Infectious Disease and Group Size: More than Just a Numbers Game
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Increased risk of infectious disease is assumed to be a major cost of group living, yet empirical evidence for this effect is mixed. We studied whether larger social groups are more subdivided structurally. If so, the social subdivisions that form in larger groups may act as barriers to the spread of infection, weakening the association between group size and infectious disease. To investigate this ‘social bottleneck’ hypothesis, we examined the association between group size and four network structure metrics in 43 vertebrate and invertebrate species. We focused on metrics involving modularity, clustering, distance and centralization. In a meta-analysis of intraspecific variation in social networks, modularity showed positive associations with network size, with a weaker but still positive effect in cross-species analyses. Network distance also showed a positive association with group size when using intraspecific variation. We then used a theoretical model to explore the effects of subgrouping relative to other effects that influence disease spread in socially structured populations. Outbreaks reached higher prevalence when groups were larger, but subgrouping reduced prevalence. Subgrouping also acted as a ‘brake’ on disease spread between groups. We suggest research directions to understand the conditions under which larger groups become more subdivided, and to devise new metrics that account for subgrouping when investigating the links between sociality and infectious disease risk.
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High-density living and increased contact can increase pathogen transmission rates among hosts.
“High-density living and increased contact can increase pathogen transmission rates among hosts (Nunn et al. 2015).”
From Red fox viromes in urban and rural landscapes (2020) · cites this paper as Nunn et al. 2015
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Newman, M. E. J. (2006) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.0601602103 · Cited by 12,256 · see citing works
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Dynamics and Control of Diseases in Networks with Community Structure
Salathé, M., & Jones, J. H. (2010) · PLoS Computational Biology · doi:10.1371/journal.pcbi.1000736 · Cited by 586 · see citing works
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“Weak ties are also central to the social bottleneck hypothesis, which proposes that clique-like structuring within social groups slows the spread of pathogens (Nunn et al. 2015).”
BackgroundHawley, D. M., Gibson, A., Townsend, A., Craft, M. E., & Stephenson, J. (2020) · Parasitology · doi:10.1017/s0031182020002048 · Cited by 42 · see citing works
“…whether this higher modularity in larger social groups represents an evolved response to limit parasite spread (as likely occurs in eusocial insects; Stroeymeyt et al., 2018), or simply a side-effect of the need for individuals to limit social interactions within larger groups (Nunn et al., 2015).”
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Bats as putative Zaire ebolavirus reservoir hosts and their habitat suitability in Africa
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“However, the frequent and tight behavioural interactions expressed by group members may also entail major fitness costs, as they typically favour the transmission of pathogens between individuals (Stroeymeyt et al., 2014; Mccabe et al., 2015; Theis et al., 2015).”
BackgroundBora, M., Bora, D., Manu, M., Barman, N., Dutta, L., Kumar, P. P., Poovathikkal, S., Suresh, K., & Nimmanapalli, R. (2020) · Pathogens · doi:10.3390/pathogens9121044 · Cited by 35
“According to studies related to infectious diseases and group sizes, it has been found that outbreaks of any disease reach higher prevalence when groups are larger and dense [40].”
BackgroundDrivers and consequences of variation in individual social connectivity
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Superorganism Immunity: A Major Transition in Immune System Evolution
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Social structure defines spatial transmission of African swine fever in wild boar
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“For 44 example, social organization into family groups (one dimension of social structure) causes local 45 clustering of hosts that can alter pathogen transmission dynamics in a broad range of wildlife 46 disease systems, such as malaria in primates, fungal infection in termites, intestinal parasites in 47 African Art…”
BackgroundOlival, K., Latinne, A., Islam, A., Epstein, J., Hersch, R., Engstrand, R., Gurley, E., Amato, G., Luby, S., & Daszak, P. (2020) · Molecular Ecology · doi:10.1111/mec.15288 · Cited by 30 · see citing works
Wild cricket social networks show stability across generations
Fisher, D. N., Rodríguez-Muñoz, R., & Tregenza, T. (2016) · BMC Evolutionary Biology · doi:10.1186/s12862-016-0726-9 · Cited by 30 · see citing works
“Similar results have been found across social networks of various species, where larger networks also tended to be more modular than smaller networks, which may limit disease transmission [63, 64].”
ResultEvans, J., Hodgson, D., Boogert, N., & Silk, M. (2021) · Behavioral Ecology and Sociobiology · doi:10.1007/s00265-021-03102-4 · Cited by 29 · see citing works
“In species with social groups that have modular social structures (Nunn et al. 2015) such as fission-fusion societies (Lehmann et al. 2007; Ramos-Fernández and Morales 2014), we would predict conformist social learning to be more favoured than in species without these structures or sub-groupings,…”
BackgroundResultThe trade-off between information and pathogen transmission in animal societies
Romano, V., Sueur, C., & MacIntosh, A. (2020) · Oikos · doi:10.32942/osf.io/vqt4g · Cited by 29
“For example, researchers have examined the association between group size and social network metrics in 43 vertebrate and invertebrate species and showed that modularity, which is expected to increase in larger groups, acted as a buffer, reducing disease spread between subgroups [24].”
BackgroundFluctuating survival selection explains variation in avian group size
Brown, C., Brown, M., Roche, E. A., O'Brien, V., & Page, C. E. (2016) · Proceedings of the National Academy of Sciences of the United States of America · doi:10.1073/pnas.1600218113 · Cited by 28
How does individual variation in sociality influence fitness in prairie voles?Preprint
Sabol, A. C., Lambert, C., Keane, B., Solomon, N. G., & Dantzer, B. (2019) · bioRxiv · doi:10.1101/676858 · Cited by 27 · see citing works
“Although high levels of sociality can have beneficial effects on individual fitness, it may also carry costs for an individual’s health or physical condition (Nunn et al., 2015).”
BackgroundBalasubramaniam, K., Beisner, B., Guan, J., Vandeleest, J. J., Hsieh, F., Atwill, E., & McCowan, B. (2018) · PeerJ · doi:10.7717/peerj.4271/supp-18 · Cited by 27
“Analogous to this ‘‘social bottleneck hypothesis’’ (Nunn et al., 2015), we might expect greater commensal microbial sharing among individuals of the same social behavioral communities, compared to individuals across communities.”
BackgroundInfluentialModelling cetacean morbillivirus outbreaks in an endangered killer whale population
Weiss, M. N., Franks, D., Balcomb, K., Ellifrit, D., Silk, M., Cant, M., & Croft, D. (2020) · Biological Conservation · doi:10.1016/j.biocon.2019.108398 · Cited by 26 · see citing works
Genomic Epidemiology and Evolution of Escherichia coli in Wild Animals in Mexico
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“Larger social groups, as observed in 321 Carnivora, Artiodactyla and Primates in Figure S4C, is known to increase the social transmissions 322 of infectious agents in animal societies, which may facilitate the dispersion of virulence genes 323 (44).”
BackgroundMendenhall, I., Skiles, M., Neves, E. S., Borthwick, S., Low, D. H. W., Liang, B., Lee, B., Su, Y. C. F., & Smith, G. J. D. (2017) · One Health · doi:10.1016/j.onehlt.2017.10.001 · Cited by 25 · see citing works
“Contact rates drive infectious disease transmission and larger colony sizes may play a role in transmission [60,61] through periodic introductions of immunologically naïve individuals [46].”
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“…Silk et al., 2014) and by a range of ecological factors including resource availability and competition (Koenig et al., 1992; Russell, 2004), predation intensity (Elgar, 1989; Beauchamp, 2008), disease risk (Nunn et al., 2015) and climate (Jetz and Rubenstein, 2011; Cockburn and Russell, 2011).”
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“Properties of the whole network might favour information transmission (e.g. density; Pasquaretta et al., 2014), reduce spreading processes (e.g. modularity; Nunn et al., 2015) or those same properties might even have dual roles regarding transmission (e.g. modularity; Romano et al., 2018).”
BackgroundResultExposure and susceptibility drive reinfection with gastrointestinal parasites in a social primate
Müller-Klein, N., Heistermann, M., Strube, C., Franz, M., Schülke, O., & Ostner, J. (2019) · Functional Ecology · doi:10.1111/1365-2435.13313 · Cited by 24 · see citing works
Formica, V., Donald, H., Marti, H., Irgebay, Z., & Brodie, E. (2020) · Journal of Animal Ecology · doi:10.1111/1365-2656.13322 · Cited by 23
Physiological and social consequences of gastrointestinal nematode infection in a nonhuman primate
Müller-Klein, N., Heistermann, M., Strube, C., Morbach, Z. M., Lilie, N. D., Franz, M., Schülke, O., & Ostner, J. (2019) · Behavioral Ecology · doi:10.1093/beheco/ary168 · Cited by 23 · see citing works
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The Behavioral Ecology of the Tibetan Macaque
Li, J.-H., Kappeler, P., & Sun, L.-X. (2020) · doi:10.1007/978-3-030-27920-2 · Cited by 23
American foulbrood in a honeybee colony: spore-symptom relationship and feedbacks
Stephan, J. G., de Miranda, J. D., & Forsgren, E. (2019) · BMC Ecology · doi:10.1186/s12898-020-00283-w · Cited by 22
The ecology of movement and behaviour: a saturated tripartite network for describing animal contacts
Manlove, K., Aiello, C. M., Sah, P., Cummins, B., Hudson, P., & Cross, P. (2018) · Proceedings of the Royal Society of London. Biological Sciences · doi:10.1098/rspb.2018.0670 · Cited by 22 · see citing works
Brookes, V., VanderWaal, K., & Ward, M. P. (2020) · Preventive Veterinary Medicine · doi:10.1016/j.prevetmed.2018.09.008 · Cited by 21
Sociality and tattoo skin disease among bottlenose dolphins in Shark Bay, Australia
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Gilbertson, M. L. J., Fountain-Jones, N., & Craft, M. E. (2018) · Behaviour · doi:10.1163/1568539x-00003471 · Cited by 21
“…contact heterogeneity include individual behavioral phenotypes (Natoli et al., 2005; Dizney & Dearing, 2013), the social structure of a population (Nunn et al., 2015; Sah et al., 2017), seasonal or temporal behavior changes (Chen et al., 2014), and behavioral responses to anthropogenic influences…”
BackgroundViral Richness is Positively Related to Group Size, but Not Mating System, in Bats
Webber, Q., Fletcher, Q., & Willis, C. (2017) · EcoHealth · doi:10.1007/s10393-017-1276-3 · Cited by 21 · see citing works
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“The size of social groups [Nunn & Heymann, 2005], subgroup structuring [Nunn et al., 2015], and position in a social network [Rimbach et al.”
BackgroundFerreira, S., Hofer, H., de Carvalho, L. M. M., & East, M. (2019) · Ecology and Evolution · doi:10.1002/ece3.5431 · Cited by 20 · see citing works
Larger, more connected societies of ants have a higher prevalence of viruses
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Hierarchical networks of food exchange in the black garden ant Lasius niger
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“We expected colony size to affect the network topology, as previous studies have shown that centralization and modularity increases with colony size (Pasquaretta et al., 2014; Nunn et al., 2015).”
BackgroundResultWhite, L. A., Siva-Jothy, J. A., Craft, M. E., & Vale, P. (2020) · Proceedings of the Royal Society B · doi:10.1098/rspb.2020.1653 · Cited by 18
Cowl, V. B., Jensen, K., Lea, J. M. D., Walker, S. L., & Shultz, S. (2020) · International journal of primatology · doi:10.1007/s10764-020-00139-6 · Cited by 18 · see citing works
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Infectious Disease and Grouping Patterns in Mule Deer
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“However, empirical evidence for this feature is mixed [4, 15], and the relationships between epidemiological and ecological processes is intricate [16].”
BackgroundLocalized removal affects white‐tailed deer space use and contacts
Tosa, M. I., Schauber, E., & Nielsen, C. (2017) · doi:10.1002/jwmg.21176 · Cited by 18
“Social behavior shapes contact patterns and, as a result, disease transmission opportunities within host populations (Anderson et al. 1986, Altizer et al. 2003, Nunn et al. 2015).”
BackgroundComparative approaches in social network ecology.
Albery, G. F., Bansal, S., & Silk, M. (2023) · Ecology Letters · doi:10.1111/ele.14345 · Cited by 17
Pathogens Shape Sex Differences in Mammalian Aging
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“Among these traits, variation in host body size [44], mating system and sexual size dimorphism [42], MHC allelic diversity [45], geographical range [46], social group size [47], population density [48], and phylogeny [6,41] have been identified as playing a role.”
BackgroundDisease transmission in animal transfer networks.
Sintayehu, D. W., Prins, H., Heitkonig, I., & de Boer, W. D. (2017) · Preventive Veterinary Medicine · doi:10.1016/j.prevetmed.2016.12.017 · Cited by 17 · see citing works
“Network analysis has become increasingly popular to examine elationships between host contact patterns and disease trans- ission dynamics (Lindstrom et al., 2012; Makagon et al., 2012; anderWaal et al., 2013b; Nunn et al., 2015).”
BackgroundMethodInfluentialKauffman, K., Werner, C. S., Titcomb, G., Pender, M., Rabezara, J. Y., Herrera, J., Shapiro, J., Solis, A., Soarimalala, V., Tortosa, P., et al. (2022) · Journal of the Royal Society Interface · doi:10.1098/rsif.2021.0690 · Cited by 16
Self-citationHuman sickness detection is not dependent on cultural experience
Arshamian, A., Sundelin, T., Wnuk, E., O'Meara, C., Burenhult, N., Rodríguez, G. G., Lekander, M., Olsson, M., Lasselin, J., Axelsson, J., et al. (2021) · Proceedings of the Royal Society B · doi:10.1098/rspb.2021.0922 · Cited by 16 · see citing works
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“We found that network modularity increased with group size, a trend commonly reported and often suggested to reduce parasite transmission in larger groups (Nunn et al., 2015; Sah et al., 2017).”
BackgroundZoonotic Blood-Borne Pathogens in Non-Human Primates in the Neotropical Region: A Systematic Review
Carrillo-Bilbao, G. A., Martin-Solano, S., & Saegerman, C. (2021) · Pathogens · doi:10.3390/pathogens10081009 · Cited by 20 · see citing works
How disease constrains the evolution of social systems
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“In a society where individuals associate in social groups, forming stable subpopulations, the likelihood of a locally transmitted infection spreading globally is low [14,29,30].”
BackgroundQuality, quantity, and the adaptive function of social relationships.
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Functional traits linked to pathogen prevalence in wild bee communities
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Primate Infectious Disease Ecology: Insights and Future Directions at the Human-Macaque Interface
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“…size of a town is positively linked with the risk of death from infectious diseases [4 – 6], and (ii) if the division into subgroups, such as number of households and villages within a given town, decreases the number of casualties as is suggested by the ‘ social bottle-neck hypothesis ’ [2].”
BackgroundResultInfluentialArcher, E. K., Bennett, N., Faulkes, C., & Lutermann, H. (2015) · Oecologia · doi:10.1007/s00442-015-3494-0 · Cited by 12 · see citing works
“…study on the same primate population (Chapman et al. 2012), several other studies in primates and carnivores have provided theoretical and empirical evidence for a role of intergroup contact rates for endoparasite transmission (Walsh et al. 2009; Drewe 2010; Nunn et al. 2015; Rimbach et al. 2015).”
BackgroundHeat stress conditions affect the social network structure of free‐ranging sheep
Borthwick, Z., Quiring, K., Griffith, S., & Leu, S. (2023) · bioRxiv · doi:10.1101/2023.02.26.530064 · Cited by 10
“Regardless, a well-connected social network with low modularity also improves the spread of pathogens and parasites, which has neg - ative well-being and health consequences (Griffin & Nunn, 2012; Nunn et al., 2015 ).”
BackgroundHow Emergent Social Patterns in Allogrooming Combat Parasitic Infections
Wilson, S. N., Sindi, S. S., Brooks, H. C., Hohn, M. E., Price, C. R., Radunskaya, A., Williams, N. D., & Fefferman, N. (2020) · Frontiers in Ecology and Evolution · doi:10.3389/fevo.2020.00054 · Cited by 10
“, modularity, clustering, host density, and mating systems) (Altizer et al., 2003b; Nunn et al., 2015), the strategies proposed in the current work are not meant to capture meaningful, real-world behaviors.”
BackgroundMulti-level societies: different tasks at different social levels
Camerlenghi, E., & Papageorgiou, D. (2025) · Philosophical transactions of the Royal Society of London. Series B, Biological sciences · doi:10.1098/rstb.2023.0274 · Cited by 9 · see citing works
“Additionally, large social unit size poses a greater risk of infectious disease [78] and is linked to higher inequality of division of labour among members of the same social unit [79].”
Variation in local population size predicts social network structure in wild songbirds
Beck, K., Farine, D., Firth, J., & Sheldon, B. (2023) · Journal of Animal Ecology · doi:10.1111/1365-2656.14015 · Cited by 9 · see citing works
“Interspecies comparisons have also suggested that network fragmentation (modularity) generally increases with increasing group size (Nunn et al., 2015 ).”
BackgroundDecline and fall: The causes of group failure in cooperatively breeding meerkats
Duncan, C., Manser, M., & Clutton-Brock, T. (2021) · Ecology and Evolution · doi:10.1002/ece3.7655 · Cited by 9
“Moreover, it is hypothesized that the increasingly modular organization of larger groups could reduce disease prevalence (Nunn et al., 2015), with social clustering that could limit disease transmission being reported in meerkats (Drewe et al., 2011).”
BackgroundPoulin, R., & Filion, A. (2021) · Behavioral Ecology and Sociobiology · doi:10.1007/s00265-021-03039-8 · Cited by 9 · see citing works
Linking parasitism to network centrality and the impact of sampling bias in its interpretation
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“…linked to increased parasitism, although this relationship is not always linear (Freeland, 1976; Altizer et al., 2003; Griffin and Nunn, 2012; Godfrey, 2013; Nunn et al., 2015, 2015; Romano et al., 2016; Rushmore et al., 2017; White et al., 2017; Briard and Ezenwa, 2021; Lucatelli et al., 2021).”
BackgroundSumner, K. M., McCabe, C. M., & Nunn, C. L. (2018) · Behaviour · doi:10.1163/1568539x-00003508 · Cited by 9 · see citing works
Self-citationInhibitory systems in brain evolution: pathways of vulnerability in neurodevelopmental disorders
Hanson, K., Greiner, D., Schumann, C. M., & Semendeferi, K. (2024) · Brain, Behavior and Evolution · doi:10.1159/000540865 · Cited by 8
“…between pathogen-rich environments and high social transmission of behavior and social learning has been demonstrated [167], though some of the risk associated with increased social transmission of behavior and culture may be mitigated by social subgrouping and fi ssion-fusion dynamics [168].”
Primate malarias as a model for cross-species parasite transmission
Voinson, M., Nunn, C. L., & Goldberg, A. (2022) · eLife · doi:10.7554/elife.69628 · Cited by 12 · see citing works
Self-citationWebber, Q., & Wal, E. V. (2020) · Behavioral Ecology and Sociobiology · doi:10.1007/s00265-020-02860-x · Cited by 8
McCabe, C. M., & Nunn, C. L. (2018) · Frontiers in Veterinary Science · doi:10.3389/fvets.2018.00071 · Cited by 8 · see citing works
Self-citationThe natural history of social bonds
Silk, J. (2025) · Annals of the New York Academy of Sciences · doi:10.1111/nyas.15318 · Cited by 7 · see citing works
Whittier, C., Nutter, F., Johnson, P. L. F., Cross, P., Lloyd-Smith, J., Slenning, B., & Stoskopf, M. (2021) · American Journal of Primatology · doi:10.1002/ajp.23350 · Cited by 7
“Increased modularity (i.e., separation of social units) has been shown to decrease pathogen spread and result in smaller outbreaks of disease in various wildlife species (Griffin & Nunn, 2012; Kurvers et al., 2014; Nunn et al., 2015; Sah et al., 2017).”
BackgroundYoung, M., Silk, M., Pritchard, A. J., & Fefferman, N. (2021) · Physical Review E · doi:10.1103/physreve.105.044315 · Cited by 7 · see citing works
“Such population level emergence of assortative mixing has important consequences for anticipating the spread of disease in a heterogeneous society, especially as homophily will also increase modularity and alter the dynamics of the spread of both disease and information [55, 56, 57, 7].”
BackgroundSocial group size influences pathogen transmission in salamanders
Ritchie, K. L., Vredenburg, V., Chaukulkar, S., Butler, H., & Zink, A. G. (2021) · Behavioral Ecology and Sociobiology · doi:10.1007/s00265-021-03057-6 · Cited by 7 · see citing works
Scaling of ant colony interaction networks
Toth, J. M., Fewell, J., & Waters, J. S. (2023) · Frontiers in Ecology and Evolution · doi:10.3389/fevo.2022.993627 · Cited by 6 · see citing works
“Notably, highly social insect colonies also show lower modularity than other animal social networks, which are generally built around maintaining individual alliances with local neighbors rather than construction around coordination of work (Waters and Fewell, 2012; Nunn et al., 2015).”
BackgroundSelf-citation
- Authors
- Nunn, C. L., Jordán, F., McCabe, C. M., Verdolin, J. L., & Fewell, J. H.
- Affiliations
- Duke University (US)
- University of Trento (IT)
- Harvard University (US)
- Arizona State University (US)
- Corresponding author
- Nunn, Charles L.
- Venue
- Philosophical Transactions of the Royal Society B: Biological Sciences
- Publisher
- The Royal Society
- Published
- 2015-05-26
- Volume
- 370
- Issue
- 1669
- Pages
- 20140111
- ISSN
- 0962-8436 (ISSN-L)
- 1471-2970
- PMC
- PMC4410377
- OpenAlex
- W1966269069
- Cited by
- 192
- References
- 0
- Citations by year
- 2026: 11 · 2025: 9 · 2024: 16 · 2023: 12 · 2022: 13 · 2021: 27 · 2020: 27 · 2019: 15 · 2018: 26 · 2017: 17 · 2016: 14 · 2015: 5
- FWCI
- 2.50
- Type
- journal article
- Language
- en
- Open access
- green
- www.ncbi.nlm.nih.gov
- Locations
- Philosophical Transactions of the Royal Society B Biological Sciences (publishedVersion)
- Europe PMC (PubMed Central) (publishedVersion)
- PubMed Central (publishedVersion)
- PubMed (publishedVersion)
- Topics
- Life Sciences › Agricultural and Biological Sciences › Ecology, Evolution, Behavior and Systematics › Animal Behavior and Reproduction
- Health Sciences › Medicine › Public Health, Environmental and Occupational Health › Mathematical and Theoretical Epidemiology and Ecology Models
- Physical Sciences › Physics and Astronomy › Statistical and Nonlinear Physics › Complex Network Analysis Techniques
- Keywords
- infectious diseases, group size, social networks, network modularity, network distance, network centralization, network clustering, disease spread, sociality, animal social structure, vertebrates, invertebrates
- MeSH terms
- Animals, Behavior, Animal, Communicable Diseases, Invertebrates, Models, Biological, Population Density, Risk Factors, Social Behavior, Species Specificity, Vertebrates
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