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Biodiversity inhibits parasites: Broad evidence for the dilution effect
Open accessAbstract
Significance: The dilution effect hypothesis suggests that diverse ecological communities limit disease spread via several mechanisms. Therefore, biodiversity losses could worsen epidemics that harm humans and wildlife. However, there is contentious debate over whether the hypothesis applies broadly, especially for parasites that infect humans. We address this fundamental question with a formal meta-analysis of >200 assessments relating biodiversity to disease in >60 host–parasite systems. We find overwhelming evidence of dilution, which is independent of host density, study design, and type and specialization of parasites. A second analysis identified similar effects of diversity in plant–herbivore systems. Thus, biodiversity generally decreases parasitism and herbivory. Consequently, human-induced declines in biodiversity could increase human and wildlife diseases and decrease crop and forest production.
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“A rich literature on infectious diseases of wildlife, livestock, and plants demonstrates increased pathogen transmission when biodiversity is lost from some ecological communities (9, 53), supporting the generality of this relationship across nonzoonotic disease systems as well.”
From Impacts of biodiversity and biodiversity loss on zoonotic diseases (2021) · cites this paper as [9]
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“In contrast, biodiversity loss has been shown to often increase the risk of zoonotic diseases, for example, through the dilution effect (9).”
From Impacts of biodiversity and biodiversity loss on zoonotic diseases (2021) · cites this paper as [9]
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“But for decades, we have also known that under some conditions, high biological diversity can decrease the transmission of zoonotic diseases that have already become established (8, 9).”
From Impacts of biodiversity and biodiversity loss on zoonotic diseases (2021) · cites this paper as [9]
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“However, a larger metaanalysis found that the dilution effect was as strong for zoonoses as for other types of diseases (9), a conclusion that was robust to criticisms from the authors of the earlier study (57, 58).”
From Impacts of biodiversity and biodiversity loss on zoonotic diseases (2021) · cites this paper as [9]
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The Relationship between Vector Species Richness and the Risk of Vector-Borne Infectious Diseases
Takimoto, G., Shirakawa, H., & Sato, T. (2022) · The American Naturalist · doi:10.1086/720403 · Cited by 5
“This recognition has motivated studies that investigated the effects of host diversity on disease risk and their meta-analytic reviews (Civitello et al. 2015; Halliday and Rohr 2019).”
BackgroundMori, K., Massolo, A., Marceau, D. J., & Stefanakis, E. (2022) · Ecological Modelling · doi:10.1016/j.ecolmodel.2022.110177 · Cited by 5
Strasburg, M., & Boone, M. D. (2022) · Ecology and Evolution · doi:10.1002/ece3.9512 · Cited by 5
INSECT RICHNESS IN DUNG PATCHES OF CATTLE RAISED IN TWO LIVESTOCK SYSTEMS
Gusmão, M. R., Oliveira, M. C. D. S., Nicodemo, M. L. F., & Pezzopane, J. R. M. (2020) · REVISTA DE AGRICULTURA NEOTROPICAL · doi:10.32404/rean.v7i1.4033 · Cited by 5
Camargo, P. R. D. S., da Silva, R. G., Barbosa, N. P. U., Cardoso, A. V., Assis, P. S., & Pelli, A. (2022) · Research Society and Development · doi:10.33448/rsd-v11i7.30214 · Cited by 6
The wild life of tick-borne pathogens
Hofmeester, T. R. (2016) · doi:10.18174/393112 · Cited by 7
Impact of global change on future Ebola emergence and epidemic potential in AfricaPreprint
Redding, D. W., Atkinson, P. M., Cunningham, A. A., Iacono, G. L., Moses, L., Wood, J. L. N., & Jones, K. E. (2017) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/206169 · Cited by 5
“Emerging Infectious Diseases 653 11(2):283.”
BackgroundBoúúaert, D. C., Smet, L. D., Brunain, M., Dahle, B., Blacquière, T., Dalmon, A., Dezmirean, D. S., Elen, D., Filipi, J., Giurgiu, A.-I., et al. (2022) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2022.03.25.485775 · Cited by 5
Idiosyncratic spatial scaling of biodiversity–disease relationships
Gilbert, N. A., DiRenzo, G. V., & Zipkin, E. F. (2025) · Ecography · doi:10.1111/ecog.07541 · Cited by 4
Reappraisal of the Dilution and Amplification Effect Framework: A Case Study in Lyme Disease
Chen, S., & McFarlane, S. E. (2025) · Ecology and Evolution · doi:10.1002/ece3.71969 · Cited by 6
“A different meta-analysis concluded a strong net dilution effect across multiple disease systems but acknowledged the inability of their meta-analysis to test these mechanisms (Civitello et al. 2015).”
InfluentialMatthews, K. E., Breed, M. F., Stirling, E., Macdonald, L. M., & Cavagnaro, T. R. (2025) · Applied Soil Ecology · doi:10.1016/j.apsoil.2025.106470 · Cited by 4
Biodiversity, Infectious Diseases, and the Dilution Effect
Stefano Bosi, Y., & Desmarchelier, D. (2020) · Environmental Modeling & Assessment · doi:10.1007/s10666-020-09688-9 · Cited by 4
“Setting the Hamiltonian H = e−θtu (c,B)+λ [(r − δ)h+ ω̄ − c], deriving the first-order conditions ∂H/∂c = 0, ∂H/∂h = −λ̇ and ∂H/∂μ = ḣ, and defining μ ≡ λe, we get (8), (9) and (10).”
Influence of colony traits on ectoparasite infestation in birds breeding in mixed-species colonies
Gameiro, J., Veiga, J., Valera, F., Palmeirim, J. M., & Catry, I. (2021) · Parasitology · doi:10.1017/s0031182021000470 · Cited by 4
“High host diversity could decrease infestation risk through dilution effects, as different host species may be differently susceptible and/or competent to different parasite species, but this effect may not always occur and likely depends on specific community compositions (Keesing et al., 2006; Randolph and Dobson, 2…”
BackgroundCOVID-19: a wake-up call to protect planetary health
Pachauri, A., Sevilla, N. P. M., Kedia, S., Pathak, D., Mittal, K., & Philo Magdalene, A. (2021) · Elsevier eBooks · doi:10.1016/b978-0-323-85512-9.00017-6 · Cited by 5
Halle, S., Garrido, M., Noy, K., Messika, I., Kedem, H., Cohen, C., Ytzhak, K., Siegal, Z., Shenbrot, G. I., Abramsky, Z., et al. (2021) · Global Ecology and Biogeography · doi:10.1111/geb.13423 · Cited by 4
Prevalence of Fox Tapeworm in Invasive Muskrats in Flanders (North Belgium)
Cartuyvels, E., Adriaens, T., Baert, K., Huysentruyt, F., & Berge, K. V. D. (2022) · Animals · doi:10.3390/ani12070879 · Cited by 4
“or dilution, where more diverse host communities inhibit the spread of parasites [4].”
BackgroundKozanitas, M., Knaus, B. J., Tabima, J. F., Grünwald, N. J., & Garbelotto, M. M. (2024) · Ecography · doi:10.1111/ecog.07012 · Cited by 5
“Differences in the epidemiological role of different hosts have been shown to modify not only the course of animal and human diseases (Civitello et al. 2015), but also the structure and composition of host populations (Johnson et al. 2008) and the evolutionary trajectories of pathogens themselves…”
BackgroundMolento, M. B., Dall’Anese, J., Pontarolo, D. V., Brandão, Y. D. O., & Yoshitani, Ú. Y. (2024) · Revista Brasileira de Parasitologia Veterinária/Brazilian Journal of Veterinary Parasitology · doi:10.1590/s1984-29612024043 · Cited by 4
Alves, M. H., Mendoza‐Roldan, J. A., Alfaro-Segura, P., Carbonara, M., Gómez, A., Leitón, N. M., Arias-Ortega, J., Solano-Barquero, A., Rojas, A., & Otranto, D. (2025) · International Journal for Parasitology Parasites and Wildlife · doi:10.1016/j.ijppaw.2025.101090 · Cited by 4
“Indeed, as a megadiverse country (Mittermeier et al., 1997), Costa Rica ’ s host diversity may suppress parasite spread through mechanisms, like regulating susceptible host populations or reducing transmission (Civitello et al., 2015).”
Zitoun, J.-L., Rousseau, R., & Gourbière, S. (2025) · Journal of The Royal Society Interface · doi:10.1098/rsif.2025.0283 · Cited by 3
Lesser snow goose helminths show recurring and positive parasite infection-diversity relations
Dargent, F., Morrill, A., Alisauskas, R. T., McLaughlin, J., Dave Shutler, A., & Forbes, M. R. (2017) · International Journal for Parasitology Parasites and Wildlife · doi:10.1016/j.ijppaw.2017.01.003 · Cited by 4
“…infection-diversity relations have neglected to investigate the effects of parasite diversity on infection dynamics (see supplementary materials in Civitello et al., 2015; Johnson et al., 2015), despite evidence for the role of parasites in modifying infection outcomes (e.g. Ezenwa et al., 2010;…”
BackgroundResultInfluentialDuffy, J. E. (2019) · Nature Ecology & Evolution · doi:10.1038/s41559-018-0784-z · Cited by 4
Ahn, S., & Goater, C. P. (2021) · Ecology and Evolution · doi:10.1002/ece3.7823 · Cited by 4
“…pairwise comparisons between heterospecific combinations and monospecific combinations involving fathead minnows at the same total fish density (i.e., 4FHMN treatment versus 2FHMN + 2LNDC) host species richness resulted in decreased rates of parasite infection (reviewed in Civitello et al., 2015).”
BackgroundSaúde e conservação dos animais silvestres na natureza
de Macedo, G. C., Herrera, H. M., Jansen, A. M., de Oliveira, C. E., Rocha, F. L., & Porfírio, G. E. D. O. (2022) · Boletim do Museu Paraense Emílio Goeldi - Ciências Naturais · doi:10.46357/bcnaturais.v16i3.806 · Cited by 4
Demography, education, and research trends in the interdisciplinary field of disease ecology
Brandell, E. E., Becker, D. J., Sampson, L., & Forbes, K. M. (2021) · Ecology and Evolution · doi:10.1002/ece3.8466 · Cited by 4
“In particular, the frequency of published research on the dilution effect has undergone several spikes following key findings (Civitello et al., 2015; Keesing, Holt, & Ostfeld, 2006) and a steady increase in publication rate.”
BackgroundEthics, zoonoses, and human-nonhuman conflict: Covid-19 and beyond
Humphreys, R., Chakraborty, R., & Varghese, N. (2022) · Ethics in Science and Environmental Politics · doi:10.3354/esep00203 · Cited by 4
Vector species richness predicts local mortality rates from Chagas disease
da Silva, G. G., Lopez, V. M., Vilarinho, A. C., DATTO-LIBERATO, F. H., Oliveira, C. J. F., Poulin, R., & Ferreira, R. G. N. (2023) · International Journal for Parasitology · doi:10.1016/j.ijpara.2023.10.002 · Cited by 4
“Two competing hypotheses, the DEH and the AEH, make contrasting predictions regarding the role of local diversity as a determinant of transmission rates (Keesing et al., 2006, 2010; Ostfeld and Keesing, 2012; Civitello et al., 2015).”
InfluentialChoo, J., Nghiem, L. T. P., Benítez‐López, A., & Carrasco, L. R. (2023) · Scientific Reports · doi:10.1038/s41598-023-47448-3 · Cited by 5
Larson, M. D., Levri, E. P., Huzurbazar, S. V., Greenwood, D. J., Wise, K. L., & Krist, A. C. (2020) · PLoS ONE · doi:10.1371/journal.pone.0239762 · Cited by 4
“Our results also contrast with a recent meta-analysis of dilution effect experiments showing that increased host diversity decreased the abundance of parasites across a wide range of parasite types, life cycles, and functional groups [16].”
BackgroundResultCappelli, S. L., Domeignoz‐Horta, L. A., Loaiza, V., & Laine, A. (2021) · doi:10.32942/osf.io/tm5eh · Cited by 4
Global change drivers and the risk of infectious diseasePreprint
Mahon, M. B., Sack, A., Aleuy, O. A., Barbera, C., Brown, E., Buelow, H. N., Civitello, D. J., Cohen, J. M., de Wit, L., Forstchen, M., et al. (2022) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2022.07.21.501013 · Cited by 4
“For example, several studies have demonstrated that infectious disease risk is modified by changes to biodiversity (2-5), climate change (6-9), and chemical pollution (10-12).”
BackgroundResultSelf-citation
Metadata
- Authors
- Civitello, D. J., Cohen, J., Fatima, H., Halstead, N. T., Liriano, J., McMahon, T. A., Ortega, C. N., Sauer, E. L., Sehgal, T., Young, S., & Rohr, J. R.
- Affiliations
- University of South Florida (US)
- ORCID
- Venue
- Proceedings of the National Academy of Sciences
- Publisher
- National Academy of Sciences
- Published
- 2015-06-11
- Volume
- 112
- Issue
- 28
- Pages
- 8667-8671
- ISSN
- 0027-8424 (ISSN-L)
- 1091-6490
- PMC
- PMC4507196
- OpenAlex
- W1787861744
- Semantic Scholar
- 30ae28188a18cd22709c7ef13c462acd9260a5ea
- Cited by
- 758
- References
- 34
- Citations by year
- 2026: 38 · 2025: 72 · 2024: 83 · 2023: 61 · 2022: 81 · 2021: 114 · 2020: 78 · 2019: 61 · 2018: 54 · 2017: 51 · 2016: 49 · 2015: 14
- FWCI
- 50.68
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Locations
- Proceedings of the National Academy of Sciences (publishedVersion)
- Europe PMC (PubMed Central) (publishedVersion)
- PubMed Central (publishedVersion)
- PubMed (publishedVersion)
- Funders
- National Science Foundation (EF-1241889, 1144244)
- HHS | National Institutes of Health (R01GM109499, F32AI112255)
- U.S. Department of Agriculture (NRI 2006-01370, 2009-35102-0543)
- U.S. Environmental Protection Agency (CAREER 83518801)
- Topics
- Life Sciences › Immunology and Microbiology › Parasitology › Parasite Biology and Host Interactions
- Health Sciences › Medicine › Public Health, Environmental and Occupational Health › Zoonotic diseases and public health
- Physical Sciences › Environmental Science › Nature and Landscape Conservation › Ecology and Vegetation Dynamics Studies
- Keywords
- dilution effect, biodiversity, parasite abundance, host diversity, zoonoses, disease risk, host-parasite interactions, plant-herbivore interactions, herbivory, biodiversity loss, human disease, wildlife disease
- MeSH terms
- Animals, Disease Susceptibility, Host-Parasite Interactions, Humans, Parasitic Diseases, Biodiversity
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