Density-dependent network structuring within and across wild animal systems

Gregory F Albery^{1

2

3}, Daniel J Becker⁴, Josh A Firth^{5

6}, Delphine De Moor⁷, Sanjana Ravindran⁸, Matthew Silk^{8

9

10}, Amy R Sweeny^{8

11}, Eric Vander Wal¹², Quinn Webber¹³, Bryony Allen^{14

15}, Simon A Babayan¹⁶, Sahas Barve^{17

18}, Mike Begon¹⁴, Richard J Birtles¹⁹, Theadora A Block^{20

21}, Barbara A Block²², Janette E Bradley²³, Sarah Budischak²⁴, Christina Buesching²⁵, Sarah J Burthe²⁶, Aaron B Carlisle²⁷, Jennifer E Caselle²⁸, Ciro Cattuto²⁹, Alexis S Chaine³⁰, Taylor K Chapple³¹, Barbara J Cheney³², Timothy Clutton-Brock³³, Melissa Collier³⁴, David J Curnick¹⁵, Richard J Delahay³⁵, Damien R Farine^{36

37

38

39}, Andy Fenton¹⁴, Francesco Ferretti⁴⁰, Laura Feyrer^{41

42}, Helen Fielding^{43

44}, Vivienne Foroughirad^{34

45}, Celine Frere⁴⁶, Michael G Gardner⁴⁷, Eli Geffen⁴⁸, Stephanie S Godfrey⁴⁹, Andrea L Graham⁵⁰, Phil S Hammond⁵¹, Maik Henrich⁵², Marco Heurich^{52

53

54}, Paul Hopwood¹⁰, Amiyaal Ilany⁵⁵, Joseph A Jackson¹⁹, Nicola Jackson⁴⁶, David M P Jacoby⁵⁶, Ann-Marie Jacoby⁵⁷, Miloš Ježek⁵⁸, Lucinda Kirkpatrick^{59

60}, Alisa Klamm⁶¹, James A Klarevas-Irby^{36

37

62

63}, Sarah Knowles⁵, Lee Koren⁵⁵, Ewa Krzyszczyk⁶⁰, Jillian M Kusch^{12

64}, Xavier Lambin⁶⁵, Jeffrey E Lane⁶⁴, Herwig Leirs⁵⁹, Stephan T Leu⁶⁶, Bruce E Lyon²⁰, David W Macdonald⁶⁷, Anastasia E Madsen^{68

69}, Janet Mann³⁴, Marta Manser³⁷, Joachim Mariën⁵⁹, Apia Massawe⁷⁰, Robbie A McDonald⁴⁴, Kevin Morelle^{58

71}, Johann Mourier^{72

73

74}, Chris Newman⁶⁷, Kenneth Nussear⁷⁵, Brendah Nyaguthii^{38

39

62

76}, Mina Ogino^{36

37

62}, Laura Ozella⁷⁷, Craig Packer⁷⁸, Yannis P Papastamatiou⁷⁹, Steve Paterson⁸⁰, Eric Payne⁸¹, Amy B Pedersen⁸, Josephine M Pemberton⁸, Noa Pinter-Wollman⁸², Serge Planes^{73

74}, Aura Raulo⁵, Rolando Rodríguez-Muñoz¹⁰, Lauren Rudd⁵, Christopher Sabuni⁷⁰, Pratha Sah³⁴, Robert J Schallert⁸³, Ben C Sheldon⁵, Daizaburo Shizuka⁶⁸, Andrew Sih⁸¹, David L Sinn⁸⁴, Vincent Sluydts⁵⁹, Orr Spiegel⁴⁸, Sandra Telfer⁶⁵, Courtney A Thomason^{40

85}, David M Tickler⁸⁶, Tom Tregenza¹⁰, Kimberley VanderWaal⁸⁷, Sam Walmsley⁴¹, Eric L Walters¹⁷, Klara M Wanelik^{80

88}, Hal Whitehead⁴¹, Elodie Wielgus⁵², Jared Wilson-Aggarwal^{10

44}, Caroline Wohlfeil⁴⁷, Shweta Bansal³⁴

Affiliations

¹ School of Natural Sciences, Trinity College Dublin, Dublin, Ireland. gfalbery@gmail.com.
² Department of Biology, Georgetown University, Washington, DC, USA. gfalbery@gmail.com.
³ Wissenschaftskolleg zu Berlin, Berlin, Germany. gfalbery@gmail.com.
⁴ School of Biological Sciences, University of Oklahoma, Norman, OK, USA.
⁵ Department of Biology, University of Oxford, Oxford, UK.
⁶ School of Biology, University of Leeds, Leeds, UK.
⁷ Department of Primate Behavior and Evolution, Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany.
⁸ Institute of Ecology and Evolution, University of Edinburgh, Edinburgh, UK.
⁹ Centre d'Ecologie Fonctionnelle et Evolutive, Université de Montpellier, Centre National de la Recherche Scientifique, École Pratique des Hautes Études, Institut de Recherche pour le Développement, Montpellier, France.
¹⁰ Centre for Ecology and Conservation, University of Exeter, Penryn, UK.
¹¹ School of Biosciences, University of Sheffield, Sheffield, UK.
¹² Department of Biology, Memorial University of Newfoundland, St. John's, Newfoundland and Labrador, Canada.
¹³ University of Guelph, Guelph, Ontario, Canada.
¹⁴ Institute of Infection, Veterinary and Ecological Sciences, University of Liverpool, Liverpool, UK.
¹⁵ Institute of Zoology, Zoological Society of London, London, UK.
¹⁶ University of Glasgow, Glasgow, UK.
¹⁷ Department of Biological Sciences, Old Dominion University, Norfolk, VA, USA.
¹⁸ Avian Ecology Program, Archbold Biological Station, Venus, FL, USA.
¹⁹ University of Salford, Salford, UK.
²⁰ Department of Ecology and Evolutionary Biology, University of California, Santa Cruz, Santa Cruz, CA, USA.
²¹ Canine Companions, Santa Rosa, CA, USA.
²² Hopkins Marine Station, Stanford University, Stanford, CA, USA.
²³ University of Nottingham, Nottingham, UK.
²⁴ Pitzer and Scripps Colleges, Claremont, CA, USA.
²⁵ Okanagan Department of Biology, Irving K. Barber Faculty of Science, University of British Columbia, Kelowna, British Columbia, Canada.
²⁶ UK Centre for Ecology and Hydrology, Edinburgh, UK.
²⁷ University of Delaware, Newark, DE, USA.
²⁸ Marine Science Institute, University of California, Santa Barbara, Santa Barbara, CA, USA.
²⁹ ISI Foundation, Torino, Italy.
³⁰ Evolutionary Ecology Group, Station d'Écologie Theorique et Experimentale du CNRS, Moulis, France.
³¹ Oregon State University, Corvallis, OR, USA.
³² Lighthouse Field Station, School of Biological Sciences, University of Aberdeen, Aberdeen, UK.
³³ Department of Zoology, University of Cambridge, Cambridge, UK.
³⁴ Department of Biology, Georgetown University, Washington, DC, USA.
³⁵ National Wildlife Management Centre, Animal and Plant Health Agency, York, UK.
³⁶ Department of Collective Behavior, Max Planck Institute for Animal Behavior, Konstanz, Germany.
³⁷ Department of Evolutionary Biology and Environmental Studies, University of Zurich, Zurich, Switzerland.
³⁸ Department of Ornithology, National Museum of Kenya, Nairobi, Kenya.
³⁹ Division of Ecology and Evolution, Research School of Biology, Australian National University, Canberra, Australian Capital Territory, Australia.
⁴⁰ Virginia Tech, Blacksburg, VA, USA.
⁴¹ Department of Biology, Dalhousie University, Halifax, Nova Scotia, Canada.
⁴² Department of Fisheries and Oceans, Bedford Institute of Oceanography, Dartmouth, Nova Scotia, Canada.
⁴³ The Royal (Dick) School of Veterinary Studies, University of Edinburgh, Edinburgh, UK.
⁴⁴ Environment and Sustainability Institute, University of Exeter, Penryn, UK.
⁴⁵ Department of Marine Biology, Texas A&M University at Galveston, Galveston, TX, USA.
⁴⁶ School of the Environment, University of Queensland, Brisbane, Queensland, Australia.
⁴⁷ College of Science and Engineering, Flinders University, Adelaide, South Australia, Australia.
⁴⁸ School of Zoology, Faculty of Life Science, Tel Aviv University, Tel Aviv, Israel.
⁴⁹ Department of Zoology, University of Otago, Otago, New Zealand.
⁵⁰ Department of Ecology and Evolutionary Biology, Princeton University, Princeton, NJ, USA.
⁵¹ Sea Mammal Research Unit, South Oceans Institute, University of St Andrews, St Andrews, UK.
⁵² Department of Visitor Management and National Park Monitoring, Bavarian Forest National Park, Grafenau, Germany.
⁵³ Chair of Wildlife Ecology and Wildlife Management, Faculty of Environment and Natural Resources, University of Freiburg, Freiburg, Germany.
⁵⁴ Department of Forestry and Wildlife Management, Faculty of Applied Ecology, Agricultural Sciences and Biotechnology, Inland Norway University of Applied Sciences, Elverum, Norway.
⁵⁵ Faculty of Life Sciences, Bar Ilan University, Ramat Gan, Israel.
⁵⁶ Lancaster University, Lancaster, UK.
⁵⁷ Division of Marine Science and Conservation, Nicholas School of the Environment, Duke University Marine Laboratory, Beaufort, NC, USA.
⁵⁸ Department of Game Management and Wildlife Biology, Faculty of Forestry and Wood Sciences, Czech University of Life Sciences, Prague, Czech Republic.
⁵⁹ Evolutionary Ecology Group-EVECO, Universiteit Antwerpen, Antwerp, Belgium.
⁶⁰ School of Environment and Natural Science, Bangor University, Bangor, Wales.
⁶¹ Department of Nature Conservation and Research, Hainich National Park, Bad Langensalza, Germany.
⁶² Mpala Research Center, Nanyuki, Kenya.
⁶³ Centre for the Advanced Study of Collective Behaviour, University of Konstanz, Konstanz, Germany.
⁶⁴ Department of Biology, University of Saskatchewan, Saskatoon, Saskatchewan, Canada.
⁶⁵ University of Aberdeen, Aberdeen, UK.
⁶⁶ School of Animal and Veterinary Sciences, University of Adelaide, Adelaide, South Australia, Australia.
⁶⁷ Wildlife Conservation Research Unit, The Recanati-Kaplan Centre, Department of Zoology, University of Oxford, Oxford, UK.
⁶⁸ School of Biological Sciences, University of Nebraska-Lincoln, Lincoln, NE, USA.
⁶⁹ Department of Ecology, Behavior, and Evolution, University of California, San Diego, San Diego, CA, USA.
⁷⁰ Institute of Pest Management, Sokoine University of Agriculture, Morogoro, Tanzania.
⁷¹ Max Planck Institute of Animal Behavior, Radolfzell, Germany.
⁷² Biodiversité Marine, Exploitation et Conservation, Université de Montpellier, Centre National de la Recherche Scientifique, Institut Français de Recherche pour l'Exploitation de la Mer, Institut de Recherche pour le Développement, Sète, France.
⁷³ PSL Research University, Université de Perpignan, Perpignan, France.
⁷⁴ Laboratoire d'Excellence CORAIL, Papetoai, French Polynesia.
⁷⁵ University of Nevada, Reno, NV, USA.
⁷⁶ Department of Wildlife, School of Natural Resource Management, University of Eldoret, Eldoret, Kenya.
⁷⁷ Department of Veterinary Sciences, University of Turin, Grugliasco, Italy.
⁷⁸ Department of Ecology, Evolution and Behavior, University of Minnesota, Saint Paul, MN, USA.
⁷⁹ Florida International University, Florida, FL, USA.
⁸⁰ University of Liverpool, Liverpool, UK.
⁸¹ Department of Environmental Science and Policy, University of California, Davis, Davis, CA, USA.
⁸² Department of Ecology and Evolutionary Biology, University of California, Los Angeles, Los Angeles, CA, USA.
⁸³ Stanford University, Stanford, CA, USA.
⁸⁴ Department of Wildlife, California State Polytechnic University, Arcata, CA, USA.
⁸⁵ Texas Tech University, Lubbock, TX, USA.
⁸⁶ University of Western Australia, Perth, Western Australia, Australia.
⁸⁷ Department of Veterinary Population Medicine, University of Minnesota, Minneapolis, MN, USA.
⁸⁸ University of Surrey, Guildford, UK.

PMID: 40908382
DOI: 10.1038/s41559-025-02843-z

Density-dependent network structuring within and across wild animal systems

Gregory F Albery et al. Nat Ecol Evol. 2025.

. 2025 Sep 4.

doi: 10.1038/s41559-025-02843-z. Online ahead of print.

Authors

Affiliations

¹ School of Natural Sciences, Trinity College Dublin, Dublin, Ireland. gfalbery@gmail.com.
² Department of Biology, Georgetown University, Washington, DC, USA. gfalbery@gmail.com.
³ Wissenschaftskolleg zu Berlin, Berlin, Germany. gfalbery@gmail.com.
⁴ School of Biological Sciences, University of Oklahoma, Norman, OK, USA.
⁵ Department of Biology, University of Oxford, Oxford, UK.
⁶ School of Biology, University of Leeds, Leeds, UK.
⁷ Department of Primate Behavior and Evolution, Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany.
⁸ Institute of Ecology and Evolution, University of Edinburgh, Edinburgh, UK.
⁹ Centre d'Ecologie Fonctionnelle et Evolutive, Université de Montpellier, Centre National de la Recherche Scientifique, École Pratique des Hautes Études, Institut de Recherche pour le Développement, Montpellier, France.
¹⁰ Centre for Ecology and Conservation, University of Exeter, Penryn, UK.
¹¹ School of Biosciences, University of Sheffield, Sheffield, UK.
¹² Department of Biology, Memorial University of Newfoundland, St. John's, Newfoundland and Labrador, Canada.
¹³ University of Guelph, Guelph, Ontario, Canada.
¹⁴ Institute of Infection, Veterinary and Ecological Sciences, University of Liverpool, Liverpool, UK.
¹⁵ Institute of Zoology, Zoological Society of London, London, UK.
¹⁶ University of Glasgow, Glasgow, UK.
¹⁷ Department of Biological Sciences, Old Dominion University, Norfolk, VA, USA.
¹⁸ Avian Ecology Program, Archbold Biological Station, Venus, FL, USA.
¹⁹ University of Salford, Salford, UK.
²⁰ Department of Ecology and Evolutionary Biology, University of California, Santa Cruz, Santa Cruz, CA, USA.
²¹ Canine Companions, Santa Rosa, CA, USA.
²² Hopkins Marine Station, Stanford University, Stanford, CA, USA.
²³ University of Nottingham, Nottingham, UK.
²⁴ Pitzer and Scripps Colleges, Claremont, CA, USA.
²⁵ Okanagan Department of Biology, Irving K. Barber Faculty of Science, University of British Columbia, Kelowna, British Columbia, Canada.
²⁶ UK Centre for Ecology and Hydrology, Edinburgh, UK.
²⁷ University of Delaware, Newark, DE, USA.
²⁸ Marine Science Institute, University of California, Santa Barbara, Santa Barbara, CA, USA.
²⁹ ISI Foundation, Torino, Italy.
³⁰ Evolutionary Ecology Group, Station d'Écologie Theorique et Experimentale du CNRS, Moulis, France.
³¹ Oregon State University, Corvallis, OR, USA.
³² Lighthouse Field Station, School of Biological Sciences, University of Aberdeen, Aberdeen, UK.
³³ Department of Zoology, University of Cambridge, Cambridge, UK.
³⁴ Department of Biology, Georgetown University, Washington, DC, USA.
³⁵ National Wildlife Management Centre, Animal and Plant Health Agency, York, UK.
³⁶ Department of Collective Behavior, Max Planck Institute for Animal Behavior, Konstanz, Germany.
³⁷ Department of Evolutionary Biology and Environmental Studies, University of Zurich, Zurich, Switzerland.
³⁸ Department of Ornithology, National Museum of Kenya, Nairobi, Kenya.
³⁹ Division of Ecology and Evolution, Research School of Biology, Australian National University, Canberra, Australian Capital Territory, Australia.
⁴⁰ Virginia Tech, Blacksburg, VA, USA.
⁴¹ Department of Biology, Dalhousie University, Halifax, Nova Scotia, Canada.
⁴² Department of Fisheries and Oceans, Bedford Institute of Oceanography, Dartmouth, Nova Scotia, Canada.
⁴³ The Royal (Dick) School of Veterinary Studies, University of Edinburgh, Edinburgh, UK.
⁴⁴ Environment and Sustainability Institute, University of Exeter, Penryn, UK.
⁴⁵ Department of Marine Biology, Texas A&M University at Galveston, Galveston, TX, USA.
⁴⁶ School of the Environment, University of Queensland, Brisbane, Queensland, Australia.
⁴⁷ College of Science and Engineering, Flinders University, Adelaide, South Australia, Australia.
⁴⁸ School of Zoology, Faculty of Life Science, Tel Aviv University, Tel Aviv, Israel.
⁴⁹ Department of Zoology, University of Otago, Otago, New Zealand.
⁵⁰ Department of Ecology and Evolutionary Biology, Princeton University, Princeton, NJ, USA.
⁵¹ Sea Mammal Research Unit, South Oceans Institute, University of St Andrews, St Andrews, UK.
⁵² Department of Visitor Management and National Park Monitoring, Bavarian Forest National Park, Grafenau, Germany.
⁵³ Chair of Wildlife Ecology and Wildlife Management, Faculty of Environment and Natural Resources, University of Freiburg, Freiburg, Germany.
⁵⁴ Department of Forestry and Wildlife Management, Faculty of Applied Ecology, Agricultural Sciences and Biotechnology, Inland Norway University of Applied Sciences, Elverum, Norway.
⁵⁵ Faculty of Life Sciences, Bar Ilan University, Ramat Gan, Israel.
⁵⁶ Lancaster University, Lancaster, UK.
⁵⁷ Division of Marine Science and Conservation, Nicholas School of the Environment, Duke University Marine Laboratory, Beaufort, NC, USA.
⁵⁸ Department of Game Management and Wildlife Biology, Faculty of Forestry and Wood Sciences, Czech University of Life Sciences, Prague, Czech Republic.
⁵⁹ Evolutionary Ecology Group-EVECO, Universiteit Antwerpen, Antwerp, Belgium.
⁶⁰ School of Environment and Natural Science, Bangor University, Bangor, Wales.
⁶¹ Department of Nature Conservation and Research, Hainich National Park, Bad Langensalza, Germany.
⁶² Mpala Research Center, Nanyuki, Kenya.
⁶³ Centre for the Advanced Study of Collective Behaviour, University of Konstanz, Konstanz, Germany.
⁶⁴ Department of Biology, University of Saskatchewan, Saskatoon, Saskatchewan, Canada.
⁶⁵ University of Aberdeen, Aberdeen, UK.
⁶⁶ School of Animal and Veterinary Sciences, University of Adelaide, Adelaide, South Australia, Australia.
⁶⁷ Wildlife Conservation Research Unit, The Recanati-Kaplan Centre, Department of Zoology, University of Oxford, Oxford, UK.
⁶⁸ School of Biological Sciences, University of Nebraska-Lincoln, Lincoln, NE, USA.
⁶⁹ Department of Ecology, Behavior, and Evolution, University of California, San Diego, San Diego, CA, USA.
⁷⁰ Institute of Pest Management, Sokoine University of Agriculture, Morogoro, Tanzania.
⁷¹ Max Planck Institute of Animal Behavior, Radolfzell, Germany.
⁷² Biodiversité Marine, Exploitation et Conservation, Université de Montpellier, Centre National de la Recherche Scientifique, Institut Français de Recherche pour l'Exploitation de la Mer, Institut de Recherche pour le Développement, Sète, France.
⁷³ PSL Research University, Université de Perpignan, Perpignan, France.
⁷⁴ Laboratoire d'Excellence CORAIL, Papetoai, French Polynesia.
⁷⁵ University of Nevada, Reno, NV, USA.
⁷⁶ Department of Wildlife, School of Natural Resource Management, University of Eldoret, Eldoret, Kenya.
⁷⁷ Department of Veterinary Sciences, University of Turin, Grugliasco, Italy.
⁷⁸ Department of Ecology, Evolution and Behavior, University of Minnesota, Saint Paul, MN, USA.
⁷⁹ Florida International University, Florida, FL, USA.
⁸⁰ University of Liverpool, Liverpool, UK.
⁸¹ Department of Environmental Science and Policy, University of California, Davis, Davis, CA, USA.
⁸² Department of Ecology and Evolutionary Biology, University of California, Los Angeles, Los Angeles, CA, USA.
⁸³ Stanford University, Stanford, CA, USA.
⁸⁴ Department of Wildlife, California State Polytechnic University, Arcata, CA, USA.
⁸⁵ Texas Tech University, Lubbock, TX, USA.
⁸⁶ University of Western Australia, Perth, Western Australia, Australia.
⁸⁷ Department of Veterinary Population Medicine, University of Minnesota, Minneapolis, MN, USA.
⁸⁸ University of Surrey, Guildford, UK.

PMID: 40908382
DOI: 10.1038/s41559-025-02843-z

Abstract

Theory predicts that high population density leads to more strongly connected spatial and social networks, but how local density drives individuals' positions within their networks is unclear. This gap reduces our ability to understand and predict density-dependent processes. Here we show that density drives greater network connectedness at the scale of individuals within wild animal populations. Across 36 datasets of spatial and social behaviour in >58,000 individual animals, spanning 30 species of fish, reptiles, birds, mammals and insects, 80% of systems exhibit strong positive relationships between local density and network centrality. However, >80% of relationships are nonlinear and 75% are shallower at higher values, indicating saturating trends that probably emerge as a result of demographic and behavioural processes that counteract density's effects. These are stronger and less saturating in spatial compared with social networks, as individuals become disproportionately spatially connected rather than socially connected at higher densities. Consequently, ecological processes that depend on spatial connections are probably more density dependent than those involving social interactions. These findings suggest fundamental scaling rules governing animal social dynamics, which could help to predict network structures in novel systems.

PubMed Disclaimer

Conflict of interest statement

Competing interests: The authors declare no competing interests.

References

1. Lloyd-Smith, J. O. et al. Should we expect population thresholds for wildlife disease? Trends Ecol. Evol. 20, 511–519 (2005). - PubMed
1. Albery, G. F. Density dependence and disease dynamics: moving towards a predictive framework. Preprint at EcoEvoRxiv https://doi.org/10.32942/osf.io/gaw49 (2022).
1. Hopkins, S. R., Fleming-Davies, A. E., Belden, L. K. & Wojdak, J. M. Systematic review of modeling assumptions and empirical evidence: does parasite transmission increase nonlinearly with host density? Methods Ecol. Evol. 11, 476–486 (2020).
1. Begon, M. et al. A clarification of transmission terms in host–microparasite models: numbers, densities and areas. Epidemiol. Infect. 129, 147–153 (2002). - PubMed - PMC
1. Ferrari, M. J., Perkins, S, E., Pomeroy, L. W. & Bjrnstad, O. N.Pathogens, social networks, and the paradox of transmission scaling. Interdiscip. Perspect. Infect. Dis. 2011, 267049 (2011). - PubMed - PMC

LinkOut - more resources

Full Text Sources
- Nature Publishing Group
Miscellaneous
- NCI CPTAC Assay Portal

Save citation to file

Email citation

Add to Collections

Add to My Bibliography

Your saved search

Create a file for external citation management software

Your RSS Feed

Density-dependent network structuring within and across wild animal systems

Affiliations

Density-dependent network structuring within and across wild animal systems

Authors

Affiliations

Abstract

Conflict of interest statement

References

LinkOut - more resources

Full Text Sources

Miscellaneous