics statement
Approval for the semi-structured interviews, as part of ethnography, was granted by the Central University Research Ethics Committee (CUREC), University of Oxford (reference: SOGE 17A-82) and by the Training, Research, and Academic Council (TRAC) of the Wildlife Institute of India, Dehradun (WII). We conducted all interviews in accordance with the relevant guidelines and regulations.

Results
A summary of salience and charisma for vultures and other avian scavengers The phenotypic and ecological salience for vultures were restricted amongst people older than 35 years-subjects not generally domiciled in top-tier cities-who encountered large scavenging-ocks in the past. Older males (>35 years) and subjects who were professionally associated with animals (municipality, zoological-park and NGOs working on animal welfare and conservation) shared direct experience, stories and knowledge about vultures and other avian scavengers (summarised in Supplementary materials1, 2). Similar to how multiple morphological cues are used by taxonomists to classify and name a species, folk names, perceptions and practices associated with avian scavengers depended on locally prevalent folktales, based on long-time coexistence. Cultural legends that explained avian scavengers' keen eyesight and ight capability re ected inductive opinions about the exaptation of morphological features enabling synurbization. The amalgamation of socially anthropomorphised approaches associated with the local political economy and biophysical ecology of commensals collectively extended the carrying capacity of urban ecosystems for selected scavengers (Fig. 3).
All anecdotes that mentioned direct vulture encounters were associated with village/locality-based carcass dumping areas carefully set aside from the residential blocks to harness their scavenging services (Fig. 1b). For rural systems, people informed that in the absence of vultures' services, carcasses are often dumped in open water-bodies (canals or rivers), buried underground with salt, or discarded in open (see below). The livestock carcasses in megacities like Delhi were collected by the municipal workers, but only from the designated 'dairy-colonies', e.g. at Ghazipur in Delhi. Thus, the absence of vultures' services in tending carcasses limited where all urban-poor could practice 'small-unit animal husbandry' in backyards, collectively stressing the perils of livestock maintenance in heterogeneously developed tropical cities. Their narratives (details in supplementary materials) had frequent mentions of competitive release impacts on stray dogs and rodents, underscoring heightened human-dog con icts and threats of disease spread.
Youngsters who never encountered vultures in wild, and/or female subjects who were less outdoorsy than male members of their families discussed the cultural salience of vultures, based on folktales, television and documentaries. Considering the sympatric opportunism exhibited by several avian scavengers for human-offal over accumulated garbage-an aspect more prominent at the sanitary land lls that currently support enormously large ocks of migratory black-eared kites Milvus migrans lineatus (n= 10,000 birds at Ghazipur: see 49 )-vernacular names for vultures and kites were used interchangeably (Supplementary material 2). Contrary to our expectations, immigrants who worked on land lls, tending waste in the proximity of multiple commensal species, re ected poorly on salience or charisma associated with avian scavengers. Given their socio-economic indulgences and poor connection with their native biocultural heritage, their indifference to mega-congregations of kites re ected the poor autonomy immigrant informal workers have over cultural expressions to nature within cities, where they settle as subalterns 39 . However, the prevalence of ritualistic animal feeding, and animal husbandry also provided livelihoods to another set of stakeholders amongst the immigrant urban poor in Delhi.

Discussion
This study elaborates on beliefs and perceptions about common avian scavengers amongst cultural mosaics represented by heterogeneously-developed tropical megacities like Delhi that comprise of multiple stakeholder units.
We contribute to advance the resolution of human-animal interactions, which are interdisciplinary and di cult to achieve via standalone ecological or ethnographic studies. Our results contribute folk-biological perspectives that are transdisciplinary and offer insights on how people ordinarily understand the biological world in rapidly urbanising ecosystems shared with opportunistic commensals. In cities like Delhi, millions of scavenging populations of myriad species live on waste and ritually offered food-subsidies by thousands of devout people who indulge in diverse practices, expressing patronage to animals. This is probably the largest human-animal interface in the world 16 . Human-animal interactions in tropical cities are distinct from their western counterparts 50 , driven by people acknowledging age-old ecosystem services by animals, potentially con guring regional urban nature-basedsolutions e.g. in solid waste disposal 16 . Cities within the Global South are conglomerates of immigrants that impact social traditions and practices at multiple scales 14 . Thus, dissociation of urban immigrants from their biocultural roots impacted their attitudes and ecological salience for non-human animals, even for frequently encountered avian scavengers 33,51,52 .
Unravelling ecology of opportunistic synurbic organisms to support their conservation will need distinction in how we characterise 'cities', which are con ned in physical geography. Urban systems, however, are not limited or de ned by political boundaries 32 . When characterising organismic ecology well entwined with the populous, we need to account for the co-production of materials, information (tangible and intangible archives), people, power relations, etc. -to establish forms of interconnection that most meaningfully constitute the urban for humans and nonhumans in shared spaces 53,54 . Unfortunately, as vulture-restoration efforts are being consolidated to achieve numerical response for a far greater quantity of human-offal, the eventual lag phase would not just be modulated by birds' life-history traits. Even the access to food-subsidies would be impacted by urban-development which entails increased power transmission lines, considering that vultures and other large birds like Great Indian Bustard are electrocuted by overhead power lines or windmill collision 7 . The functional response of vultures in human-use landscapes, unlike the small populations currently con ned to protected areas, would oversee newly co-produced ecosystems, contextualising ecological opportunism that ts the socio-cultural and socio-economic milieu. Based on the four domains of population, demography, behaviour and ethnoecology that we explore in more detail below contextualise urban scavengers and people in social-ecological systems 34,36,37,55 SES.
First, as an upper-trophic wide-ranging raptor, vultures fall under those avian groups that successfully colonised and thrived in human-dominated landscapes, attracted to the frequent allocation of anthropogenic resources. Raptor populations are typically limited by food and nest-sites 3 , wherein, the latter constrains the level of resource-tracking by individuals attuned to opportunism (e.g. 15 ). However, unlike the commensal-scale urban niche for kites and crows, vultures' services were resolutely harnessed by locals through the allocation of designated feeding sites -an erstwhile practice analogous to today's "vulture-restaurants'' 56 . It not only had implications for solid waste and disease management but also enabled the maintenance of the world's largest livestock population in povertystricken regions 7,8,57 . Till the last century, this vulture-mediated nature-based-solution for solid waste disposal simultaneously addressed economic, sustainability and socio-cultural goals-a regionally motivated feat incorporated by highly variable communities across South Asia 7,25 and elsewhere 10 . Therefore, for the Indian subcontinent's likely future, the largest milk-producing region of the world, we reckon to couple: (i) dairy-colonies' displacement as the urban gentri cation drive 58 ; and (ii) releasing vultures from conservation-breeding facilities 7,25 .
The latter would, of course, depend on careful, multi-stakeholder and multi-scalar planning to afford accessibility of safe feeding, roosting and nesting sites for vultures, whose availability could easily be exploited as a tool to simultaneously address urban waste and poverty 15,16 . For instance, north-western Delhi, where informal livestockrearing colonies' resettlement is planned 58 , lies in proximity to the Aravallis range that is currently an active, safe nesting zone (Authors' unpublished data). Correspondingly, policies pertaining to vulture restoration shall solicit simultaneous contributions from the federal and Non-Governmental Organisations (NGOs) that concern the environment and forests, animal husbandry practices and resettlement of the urban poor.
Second, the demographic shifts in vulture populations have accompanied changes in human population structure across the globe, especially in the Global South (Census Organisation of India, 2011; 41 ). Our surveys informed that migration to cities selectively de-couples younger generations from the joint-families, disrupting direct as well as oblique biocultural transmissions. Previously, these formed the basis for mutual tolerance 59 . Cities' physical urban sprawl and far-ranging urban impacts will likely affect successful vulture-reintroduction. Therefore, demographic repair for vultures should factor in cultural dynamics in humans to support re-attainment of mutually conducive niches in rapidly urbanising ecosystems 59,60 .
Third, while our subjects offered deep insights into how religious beliefs motivated them to patronise and tolerate vultures in localities, unfortunately, we have a poor understanding of how these birds dealt with the behavioural bottleneck that involved trade-offs in urban environments in terms of defending their young against humans, while exploiting anthropogenic food sources 45 . Recent studies on birds that occupy and exploit anthropogenic resources stress the factor of animal-personalities generated under the selection of individuals that could breed within the higher-end spectrum of adaptation to urban life 5 . But unlike multiple birds that have been studied to aggressively attack human beings to defend offspring, e.g. kites 17 , and magpies 61 , vultures are not known to attack humans 39,62,63 . Therefore, we speculate that vultures-populations breeding within cities were constituted by individuals that had synurbic personalities, developed via selection for tolerance to human proximity as a behavioural strategy 15,17 . Unfortunately, this urban opportunism was also the basis for ecological trap 64 that fuelled rapid decline of populations via consumption of cattle-carcasses which had diclofenac-residues in urban areas 7,25 . In terms of the erstwhile functional ecology of avian-scavengers that conjugated with animal-husbandry practices, behaviouralphenotype(s) 65,66 of the captive-bred vultures might preclude their settlement in human-use landscapes, rendering the small populations exposed to multiple threats at the currently altered human-wildlife interface 66-68 . Alternatively, urban opportunism on cattle carcasses can also be species-speci c tolerance of proximity to humans, selecting for White-rumped vultures Gyps bengalensis that constituted >80% of all vultures in South Asia before decline 7,25 . Collectively, based on aforementioned links in human and non-human agencies, changes to human-vulture interface have contributed to a long-drawn social-ecological trap for both agencies 69 . Given that media assisted in spread of anthropomorphic socio-cultural legends about vultures, as well as misinformation regarding what led to their loss and/or treatment as bad omens, socio-cultural shifts can erode patronising attitudes that previously allowed Indians to tolerate vultures 70 .
Fourth, we weave the aforementioned population, demographic and behavioural perspectives for the human and non-human animal agencies that co-produce and constitute the 'urban' in the Global South. Under the domains of SES, for cities under variably urbanised status, these constitute regional ethnoecology. Previous studies 71,72 , including a few in the region 5,45 , have shown that commensal species' functional ecologies are ascribed via centuries of coexistence, enabling metabolism of human offal by opportunistic species at multiple trophic levels.
For instance, food waste is a valuable subsidy to livestock and poultry owners in South-Asian cities and towns 16 .
Thereby, the urban ecology of the human-animal interface in urbanising systems is an unintended, informal selection of animal rearing practices. These ethnic-practices sustain and support millions of poor people, con guring domains of political economies and political ecologies with respect to which animals can cohabit in rapidly changing tropical SES 73 . Along with commensals, citizens in urban tropics share living spaces with livestock in the backyard, which consume edible waste as nutrient-rich feed. It reduces the costs of milk and meat production [74][75][76] . Scavengers like vultures, dogs, kites, rats and crows, thereafter, consume the organic remains, as human-mediated commensal agencies, with highly variable synurbic status 77 . Therefore, for the old world, the fastest declines witnessed in the vulture populations should not be treated as a single event isolated from simultaneously occurring changes in the human-wildlife interface and socio-economy. Such events are driven by, and, in turn, further cascades urban growth and development that shapes local culture, built environment and socialecological processes 74,78 . Meanwhile, tropical cities have witnessed massive increase in their solid waste since the vulture decline e.g. Delhi's waste grew by 300% since 2002 79 . Unlike their western counterparts, the ecology of human-nature interactions in tropical megacities 33 respond to the geography of human religion, hygiene and poverty via (i) interactions of socio-economic and socio-cultural processes amidst, (ii) rapid spatio-temporal alterations over availability and accessibility to food subsidies that are inextricably entwined with; (ii) population and behavioural dynamics that characterise functional urban ecology of non-human animals in human-dominated landscapes 36 (Fig. 4).
Further, vultures' loss has had its share in contributing to the burden of diseases that spill from animals to humans (zoonoses) 80 . Given that South Asia and Africa share the highest zoonotic burden, human-animal interactions within nite spaces have reportedly been associated with an increase in the population of stray dogs and other warm blooded commensals that cause rabies 80,81 . These commensals that share urban opportunism underwent competitive release in the absence of vultures 80 . The latter's resurrection will, therefore, be impacted by competition over anthropogenic resources, involving commensals' opportunism that is currently contingent on regional cultural geography 16 . Considering the perpetual increase in solid waste and its poor disposal in South Asia 48,82 that supports dense populations of free-ranging animals, successful seeding of captive-bred vultures will be challenging, considering the projected urbanisation of 500-million citizens over the next 30 years 30,35,36,41,83 .
Furthermore, a comparison of current precarity for vultures in Asia, Europe and the African continents (see Table 1) re ects the distinguishable socio-cultural as well as socio-economic factors for respective regional ethnoecological  A comparison matrix of current conservation threats, policy tools for vultures in Asia, Europe and the African continents, and the impacts of regional losses re ects the distinguishable socio-cultural as well as socioeconomic factors for respective regional ethnoecological practices. Adapted from CMS Raptor MOU Technical

Conclusion
In conclusion, social-ecological framework used in this study models the contingency of biodiversity conservation in the human-dominated landscapes on ecological as well as social, cultural and economic factors. The loss of vultures quintessentially depicts how limited social-ecological understanding delayed actions on the conservation and management of a once-common group of species. For poverty-stricken urbanising ecosystems of the developing world, the organismic criterion of habitat selection by opportunistic animals is contingent on foodaffording socio-economic drivers and cultural beliefs at the local to regional scales. Therefore, integrating human socio-cultural estimates while seeding individual animal units during restoration practices would affect populationlevel behavioural consequences and back 67,68 . In turn, expected progressive rationalisation of refuse disposal and detachment of younger generations from ritual practices and beliefs, which modulate tolerance for non-humans, make wildlife restoration a moving target within developing countries. Further, conservation practices that entail    misinformation about their local-extinction due to "technology-assisted" migration to Japan and USA. These socialecological disassociations represented by broken red lines impact the non-human charisma and salience involving