Current unprecedented declines in biodiversity reduce the capacity of ecological areas to carry out many fundamental ecomechanism services. Here we evaluate evidence that lessened biodiversity affects the transmission of transmittable conditions of people, other pets and also plants. In principle, loss of biodiversity might either boost or decrease disease transmission. However before, mounting proof indicates that biodiversity loss typically rises disease transmission. In comparison, locations of normally high biodiversity might serve as a resource pool for brand-new pathogens. Overall, despite many kind of remaining inquiries, present evidence suggests that keeping undamaged ecodevices and also their endemic biodiversity need to mostly mitigate the ubiquity of contagious illness.
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In June 2010, a brand-new organization, the Intergovernpsychological Science-Policy Platdevelop on Biodiversity and Ecodevice Services (IPBES)—patterned after the Intergovernpsychological Panel on Climate Change (IPCC)—was established to assess changes to the diversity of life on the Planet and just how these alters will certainly impact human well-being1. Human well-being would certainly be adversely influenced by biodiversity losses if ecosystems with diminished biodiversity are less able to carry out the ecodevice services—such as carbon sequestration, nutrient cycling and also resistance to drought—on which humans rely. In current years, a consensus has actually emerged that ecodevice features decrease as biodiversity is lost2. Here we research how biodiversity affects the transmission and appearance of contagious illness and also evaluate the proof that diminished disease transmission is a vital ecomechanism service provided by high biodiversity.
Biodiversity includes the diversity of genes, species and also ecodevices. Increases in huguy populaces have led to an unprecedented and precipitous loss of biodiversity3. Current extinction prices are estimated to be at leastern 100–1,000 times background extinction prices and future extinction rates (over the following 50 years) are approximated to be 10 to 100 times current extinction rates3. A big proportion of species in all assessed taxa are currently intimidated through extinction (12% of birds, 23% of mammals, 32% of amphibians; 31% of gymnosperms; 33% of corals4) and also the ideal estimate of population fads of birds, mammals, amphibians, reptiles and fish suggests that considering that 1970 worldwide population sizes have declined by almost 30% (ref. 5). Global and also neighborhood extinction rates of some taxa, specifically microbes, have not been well defined. For the many type of organisms that are symbionts of other organisms, extinction of their hosts deserve to reason their extinction too6. Collectively, these declines and extinctions are resulted in by transforming the Earth’s ecounits to accomplish flourishing needs for food, fresh water, fibre, wood and also fuel, and also by climate readjust.
Changes in biodiversity have actually the potential to impact the risk of contagious illness exposure in plants and also animals—including humans—bereason contagious conditions by definition involve interactions among species. At a minimum, these species include a host and a pathogen; regularly many kind of even more species are connected, including added hosts, vectors and also various other organisms through which these species connect. Intriguingly, biodiversity might play a double function in the appearance and transmission of transmittable illness. On the one hand, high biodiversity may carry out a larger potential resource of novel pathogens, but on the other hand also, biodiversity deserve to minimize further pathogen transmission for both long-establiburned and also newly arising illness. We initially review the results of biodiversity on the transmission of establiburned illness and also then turn to illness introduction.
Transmission of pathogens in between species
Biodiversity loss can influence condition transmission through a number of mechanisms (Box 1). If the result of each species on pathogen transmission were totally idiosyncratic, one would certainly mean that diversity declines would certainly be equally most likely to reason a decrease or a boost in condition transmission in the continuing to be species. However, in recent years, a regular picture has actually emerged—biodiversity loss often tends to boost pathogen transmission and also condition incidence. This pattern occurs across environmental units that vary in kind of pathogen, hold, ecosystem and also transmission mode (Table 1). As an instance, West Nile virus is a mosquito-transmitted virus for which a number of species of passerine birds act as hosts. Three recent research studies detected solid correlationships between low bird diversity and boosted humale hazard or incidence of West Nile encephalitis in the United States7,8,9. Communities via low avian diversity tfinish to be overcame by species that amplify the virus, inducing high infection ubiquity in mosquitoes and people, while areas through high avian diversity contain many species that are much less experienced hosts. For hantavirus pulmonary syndrome, a straight transmitted zoonotic condition, correlational and also speculative studies have actually shown that a lower diversity of tiny mammals rises the ubiquity of hantavirprovides in their hosts, thereby increasing threat to human beings (Box 2). Diversity has actually a comparable result for plant diseases, with species losses increasing the transmission of 2 fungal rust pathogens that infect perennial rye grass and various other plant species10.
Recent attention has concentrated on assessing the mechanisms by which lessened biodiversity rises pathogen transmission (Box 1). Biodiversity loss can plainly increase transmission if it reduces predation and competition on reservoir hosts, thereby increasing their density. However, dispute has actually centred roughly whether the loss of species ca boost transmission in other ways11. This is bereason area researches prefer those on West Nile virus, hantaviroffers and also rye grass have typically not controlled for alters in organize density that deserve to outcome from alters in ‘species richness’ (the number of species present in a community, which is a measure of taxonomic diversity). As a repercussion, it has been hard to sepaprice the results of greater thickness from those of reduced diversity. Recent experiments confirm that boosts in disease transmission deserve to happen as soon as species richness declines also if host thickness remains consistent. One of the finest examples originates from a examine of Schistosoma mansoni, a trematode that causes schistosomiasis in people. The parasite alternately infects snails and human beings through free-living infectious stperiods. Host snails were placed in tanks at a continuous thickness either alone or via one or two various other species of non-hold snails and then exposed to the parasite12. In single-species therapies, hold snails were 30% even more most likely to be infected bereason parasites in multi-species therapies frequently finished up in dead-finish hosts. Increased parasite–hold enrespond to rates brought about by decreased diversity are enough to increase condition transmission for Schistosoma.
The loss of species can increase encounter rates in between pathogens and also hosts, as in the Schistosoma example, once the shed species are not hosts for the pathogen. But if the lost species are indeed hosts capable of transmission, this declining diversity might likewise mitigate the complete number of hosts, thereby decreasing transmission if all else stays equal13,14. Certainly reductions in the variety of hosts have the right to reduce the number of vectors15 and additionally their infection prevalence16,17, yet empirical examples are relatively rare, in part bereason the concern has been neglected, and also bereason all else rarely stays equal. For instance, the loss of hosts can cause compensatory increases in the abundances of other hosts, such that total hold abundance changes little loved one to total host abundance in even more diverse communities. Even as soon as total hold abundance does decrease in less diverse units, distinctions in organize top quality among species can alter straightforward correlationships in between hold abundance and also infection risk18.
Pathogen transmission is not constantly a duty of host density. For instance, the variety of contagious bites delivered by extremely mobile vectors choose mosquitoes deserve to be independent of the density of the organize population14. Transmission of straight transmitted pathogens like hantavirsupplies can also be independent of host thickness if transmission entails behavioural encounters, for example, aggressive interactions in between rodents, and also if the frequency of these encounters does not differ a lot through hold density14,19. In units favor these, the loss of hold species can actually rise transmission if the shed hosts are suboptimal for parasite development and reproduction; this is because these suboptimal hosts absorb pathogens but are negative at transmitting them.
In amount, reducing biodiversity ca rise disease transmission once the shed species are either not hosts for the pathogen or are suboptimal ones. For pathogens for which transmission is a role of organize density, loss of diversity is the majority of likely to increase transmission if the loss causes a boost in the thickness of proficient hosts. The number and also diversity of examples of pathogens for which species loss leads to boosts in full transmission argues that these conditions are commonly met (Table 1). Further research studies in various other disease devices would much better develop the generality of these relationships.
The loss of biodiversity can impact the transmission of infectious diseases65 by changing:
(1) The abundance of the organize or vector. For plants, seeding experimental areas via plant species that are not hosts for fungal pathogens lessened threefold the pathogen fill of species that are hosts, supposedly by reducing organize thickness through competition66. On the other hand also, a better diversity of organize species can sometimes rise pathogen transmission by increasing the abundance of vectors67.
(2) The behaviour of the organize, vector or parasite. In an extra varied area, among the parasitic worms that reasons schistosomiasis (which infects 200 million world worldwide) is more likely to end up in an unsuitable intermediate host. This have the right to reduce the probability of succeeding infection of human beings by 25–99% (ref. 68). For hantavirus in Utah, USA, rodent hosts on more varied plots are more most likely to come in call via heterocertain mammals and less most likely to come in call via conspecifics, reducing the probability of transmission of the virus55. In principle, higher diversity might affect behaviours with a resulting boost in illness transmission65 or could change the evolutionary dynamics of virulence and also transmission pathmeans.
(3) The condition of the organize or vector. In experimental rice areas in China, rice plants in genetically varied mixtures had actually drier leaves bereason the mixture adjusted microclimatic conditions69. As a repercussion, infection through rice blast fungus was less common in diverse fields. Genetically diverse plantings deserve to additionally lead to induced resistance in host plants because they are exposed to equivalent pathogens that are experts on the various other cultivars70.
For some condition devices (for instance, Lyme disease), multiple mechanisms operate in concert, leading to a compounding impact of biodiversity loss on boosted disease transmission (Table 1).
Hantaviroffers are a group of negative-stranded RNA viruses linked through murid rodents. They can cause major morbidity and mortality in humans, via case-fatality prices close to 40% (ref. 71). Infected rodents burned hantavirus in saliva, urine and also faeces; transmission to humans occurs via inhalation of aerosolized excreta and through rodent bites72. The risk of humale expocertain increases as the thickness and infection pervasiveness of rodent reservoirs increase72.
In a field research in Oregon, USA, the just variable substantially connected to infection prevalence in deer mouse host populations was mammalian species diversity, with the ubiquity of the hantavirus Sin Nombre virus increasing from 2% to 14% as diversity decreased. Deer computer mouse populace density was not statistically linked via Sin Nombre virus infection ubiquity, suggesting that high diversity diminished intracertain encounters quite than hold abundance56. A research in Utah, USA55, additionally found a negative correlation in between small-mammal diversity and also Sin Nombre virus infection pervasiveness in deer mice. As in Oregon, high diversity lessened infection prevalence apparently by reducing intraparticular encounters rather than by reducing hold density, a result sustained by experiments19.
The conclusions of these studies were supported by an speculative study of hantaviruses in tiny mammal communities of Panamá23. In replicated plots, small-mammal diversity was reduced by trapping and removing species that are not hosts for the virus; infection ubiquity in hosts was compared on manipulated and also unmanipulated plots (Box 2 figure). Experimentally lessened small-mammal diversity resulted in a boost in the thickness of organize species and additionally in seroconversion rates and seroubiquity within hosts (Box 2 figure).
Species diversity versus species identityThe loss of particular species in a community clearly has the potential to increase disease transmission. But does reducing diversity itself boost transmission, or is boosted transmission the consequence of the removal of particular species? The answer depends on how species complace transforms as richness changes20,21. For instance, if those organize species many responsible for amplifying the pathogen tfinish to persist or even grow as biodiversity is shed, then condition danger will certainly continuously rise as biodiversity declines. On the other hand, if amplifying species tfinish to disappear as biodiversity declines, then biodiversity loss will tend to reduce condition hazard. These theoretical possibilities indicate the prestige of knowledge both the non-random sequences by which species are lost from neighborhoods, and also whether the species that tfinish to happen only in even more species-affluent areas tend to amplify or buffer pathogen transmission.
In a number of instance studies, the species a lot of likely to be lost from environmental communities as diversity declines are those many likely to alleviate pathogen transmission. In the Lyme illness device of eastern North America, for example, the white-footed computer mouse is all at once the a lot of plentiful hold species, the many skilled organize for the Lyme bacterium, and also the highest-quality hold for immature tick vectors18 (Fig. 1). As an effect, this hold species infects a high proportion of the ticks within woodland communities. The white-footed mouse is likewise an ecologically durable species, current in both species-affluent and species-poor communities22. In comparison, Virginia opossums are negative hosts for the pathogen, kill the huge majority of ticks that attempt to feed on them, and also are lacking from many kind of low-diversity forest fragments and degraded woodlands wright here mice are abundant18,22. Because of this, as biodiversity is shed, the host through a solid buffering effect—the opossum—disappears, while the host via a solid amplifying effect—the mouse—continues to be. The major hosts for the pathogens that cause West Nile encephalitis, hantavirus pulmonary syndrome, and also bartonellosis likewise appear to be sturdy species that boost in abundance as biodiversity is lost7,23,24.
Lyme illness is transmitted to humans by the bite of an infected blacklegged tick (Ixodes scapularis). Immature ticks can obtain the infection if they feed on an infected host and have the right to end up being transmittable to people if they ultimately survive to the following life stage. White-footed mice are abundant in northeastern woodlands and feed many kind of ticks18. Ticks that attempt to feed on Virginia opossums are likely to be groomed off and eliminated. Green-and-yellow circles show the expect variety of ticks per hectare fed by mice or opossums; yellow shading reflects the propercent of ticks infected after feeding. Blue circles show the expect number of ticks per hectare groomed off and also eliminated. Ticks that feed on mice are extremely likely to come to be infected through the bacterium that causes Lyme disease, whereas those that feed on opossums are not.
Whether an organism’s hold competence and its resilience to components that mitigate biodiversity are causally related is an unresolved however instrumental problem. Traits that make a hold long lasting to biodiversity loss might likewise make them vulnerable to pathogen infection and also transmission. Such a relationship would describe the frequency through which the attach between diversity loss and illness transmission has been observed in jiyuushikan.org (Table 1). For plants, species that are fast-growing and nutrient-rich via reasonably high metabolic rates—attributes of ‘weedy’ species—can be even more knowledgeable hosts for arthropod vectors and plant pathogens than those via much less weedy traits25. Plants with these weedy traits are additionally more most likely to become even more abundant once plant diversity declines26. Consequently, the extremely species that have traits permitting persistence in degraded and species-bad ecounits are also more likely to bring high pathogen and vector burdens. A similar pattern may occur in vertebrates—resilience in the confront of disturbances that cause biodiversity loss, such as habitat damage and fragmentation, is promoted by life-background features such as high refertile output and intrinsic rates of increase27. Vertebrates via these features tfinish to invest minimally in some facets of adaptive immunity28,29,30; we hypothesize that this may make them even more competent hosts for pathogens and also vectors. Understanding the interrelationships among pathogen transmission, biodiversity loss and also intercertain distinctions in immune feature is a crucial area for future study. Such researches would illuminate just how typically sturdy species are additionally those that increase pathogen transmission, and can carry out basic rules about the impact of biodiversity loss on illness transmission.
Diversity within individual hosts
Could alters in biodiversity within the bodies of organisms also alter pathogen transmission? Recent enhancements in the ability of researchers to detect unculturable microbial species have permitted documentation of the remarkable diversity of microbes upon and within plants and pets. In huguy bodies, for instance, 90% of all cells are microbial31. A number of researches have started to display web links between conditions and also the diversity of an organism’s ‘microbiome’.
Changes in the complace of microbiomes are frequently associated through infection and also condition. For instance, corals experiencing from white torment condition have actually microbial areas distinctly various from those in healthy and balanced corals32. In people, bacterial vaginosis outcomes from changes in the composition of the vaginal microbial community33, and also this subsequently boosts the danger of HIV infection34. Although transforms in microbial species complace linked via infection are well-recorded, few studies have actually investigated the effects of alters in diversity itself. In a current examination, patients through recurrent episodes of infection caused by the bacterium Clostridium difficile had substantially reduced diversity of intestinal microbes than did regulate patients35. Correlational studies such as these, though intriguing, make it hard to identify whether changes in microbial neighborhoods are the cause or the consequence of infections. But some speculative studies plainly demonstrate that boosting microbial biodiversity deserve to defend versus infection. For instance, kids via a history of ear infections given a mixture of five strains of Streptococcus were less likely to build succeeding infections compared to a control group36. Similarly, reducing microbial diversity within a hold can increase transmission. When mice through persistent infections of C. difficile were treated with antibiotics that diminished the diversity of intestinal microbes, they started shedding C. difficile spores at high rates37.
In some of these examples, a well-off microbial neighborhood shows up to manage the abundance of endemic microbial species that have the right to end up being pathogenic once overly abundant35. In other cases, high microbial species diversity can prevent emigration by invasive pathogenic species. For instance, the more diverse the microbiome surrounding the roots of wwarm plants, the even more safeguarded the plants were versus intrusion by the pathogenic bacterium Pseudomonas aeruginosa38. Similarly, piglets increased in organic settings supporting a high diversity of microbes were even more resistant to invasion by pathogenic gut microbes than those increased in more sterile environments39.
The results of microbial diversity within and upon host bodies present intriguing similarities to the impacts of macroscopic species diversity on illness transmission in aquatic and terrestrial ecosystems. Additional expedition of these similarities, and especially the particular mechanisms operating within hosts, is an essential study frontier because alters in microbial diversity might acfirm biodiversity loss in their hosts.
For pathogens currently establiburned within ecological communities, we have shown that biodiversity loss generally increases the rate of transmission. But what role, if any type of, does biodiversity have in the procedures by which new pathogens emerge? Between 1940 and also 2004, over 300 arising condition occasions were determined in people about the world40. Concomitantly, other arising transmittable conditions additionally showed up in wildlife, domesticated animals, and chop and wild plants. Emerging contagious illness incorporate those in which the pathogen has actually progressed right into a new strain within the exact same hold species, for example, via the evolution of drug resistance (methicillin-resistant Staphylococcus aureus or MRSA) or switched to brand-new hold species (for example, huguy immunodeficiency virus or HIV, significant acute respiratory syndrome or SARS). In some situations, the switch to new host species is accompanied by a readjust in geographical selection (for example, West Nile virus in the Americas).
For pathogens that create in brand-new species, the development procedure entails multiple steps, including the initial intrusion right into the new organize (‘spillover’), the production of transmission stages within the brand-new organize, and the facility of the pathogen in the organize populace as a whole41,42. The impact of biodiversity may vary for each of these measures. For the initial intrusion, biodiversity may act as a resource pool. This hypothesis is supported by surveys of arising conditions of humans: most are zoonotic—jumping to human beings from various other vertebprice animals43. In one current evaluation, the probability of appearance of pathogens from wildlife to people was positively associated with mammalian wildlife species richness as soon as data were corrected for reporting bias40. Other ecological and also sociofinancial determinants that bring people right into closer call with potentially new pathogens (for instance, woodland clearing for farming, wildlife hunting) might likewise add to this pattern. Indeed, nearly fifty percent of the zoonotic illness that have emerged in humans because 1940 resulted from alters in land use, from changes in agricultural or various other food production practices, or from wildlife searching (Fig. 2). These huguy activities boost prices of contact in between humans and animals, which might be a vital element underlying spillover.
Figure 2: Drivers and areas of introduction occasions for zoonotic infectious diseases in people from 1940–2005.
a, Worldwide portion of appearance events brought about by each driver; b, Countries in which the development occasions took location, and the motorists of appearance. The dimension of the circle represents the number of appearance events: for range, the number of occasions in the United States was 59. Globally, nearly fifty percent of these illness resulted from alters in land usage, changes in agricultural and other food manufacturing practices, or through wildlife searching, which says that call rates between people and also various other pets are a critical underlying reason of zoonotic illness development. ‘Other’ includes international travel and also commerce, changes in huguy demographics and behaviour, changes in the medical industry, climate and weather, breakdvery own of public health and wellness steps, and also unstated causes. Analysed from information in ref. 40.
Once spillover of the pathogen into a brand-new hold has actually emerged, high densities of that hold species may facilitate pathogen establishment and transmission within the brand-new host41. For example, Nipah virus spilled over from wild fruit bats to residential pigs in Malaysia; high densities of pigs in regional farms show up to have actually assisted in establishment of pig-to-pig transmission, and also the pathogen then spilled over from pigs to humans44. Such high densities of tamed species are almost constantly associated through low biodiversity.
In contrast to appearance with host-switching, 20% of appearance events in between 1940 and 2004 arose via the development of drug resistance40. For these situations, biodiversity of microbial communities within hosts might have actually a protective effect; humale usage of antibiotics is thshould choose for resistant microbes by eliminating the good diversity of non-resistant microbial strains and also species that suppush resistant strains in the lack of antibiotics. Investigations using recent advances in microbial detection assistance this idea45,46. Therefore, diminished microbial diversity might be an important underlying reason of the introduction of drug-resistant pathogens; this as well requires even more examination.
The addition of particular species—for instance, herbal opponents or competitors—have the right to mitigate the effects of establiburned pathogens. For example, speculative addition of a normally developing bacterium, Janthinobacterium lividum, to the skin of the intimidated frog Rana mucosa got rid of frog mortality from speculative infection with chytridiomycosis, which is damaging amphibian populaces worldwide47. For corals, application of pheras isolated from organic areas deserve to regulate the spreview of bacterial infections48. The growing interemainder in ‘probiotics’ for human beings and also harvested species provides an additional instance of this approach49.
More broadly, biodiversity itself seems to defend organisms, consisting of human beings, from transmission of infectious conditions in many kind of situations (Table 1). Preserving biodiversity in these instances, and probably primarily, may mitigate the incidence of established pathogens. To keep high diversity in jiyuushikan.org, conservation scientists have arisen robust methods that reflect the crucial principle that larger locations sustain bigger numbers of species50. Methods of conserving microbial diversity within and upon bodies or in the atmosphere are less well arisen, however preventing the overuse of antimicrobial compounds is essential. Critically, future research on the partnership in between biodiversity and also condition must protect against conflating the effects of biogeographical trends of biodiversity (for example, better diversity in lower latitudes) via those of anthropogenic reductions in extant biodiversity, because policy and administration alternatives can much even more readily affect the last than the former.
For arising illness, the observation that a much more varied microbiome within a hold suppresses strains that are resistant to antimicrobial compounds argues that avoiding the over-usage of these compounds in medicine and agriculture can prevent the introduction of resistant strains. For pathogens that emerge by switching organize species, 3 administration approaches are warranted. First, potential appearance ‘hotspots’ might be predictable on the basis of land-use adjust and underlying biodiversity patterns; these locations should be targeted for surveillance of endemic wildlife pathogens that have actually the potential to jump hold species40,51. 2nd, keeping and protecting intact habitats in these hotspots offers a basic, straight way of reducing human–animal call and reduces the likelihood of emergence of brand-new pathogens, although techniques for achieving lessened contact are not constantly straightforward51. And third, to minimize the probcapability that pathogens come to be establiburned and transmissible within a brand-new hold populace when spillover occurs, the husbandry of high-density monosocieties of domestic pets, specifically in locations at high risk of spillover, have to be subject both to even more extensive security and also to actions that minimize call between wildlife and livestock. Managing potential appearance hotspots by attempting to remove them is most likely to backfire bereason the species the majority of sturdy to habitat destruction and also degradation might be those that amplify pathogen transmission.
In spite of many type of recent advances in our knowledge of biodiversity and condition, much remains to be learned. First, we must rise the variety of condition devices for which we understand the effects of biodiversity loss on disease transmission across a variety of spatial and also temporal scales. We have to additionally focus on exactly how to implement specific policies increated by this scientific research. Future study, for instance, need to monitor changes in public health in regions in which conservation measures are applied compared to recommendation sites. A significant challenge will certainly be to untangle the facility means in which other global anthropogenic trends—such as climate readjust, biotic exreadjust, nutrient contamination, armed conflict and economic collapse—interact through biodiversity loss to affect illness dynamics, and also which of these fads have actually the best impacts upon human health. In spite of continuing to be questions, relationships between biodiversity and also condition are now sufficiently clear to boost the urgency of regional, regional, and worldwide initiatives to preserve herbal ecounits and the biodiversity they contain.
Marris, E. New UN science body to monitor biosphere. jiyuushikan.org 10.1038/news.2010.297 (2010)25
Cronin, J. P., Welsh, M. E., Dekkers, M. G., Abercrombie, S. T. & Mitchell, C. E. Host physiological phenoform defines pathogen reservoir potential. Ecol. Lett. 10.1111/j.1461–0248.2010.01513.x (2010)71
CDC. Hantavirus pulmonary syndrome in 5 pediatric patients—four claims, 2009. Morbidity Mortality Week. Rep. 58, 1409–1412 (2009)
We acunderstanding the support of the joint NSF-NIH Ecology of Infectious Disease programme and the EPA Biodiversity and also Person Health programme. M. Gillespie offered aid in the preparation of the manuscript.
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Department of Biology, Bard College, Annandale, 12504, New York, USA
Department of Biological Sciences, Virginia Tech, Blacksburg, 24061, Virginia, USA
Lisa K. Belden
EcoHealth Alliance, New York, 10001, New York, USA
Peter Daszak & Tiffany type of Bogich
EEB, Eno Hall, Princeton University, Princeton, 08544-3417, New Jersey, USA
Department of Ecology & Evolutionary Biology, Cornell College, Ithaca, 14853, New York, USA
C. Drew Harvell
Department of Biology, College of Florida, Gainesville, 32611, Florida, USA
Robert D. Holt
Center for Infectious Disease Dynamics, Pennsylvania State University, College Station, 16802, Pennsylvania, USA
College of Veterinary Medicine, Oregon State University, Corvallis, 97331-4801, Oregon, USA
Institute of Zoology, Zoological Society of London, London, NW1 4RY, UK
Kate E. Jones
Department of Biology, The College of North Carolina at Chapel Hill, Chapel Hill, 27599, North Carolina, USA
Charles E. Mitchell
Harvard Medical School, Harvard University, Cambridge, 02138, Massachusetts, USA
Samuel S. Myers
Cary Institute of Ecomechanism Studies, Millbrook, 12545, New York, USA
Ricdifficult S. Ostfeld
F.K. and R.S.O. conceived the review. F.K., L.K.B., P.D., A.D., C.D.H., R.D.H., P.H., A.J., K.E.J., C.E.M., S.S.M. and also R.S.O. created and also edited the message. T.B. prepared Fig. 2.