Keywords: emerging infectious disease, zoonotic risk, zoonotic spillover, SARS‐CoV‐2, coronavirus, pandemic prevention, wildlife trade

(1). Importance of zoonotic pathogens, including COVID‐19

The pandemic potential of zoonotic pathogens is strictly linked to their ability to generate sustained human‐to‐human transmissibility. This is the case with SARS‐CoV‐2 (the virus responsible for COVID‐19) which caused a global pandemic affecting over 200 countries and territories in under 100 days and resulting in over 175 million known infections and 3811561 assigned human deaths by 15th June 2021 [WHO Coronavirus Disease (COVID‐19) Dashboardcovid19.who.int]. Although there is debate about its exact source and infection pathway, COVID‐19 appears to have been the result of zoonotic transmission from an original wildlife host, possiblyvia an intermediate animal host, following close contact with people (Andersenet al., 2020; Wuet al., 2020). While the direct reservoir of SARS‐CoV‐2 might never be identified, it is clear that close proximity of different wild and domestic animal species in a wildlife market setting (often conflated with ‘wet market’, which may, or may not, have wildlife and simply refers to the existence of fresh produce) may enable recombination between more distant coronaviruses and the emergence of recombinants with novel phenotypes (Liet al., 2020). This is particularly relevant given that multiple relatives of SARS‐CoV‐2 and SARS‐CoV (the cause of the 2003 SARS epidemic) circulate in wildlife species in Southeast Asia and southern China (Zhouet al., 2021). Preventing such situations as well as reducing direct human contact with wild animals appears critical for preventing new coronavirus zoonoses.

Several other major recent pandemic and epidemic disease outbreaks have zoonotic origins including HIV‐AIDS, Ebola and SARS (Wang & Eaton, 2007; Allenet al., 2017); 60% of human emerging disease events are caused by zoonotic pathogens, with most (72%) originating in wildlife (Joneset al., 2008). Furthermore, most known human pathogens are zoonotic (80% of viruses, 50% of bacteria, 40% of fungi, 70% of protozoa and 95% of helminths) (Taylor, Latham & Woolhouse, 2001). Although most zoonotic pathogens are not capable of sustained human‐to‐human transmission, some can cause major disease outbreaks; thus, preventing the transfer of pathogens from other animal species into humans is a key societal challenge. Predicting and reducing the risk of such outbreaks is imperative if we are to avoid future detrimental impacts on human health and the global economy, such as those caused by the COVID‐19 pandemic.

(2). Disease transfer pathways

Transfer of pathogens from animals to humans can occur through diverse pathways involving interactions with free‐living or captive wildlife, livestock or other domesticated animals (Table 1; Fig. 1). It is important to note that, in practice, the distinctions between these categories of animals are often poorly understood and category differentiation may be imprecise but differences have significant implications for exposure to various pathogens.

The rate of zoonotic pathogen emergence is increasing globally, and human population density is a strong predictor of emerging disease events (Joneset al., 2008), indicating that pathogen emergence is driven by human‐induced changes bringing wildlife, livestock and humans into closer and more frequent contact (Morseet al., 2012). The risk of disease transmission depends on both intrinsic factors (e.g. pathogen life history, host availability and immunity and transmission route) and external factors (e.g. land‐use, human population changes, socio‐economic provisions) such that the nature of risks varies among locations and over time (Beckeret al., 2020), and influences both the probability of pathogen transmission from animal to human, and the probability of an infected person developing the disease (Han, Kramer & Drake, 2016).

The spillover of a pathogen from animals requires a series of stages, including the reservoir host being at sufficient density to retain the pathogen, pathogen release, human exposure to the pathogen, and the pathogen overcoming structural barriers, innate immune responses and molecular compatibility (Plowrightet al., 2017). The global connectivity of human society greatly increases the movement of humans, disease vectors (Tatem, Hay & Rogers, 2006) and various pathogen‐infected animals (Can, D'Cruze & Macdonald,2019), magnifying the likelihood of the spillover and spread of a pathogen, particularly in areas of high human population size and density.

(3). Disease transfer involving wild animals

Most emerging infectious diseases are thought to have originated in wild animals, especially non‐human primates, rodents and bats (Wolfe, Dunavan & Diamond, 2007; Joneset al., 2008; Hanet al., 2016), although many of these were transmitted to humansvia intermediate hosts, such as companion, farmed or feral animals (Wolfeet al., 2007). As an example, there is a broad consensus that human viruses responsible for HIV‐AIDS resulted from multiple cross‐species spillovers of simian immunodeficiency viruses involving the chimpanzeePan troglodytes, westerngorilla Gorilla gorilla and sooty mangabeyCercocebus atys. Lentiviruses, such as HIV, penetrate mucous membranes; therefore contact with non‐human primate body fluids associated with the hunting, butchering and consumption of animals likely led to spillovers. One transmission event, probably occurring between 1910 and 1930, gave rise to the HIV strain behind pandemic AIDS (Sharp & Hahn, 2011). Ebola has been suggested to have been transmitted from bats (Leroyet al., 2005; Saézet al., 2015), either directly orvia an intermediate host, while MERS (Middle‐East Respiratory Syndrome) most probably originated in a species of bat, with the dromedary camelCamelus dromedarius as an intermediate host (Mohd, Al‐Tawfiq & Memish, 2016).

Our knowledge of pathogen prevalence in wildlife populations is heavily biased by host species, pathogen type and sample availability, but studies can provide deep insights into the diversity of potential zoonotic pathogens (e.g. for bat‐borne coronaviruses; Anthonyet al., 2017). Species are defined as being capable of harbouring a particular zoonotic pathogen following the detection of that pathogen, but only pathogens that can be reliably detected and identified are recognised. In the natural host, however, infection loads may be low and not readily detectable even with modern molecular methods; testing might also be conducted on samples that are suboptimal for certain viruses that then could be missed. Furthermore, most host taxa have not been included in such studies and remain completely untested. Fewer than 300 viruses from 25 high‐risk viral families are known to infect people, yet it is estimated that there are around 1.7 million viruses from these same viral families that have not yet been discovered in mammals and birds. Of these, some 700000 are considered to have zoonotic potential (Carrollet al., 2018).

Zoonotic diseases can emerge from a wide variety of wild species, including marine turtles (Aguirreet al., 2006) and marine mammals (Waltzeket al., 2012), but the risk appears variable, with the highest risk taxa being rodents (Hanet al., 2015,2016), non‐human primates (Pedersenet al., 2005) and bats (Luiset al., 2013). Transmission in this context can occur through a range of direct and indirect pathways (e.g. infected faeces, urine, saliva, invertebrate vectors), from interactions with species in natural habitats or during the supply, transport and use of wildlife or wildlife products.

Based on the number of different infected host species, as well as phylogenetic relatedness among hosts, zoonotic pathogens can be characterised as specialists (a single wildlife host species) or generalists (multiple wildlife host species). The latter often can persist by being maintained in multiple wildlife species. Understanding pathogen ecology and evolution offers many advantages in terms of strengthening surveillance programs aimed at preventing or reducing human exposure and zoonotic infection, as well as informing early warning systems for outbreak detection.

Many host species characteristics contribute to a heightened risk of zoonotic transfer. Several zoonoses of high impact for humans originated from non‐human primates, probably in part because the phylogenetic barrier to transmission to humans is low (Wolfeet al., 2007). In rodents, reservoir species are associated with a fast‐paced life‐history strategy, rapid maturation and high fecundity (Hanet al., 2015); these characteristics mean that some species carry multiple pathogens, increasing the probability of a rodent being infected with a pathogen with zoonotic potential. The high population size and density of many rodents is probably also an important factor. It is hypothesised that pathogen‐prone rodent species may have low investment in immune defences, but ‘outrun’ the risk of a lethal infection by producing offspring quickly (Hanet al., 2015).

The comparatively high zoonotic disease risk from bats (Luiset al., 2013) has been linked to large colony sizes and high mobility due to flight, resulting in even larger effective population sizes and many bat species having the capacity to host a wide range of pathogens. The highest predictor of zoonotic viral richness in bats was the distribution overlap with sympatric species, suggesting that interspecific transmission probably plays a key role in the pathogen complement harboured by bats (Luiset al., 2013). Evolved physiological adaptations to flight might also have enabled bats to harbour a larger range of pathogens in the absence of disease (O'Sheaet al., 2014), as their immune systems seem to have been modified to enable protective cellular mechanisms, a dampened interferon response, and a key innate defence pathway that is functionally different from non‐bat mammals, implying that bats may be more effective at co‐existing with a large number of viruses (Xieet al., 2018). The rigours of flight mean that bat metabolisms can increase by up to 16 times the basal rate, producing sufficient heat to kill most mammals (Speakman & Thomas, 2003; O'Sheaet al., 2014). As a consequence of these costs, bats have adapted mechanisms to prevent degradation of cellular mechanisms by heat or oxygen radicals, affecting their ability to withstand infections (Healyet al., 2014; Huanget al., 2019). Together, these properties allow them to be asymptomatic reservoirs of multiple viruses, thus enabling these viruses to persist within bat populations. Stress responses in most mammals are shown to increase the probability of spillover (Haraet al., 2011), yet such studies in bats are only now being conducted (Subudhi, Rapin & Misra, 2019). Understanding how the role of habitat loss and degradation combines with natural stressors (e.g. reproduction and migration) is urgently needed. However, studies show that at least some mechanisms of physical stress in bats do increase viral shedding; for example, bats infected with the fungal pathogenPseudogymnoascus destructans can have a viral load that is increased by up to 60 times that of bats without this fungal infection (Davyet al., 2018).

In addition to these hypotheses, a simpler explanation is that both rodents and bats are also highly diverse vertebrate orders (2361 rodent species and 1420 bat species) and more species‐rich reservoir groups host more virus species and therefore a larger number of zoonotic pathogens (Letkoet al., 2020; Mollentze & Streicker, 2020).

Other, non‐taxon‐specific characteristics also contribute to zoonotic pathogen risk. For example, migratory species can have a profound effect on pathogen dispersal, but these effects are complex and context dependent (Altizer, Bartel & Han, 2011; Poulin & de Angeli Dutra, 2021). Migration can play a key role in introducing disease to populations naïve to the pathogen, and the heavy toll of migration can reduce immune function, so increasing infection burden. Conversely, migration can allow individuals to escape infected areas and, hence, reduce host population pathogen levels. The unsuccessful migration of infected individuals might also lead to overall reductions in pathogen prevalence (Huberet al., 2020).

Where pathogens are thought to be of wildlife origin, their emergence is often associated with a high diversity of pathogens in a wide range of host species (Joneset al., 2008; Allenet al., 2017; Anthonyet al., 2017), and many emerge in biodiverse tropical regions. However, the role of biodiversity in zoonotic disease emergence is complex. Studies have identified a general trend, known as the ‘dilution effect’, where increasing host diversity can reduce a given parasite abundance in both wild animals and humans (Civitelloet al., 2015; Huanget al., 2017), although this effect depends on specific conditions, context and on the metrics used (Salkeld, Padgett & Jones, 2013; Roberts & Heesterbeek, 2018). While empirical evidence exists for the dilution effect in several multi‐host pathogen systems, the mechanism is often unclear, for example whether it is due to actual dilution or to reduced host density (Begon, 2008).

Pathogen transmission from wildlife to humans is influenced by extrinsic factors, such as land‐use change (Allenet al., 2017) and agricultural intensification (Joneset al., 2013). Such factors play a particularly important role in driving the emergence of zoonotic diseases in biodiverse tropical forest regions (Keesinget al., 2010), where expanding human populations (and associated agriculture or other activities) into natural habitats leads to increased opportunities for human‐to‐wildlife contact (Hanet al., 2016; Bloomfield, McIntosh & Lambin, 2020) and increased pathogen transmission at human–livestock–wildlife interfaces (Gebreyeset al., 2014). Multi‐host pathogen models indicate that pathogen transmission between species inhabiting intact and converted habitat is highest when rates of habitat conversion are intermediate (Johnsonet al., 2020). However, the potential severity of epidemics increases at higher rates of habitat conversion (Faustet al., 2018). Land‐use change that reduces local biodiversity may increase spillover risk (Civitelloet al., 2015), as exemplified by Lyme disease in North America (Ostfeld & Keesing, 2000,2012). While the diversity of wildlife species declines with habitat degradation, it has been shown that those species more able to live in human‐modified habitat tend to have a higher rate of carriage of zoonotic pathogens than those that decline or disappear (Gibbet al., 2020).

Hunting, whether commercially or for subsistence, and the transport, sale, preparation and use of wild animals and wild animal products are also important points of human–wildlife contact. Direct wildlife use, especially for human consumption, is most commonly an issue for zoonotic emergence in the ‘paleotropics’, where high species diversity of high‐risk taxa (e.g. of bats and primates) is combined with subsistence hunting and use (Hanet al., 2016). Harvesting rates of wild meat in tropical areas are currently primarily driven by an increase in demand in fast‐growing urban centres (Coadet al., 2019). Interactions between wildlife and humans can have complex and hard‐to‐predict effects, by increasing both stress and movement of species and therefore increased spread of disease and spillover risk. For example, the culling of badgersMeles meles is known to increase the movement of surviving animals and, hence, the spread of bovine tuberculosis (Woodroffeet al., 2006).

It seems likely that it is human interactions with, and destruction of, biodiversity that leads to increased likelihood of zoonotic disease emergence. However, the pathways to overcome this are likely to be complex. It has been suggested that protecting ecosystems not currently posing a major threat of disease to humans or wildlife might prevent increases in disease emergence, yet when managing a specific disease for which the ecology is reasonably well understood, it might be more effective to manage the particular species (vectors or amplifying or diluting hosts) or habitats that are known to decrease or increase the likelihood of pathogen spillover, for example, through vaccination, culling, predator supplementation or habitat manipulation (Rohret al., 2019).

(4). Disease transfer involving domestic animals

Many zoonotic outbreaks result from pathogen transmission from domestic animals. Human pathogens considered to originate from the domestication process of animals include diphtheria, influenza A, measles, mumps, pertussis, rotavirus, and smallpox (Wolfeet al., 2007) and contacts between humans and domestic animals have led to recent zoonotic emergence events, such as the H1N1 (Swine Flu) pandemic in 2009 and MERS in 2012. The emergence of many of these diseases has been facilitated by the increased human population size and the development of commercial agriculture and livestock domestication (Wolfeet al., 2007; Joneset al., 2013), as well as agricultural encroachment leading to increased livestock–wildlife interactions and more opportunities for livestock acting as bridging species for novel zoonotic pathogens such as Nipah (Pulliamet al., 2011). Due to thousands of years of domestication, high contact rates and significant amounts of study, ungulates are the mammalian group with which humans are known to share the most pathogens (Cleaveland, Laurenson & Taylor, 2001). In temperate regions, the majority of the heaviest‐burden zoonotic diseases impact humans through domestic livestock. Transmission routes of pathogens from livestock are facilitated by poor hygiene and biosecurity measures, such as lack of protective equipment for farm workers (Ramirezet al., 2006), and can occur through a variety of direct and indirect interactions during the rearing of livestock. Numerous other zoonotic diseases involve a vector stage, such as Rift Valley fever or Crimean‐Congo haemorrhagic fever (CCHF), where the host is a tick and the vector is a mammal (Wilsonet al., 1991).

(5). Disease transfer involving wild or exotic pets

Although information is comparatively scarce, wild or exotic pets (i.e. not domesticated animal pets) are another possible source of novel zoonoses with epidemic potential. For example, in 2003 six states in the USA experienced an outbreak of monkeypox, the first cases of human monkeypox reported outside the African continent. This outbreak resulted from prairie dogsCynomys sp. sold as pets after being housed in close proximity to infected rodents recently imported from Ghana (Centers for Disease Control and Prevention, 2003). In 1999, an Egyptian rousette batRousettus aegyptiacus sold in a pet shop in France was diagnosed with Lagos bat lyssavirus encephalitis, resulting in the treatment of 120 exposed persons (Chomel, Belotto & Meslin, 2007). Variegated squirrelsSciurus variegatoides imported from Latin America as exotic pets were the identified host of a novel zoonotic Bornavirus (VSBV‐1) that infected, and led to fatal progressive encephalitis or meningoencephalitis, in three squirrel breeders in Germany in 2011–2013 (Hoffmannet al., 2015).

Some actions to combat zoonotic disease depend on an understanding of links between animal welfare and pathogen transmission, including the immune response, which is directly influenced by welfare (Broom & Fraser, 2015). For example, poor welfare during the transport of cattle or sheep can result in the opportunistically pathogenic bacteriumMannheimia (Pasteurella)haemolytica causing disease (Broom & Kirkden, 2004). Disease and mortality rates are higher in farm animals that have poor levels of welfare, and in wild animals if stressed when brought into captivity (EFSA, 2006; Ledayet al., 2018). If wild animals are captured and kept, capture stress as well as transport stress and other stressors (e.g. being caged with or next to conspecifics or other species; close proximity to people; rough handling; inadequate food or water; exposure to the elements; poor hygiene, etc.) increases their susceptibility to infection with pathogens and the likelihood that they will shed pathogens with or without the development of clinical disease, perhaps thereby infecting humans or other animals (Broom & Kirkden, 2004; Broom & Johnson, 2019).

(6). State of knowledge on coronaviruses – origins and transfer

Seven coronaviruses (CoVs) are currently known to infect humans (Andersenet al., 2020), four of which are regularly found in human populations in which they cause only mild symptoms (Cormanet al., 2018). However, the betacoronaviruses, SARS‐CoV‐1, MERS and SARS‐CoV‐2, have been associated with fatalities and, in the case of MERS, with high fatality rates (Zhouet al., 2020b). There is evidence that CoVs that infect humans have their origins in either bat or rodent species (Salataet al., 2019). Intermediate species have included both domestic and captive wild animals (including cattle and swine), masked palm civetsPaguma larvata and dromedaries (Drexler, Corman & Drosten, 2014; Cormanet al., 2018). Diverse CoVs have been found in bats in China, with 6.5% of all bats in one study testing positive for at least one coronavirus (Tanget al., 2006), and with SARs‐like viruses confirmed in horseshoe batsRhinolophus spp. in 2005 (Liet al., 2005). Since then, further betacoronaviruses have been detected in wild‐caught rhinolophid bats across the Old World (Gouilhet al., 2011). Over 200 novel coronaviruses have been identified in bats and approximately 35% of the sequenced bat virome is composed of coronaviruses (Banerjeeet al., 2019).

Although many CoVs are limited to bats, some are found in a more diverse selection of mammals. Coronavirus studies have predominantly focused on non‐bat hosts, which include both mammals (alpha and betacoronavirus) and birds (gamma and deltacoronavirus), yet few betacoronaviruses have been detected outside bats in the wild. In bats, CoVs have been found in bat families across the globe (Drexleret al., 2014), with groups such as Hipposideridae known to host these viruses asymptomatically for extended periods. Typically, CoVs have a highly restricted host range: even bats within the same cave often show different viruses in different species, with only those detected inMiniopterus spp. known to be capable of jumping between hosts (Gouilhet al., 2011), normally as a consequence of roosting in direct physical contact with other bat species. The destruction or disturbance of caves by people can lead to different species and taxa being forced to share caves and, given their different abilities to host or pass on betacoronaviruses, this increases the risk of virus spread across species. Furthermore, as rhinolophid bats (in which SARS‐like viruses have been detected) can develop clinical coronaviral disease, spillover risk may be higher from this group (Wonget al., 2019). The shared use of bat caves by non‐bat hosts, such as Viverridae (civets and genets), increases the risk of spillover into new host species (Songet al., 2005), especially as betacoronaviruses are most commonly detected in faeces (Wonget al., 2019). Rapid adaptation of SARS‐CoVs has been observed to occur between hosts (such as masked palm civets and humans) and host shifts have also been observed to occur amongst bat species in wild populations (Cuiet al., 2007; Latinneet al., 2020; Zhouet al., 2020a).

The wildlife trade, both legal and illegal, has received enormous global media attention for its potential role in the emergence of novel zoonoses like SARS and COVID‐19. Illegal wildlife trade alone is estimated to be a multibillion‐dollar industry, comparable to the international trades in narcotics and weapons, and which raises significant human and animal health concerns, especially given its volume, the complete lack of regulation and the fact that the origins of wildlife sometimes match recognised emerging infectious disease hotspots (Smithet al., 2017). However, there remains considerable uncertainty around which species have been involved in the transfer of the causative coronaviruses to humans, as well as exactly when such transfers occurred (Cormanet al., 2018; Salataet al., 2019). The virus with the closest match to SARS‐CoV‐2 has been found in several species of horseshoe batRhinolophus spp., which likely represent the ancestral or evolutionary (natural) host of the virus (Zhouet al., 2020a), although the ability of this closest known relative of SARS‐CoV‐2 to infect humans is poor (Wanet al., 2020). A related virus has been detected in Malayan pangolinManis javanica (Zhang, Wu & Zhang, 2020). While different overall from any known bat CoV, the pangolin CoV receptor binding domain part of the spike protein (which allows the virus to infect a new host) is almost identical to that found in the human virus (Zhouet al., 2020a), but a polybasic cleavage site in SARS‐CoV‐2 is absent from both known pangolin and bat CoVs, so the origin and route to human infection of SARS‐CoV‐2 remains unknown (Andersenet al., 2020). The potential for the transmission of zoonotic CoVs through wildlife markets and farm systems, however, has been clearly noted for SARS‐CoV (Wonget al., 2019) and actions to prevent such zoonotic spillover risk in future have been widely discussed (e.g. Kellyet al., 2020; Nabiet al., 2020; Ribeiroet al., 2020; Wanget al., 2020).

(1). The need for solution scanning

Creating a future in which society is more resilient to zoonotic diseases will require coordination and planning among different professionals, considering a broad range of prevention options related to wild, feral and domestic animals, all of which have potential to be the source of future epidemics in humans. These will range from medical and veterinary interventions to simple behavioural and societal interventions that could greatly reduce the risk of pathogen transfer (Morseet al., 2012). Solution scanning uses published research and guidance, the experience of experts and practitioners, and brainstorming to identify a range of potential solutions to a specific problem (Sutherlandet al., 2014). This includes solutions that have not been reviewed, might not have evidence of effectiveness or may indeed be ineffective, inconvenient, controversial or have negative side effects. It is therefore important that practitioners and policy makers identify possible interventions as a starting point in decision‐making before evidence synthesis and consideration of each option's advantages and disadvantages are conducted. The need for this is illustrated by previous research that showed that 92 conservation practitioners responsible for addressing a problem were only aware of 57% of the possible actions (Walsh, Dicks & Sutherland, 2015).

We conducted a solution scan to identify options for reducing the animal–human transfer of zoonotic pathogens with high potential for human‐to‐human transmission, such as COVID‐19. The solution scan was compiled by documenting our own experience of actions, consulting guidance and scientific literature, and contacting a number of experts and practitioners working in different countries, contexts and institutions in order to explore the range of options available. The approach is based on methods developed as part of subject‐wide evidence synthesis (Sutherlandet al., 2020). This solutions scan was initiated as a collaboration between BioRISC (the Biosecurity Research Initiative at St Catharine's College, Cambridge), Conservation Evidence based in the Department of Zoology, University of Cambridge, and numerous other researchers worldwide. The options considered here are diverse, reflecting the variety of possible transmission pathways at the wildlife–livestock–human interface that are discussed in Section II from wildlife, domesticated and captive animals (Table 1). Our review excludes other sources of potential zoonotic disease emergence which could be responsible for future pandemics including (i) the evolution of antimicrobial resistance (AMR), (ii) pathogen release from laboratories, or (iii) the intentional creation of life. It is important to note that AMR is a topic of enormous importance: AMR could be responsible for 10 million deaths per year by 2050 if left unchecked (O'Neill, 2014). However, we exclude AMR from this study due to differences in the type of pathways of disease emergence, including pathogen selection in effluent pollution and the overuse and misuse of antibiotics in farming and medical settings. Nevertheless, many of the animal husbandry options considered herein, in particular those related to improvements in hygiene and health standards, are directly relevant to enabling reduced use of antimicrobials on farms and thus avoiding the selection of resistant strains.

(2). Interpreting the list of options

We stress that the list provided below is a list of options for consideration and testing, not a list of recommendations or prescriptions. Many options listed may not be feasible, practical or affordable in some situations. For example, the options available to small‐scale subsistence farmers will differ from those available to large‐scale commercial farms. Similarly, options available in countries with poor animal health and hygiene infrastructure, weak governance and low capacity to regulate or control the local wild meat trade, international wildlife trade, and few available medical testing facilities, will differ from those available to countries with state‐of‐the‐art facilities, diagnostic equipment or strict law enforcement.

It should be recognised that solution scanning is a dynamic process which means that the current list should not be viewed as exhaustive. While effort has been made to compile as complete a list as possible using expert opinion and international collaboration, the vast literature available on specific livestock groups has not been systematically searched, and there may be further options available. This is also true for the wildlife trade literature given the diversity of ways in which animals are used across the globe (e.g. wild meat, exotic pets, medicines, curios, medical research, etc.). Additions to the list are welcome and can be sent tobiorisc@caths.cam.ac.uk, and will be added to an updated list of options available athttps://covid-19.biorisc.com.

(1). Supply side

Supply‐side measures are any that are applied to the production or sourcing of animals (i.e. rearing of farmed wildlife or livestock or hunting – endeavouring to kill or capture wild animals). They focus on preventing or reducing production, or altering the production process to reduce risk (’tSas‐Rolfeset al., 2019).

Supply‐side interventions to prevent zoonotic emergence from wildlife may focus on: (i) entirely preventing hunting and collection of high‐risk species (area‐based or species‐based restrictions); (ii) controlling the rate of hunting and collection through limits to numbers or specific characteristics of the animals taken; or (iii) regulating hunting, consolidation and trade through enforced standards. Preventing or reducing hunting, collecting and disturbance of wild species, especially of high‐risk species, should decrease the transmission of zoonotic diseases to livestock and people (Johnsonet al., 2020). For example, it can reduce the rate of contact between hunters and animals in the wild, and therefore reduce direct animal‐to‐human transmission. It can also reduce the movement of wild animals out of natural habitats, to places where they have more contact with people and other animals (e.g. markets and other vendors) (Swiftet al., 2007). For species that are lower risk for direct transmission, regulated harvesting, with licensing, standards and health or hygiene checks at critical control points, could reduce high‐risk practices that cause transmission (e.g. unhygienic slaughter, interaction of domestic and non‐domestic animals), and in some cases may be more effective than complete bans (e.g. Fourniéet al., 2013).

For livestock, supply‐side interventions to prevent zoonotic emergence may focus on: (i) improving animal welfare, health and immune function in farms by limiting density, ensuring veterinary care and reducing long‐distance transport or practices that involve large‐scale and long‐distance live animal movements (e.g. by promoting ranching instead of nomadism); (ii) preventing the mixing of domesticated and wild‐sourced or farmed wildlife animals in farms; or (iii) improving the biosecurity of farming practices (e.g.via the use of disposable protective clothing and introduction of health surveillance for farm workers as well as controlled slaughter and hygiene conditions).

(2). Transport and sale

Transactional interventions focus on any points in the supply chain between source and consumer, for the transport or sale of animals and their products. Broadly, transactional interventions involve regulation and legislative control (e.g.https://www.nfacc.ca/codes-of-practice), and tracking and detection of products as they move through the supply chain, and these can be applied during transit or sale. Here we present a series of options varying in the strength of restrictions.

(3). Consumption

Demand‐side interventions can reduce the frequency or the quantity of a given high‐risk product being used or shift demand away from high‐risk products and towards lower risk ones (UNEP, 2019). These interventions can take the form of either coercive measures, such as laws or law enforcement aimed at dictating consumer behaviour, or closer engagement with consumers to identify interventions that promote voluntary behaviour change (’tSas‐Rolfeset al., 2019). Demand‐side measures can target consumers directly or indirectly, through the use of key influencers such as peers, doctors, or religious leaders (’tSas‐Rolfeset al., 2019).

(4). Enabling environment (applies to all)

(1). Using options to make decisions

COVID‐19 has demonstrated the rapid capacity of a virus to transform human lives globally. In an interconnected world such events cause mass mortality and overload health systems, yet without systemic change the rate of emergence of zoonotic diseases with pandemic potential will continue to increase and we need to learn lessons not only on how to moderate and control pandemics post‐emergence, but to minimise the risk of their emergence by reducing the likelihood of animal–human disease spillover. To cause spillover infections in humans, all zoonotic pathogens must overcome a hierarchical series of barriers. Understanding and targeting these barriers (e.g.via integrating studies of animal ecology, landscape ecology, microbiology, disease ecology, immunology and others) could substantially improve our ability to predict or prevent spillover events, for instance by reducing human exposure (Plowrightet al., 2017).

We herein present a list of options, not conservation or sustainability focused, but specifically to achieve a reduced risk of disease transmission from animals to humans. These are not reactive strategies for use during a disease outbreak, but a set of proactive options to change the way society currently operates and to prevent disease emergence in humans at the outset. These options can be used in the design of strategies to minimise risks of zoonotic disease transfer by inter‐governmental, governmental, corporate or non‐governmental organisations or individuals. We emphasise that our solution scan does not provide a list of recommendations or prescriptions. It instead provides a starting point for discussion and analysis of the available options for intervention with different decision‐making and regulation bodies at global, regional, national and local scales.

When deciding on which measures should be adopted, policy‐makers and practitioners should first determine the context, the pathways of zoonotic disease transfer to be addressed and the risks associated with them. This will then allow relevant and practical options to be prioritised, assessed and tested. Once this has occurred, decisions can be made on which interventions to deploy, feeding into strategies to reduce the risks of zoonotic disease transfer.

(2). High‐level options currently proposed in the response to COVID‐19

There have been numerous calls for radical changes, primarily for complete bans on hunting, wildlife trade, wildlife trade for consumption or so‐called ‘wet markets’. Such calls may sound reasonable but miss the important roles of fisheries and sustainable products, wildlife's contribution to global diets and that wet markets are an important source of affordable fresh produce, where wildlife generally has a minor role (Ribeiroet al., 2020; Wanget al., 2020).

Whilst acknowledging the need and appetite for change, we believe such broad measures are unlikely to be enacted or obeyed. The solution seems likely to be a broader, more pragmatic and realistic approach that considers the variety of disease pathways, the different mechanisms by which interventions can act, the complexity of options that can target such mechanisms and the likely on‐ and off‐target effects of those actions. This solution scan can help in the early stages of designing such approaches, presenting a wide range of options, including stronger enforcement on illegal wildlife trade and bans on high‐risk wildlife trade, which can be considered to tackle the transfer of emerging zoonotic diseases with epidemic potential into humans.

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