Density-dependent network structuring within and across wild animal systems

Albery, Gregory F.; Becker, Daniel J.; Firth, Josh A.; De Moor, Delphine; Ravindran, Sanjana; Silk, Matthew; Sweeny, Amy R.; Vander Wal, Eric; Webber, Quinn; Allen, Bryony; Babayan, Simon A.; Barve, Sahas; Begon, Mike; Birtles, Richard J.; Block, Theadora A.; Block, Barbara A.; Bradley, Janette E.; Budischak, Sarah; Buesching, Christina; Burthe, Sarah J.; Carlisle, Aaron B.; Caselle, Jennifer E.; Cattuto, Ciro; Chaine, Alexis S.; Chapple, Taylor K.; Cheney, Barbara J.; Clutton-Brock, Timothy; Collier, Melissa; Curnick, David J.; Delahay, Richard J.; Farine, Damien R.; Fenton, Andy; Ferretti, Francesco; Feyrer, Laura; Fielding, Helen; Foroughirad, Vivienne; Frere, Celine; Gardner, Michael G.; Geffen, Eli; Godfrey, Stephanie S.; Graham, Andrea L.; Hammond, Phil S.; Henrich, Maik; Heurich, Marco; Hopwood, Paul; Ilany, Amiyaal; Jackson, Joseph A.; Jackson, Nicola; Jacoby, David M.P.; Jacoby, Ann-Marie; Ježek, Miloš; Kirkpatrick, Lucinda; Klamm, Alisa; Klarevas-Irby, James A.; Knowles, Sarah; Koren, Lee; Krzyszczyk, Ewa; Kusch, Jillian M.; Lambin, Xavier; Lane, Jeffrey E.; Leirs, Herwig; Leu, Stephan T.; Lyon, Bruce E.; Macdonald, David W.; Madsen, Anastasia E.; Mann, Janet; Manser, Marta; Mariën, Joachim; Massawe, Apia; McDonald, Robbie A.; Morelle, Kevin; Mourier, Johann; Newman, Chris; Nussear, Kenneth; Nyaguthii, Brendah; Ogino, Mina; Ozella, Laura; Packer, Craig; Papastamatiou, Yannis P.; Paterson, Steve; Payne, Eric; Pedersen, Amy B.; Pemberton, Josephine M.; Pinter-Wollman, Noa; Planes, Serge; Raulo, Aura; Rodríguez-Muñoz, Rolando; Rudd, Lauren; Sabuni, Christopher; Sah, Pratha; Schallert, Robert J.; Sheldon, Ben C.; Shizuka, Daizaburo; Sih, Andrew; Sinn, David L.; Sluydts, Vincent; Spiegel, Orr; Telfer, Sandra; Thomason, Courtney A.; Tickler, David M.; Tregenza, Tom; VanderWaal, Kimberley; Walmsley, Sam; Walters, Eric L.; Wanelik, Klara M.; Whitehead, Hal; Wielgus, Elodie; Wilson-Aggarwal, Jared; Wohlfeil, Caroline; Bansal, Shweta

Albery, Gregory F. ORCID: https://orcid.org/0000-0001-6260-2662; Becker, Daniel J. ORCID: https://orcid.org/0000-0003-4315-8628; Firth, Josh A. ORCID: https://orcid.org/0000-0001-7183-4115; De Moor, Delphine ORCID: https://orcid.org/0000-0003-2474-6125; Ravindran, Sanjana; Silk, Matthew ORCID: https://orcid.org/0000-0002-8318-5383; Sweeny, Amy R.; Vander Wal, Eric; Webber, Quinn ORCID: https://orcid.org/0000-0002-0434-9360; Allen, Bryony; Babayan, Simon A. ORCID: https://orcid.org/0000-0002-4949-1117; Barve, Sahas ORCID: https://orcid.org/0000-0001-5840-8023; Begon, Mike; Birtles, Richard J.; Block, Theadora A.; Block, Barbara A.; Bradley, Janette E.; Budischak, Sarah; Buesching, Christina; Burthe, Sarah J. ORCID: https://orcid.org/0000-0001-8871-3432; Carlisle, Aaron B.; Caselle, Jennifer E.; Cattuto, Ciro; Chaine, Alexis S.; Chapple, Taylor K.; Cheney, Barbara J.; Clutton-Brock, Timothy ORCID: https://orcid.org/0000-0001-8110-8969; Collier, Melissa; Curnick, David J. ORCID: https://orcid.org/0000-0002-3093-1282; Delahay, Richard J.; Farine, Damien R.; Fenton, Andy; Ferretti, Francesco ORCID: https://orcid.org/0000-0001-9510-3552; Feyrer, Laura ORCID: https://orcid.org/0000-0002-9300-1947; Fielding, Helen; Foroughirad, Vivienne; Frere, Celine; Gardner, Michael G.; Geffen, Eli; Godfrey, Stephanie S.; Graham, Andrea L. ORCID: https://orcid.org/0000-0002-6580-2755; Hammond, Phil S.; Henrich, Maik; Heurich, Marco ORCID: https://orcid.org/0000-0003-0051-2930; Hopwood, Paul; Ilany, Amiyaal ORCID: https://orcid.org/0000-0001-8089-569X; Jackson, Joseph A. ORCID: https://orcid.org/0000-0003-0330-5478; Jackson, Nicola ORCID: https://orcid.org/0000-0002-3082-6829; Jacoby, David M.P. ORCID: https://orcid.org/0000-0003-2729-3811; Jacoby, Ann-Marie; Ježek, Miloš ORCID: https://orcid.org/0000-0002-4127-9947; Kirkpatrick, Lucinda ORCID: https://orcid.org/0000-0002-0161-2469; Klamm, Alisa; Klarevas-Irby, James A. ORCID: https://orcid.org/0000-0001-7499-4085; Knowles, Sarah ORCID: https://orcid.org/0000-0002-2368-4623; Koren, Lee ORCID: https://orcid.org/0000-0002-7425-501X; Krzyszczyk, Ewa; Kusch, Jillian M. ORCID: https://orcid.org/0000-0003-0078-5621; Lambin, Xavier ORCID: https://orcid.org/0000-0003-4643-2653; Lane, Jeffrey E.; Leirs, Herwig ORCID: https://orcid.org/0000-0002-7612-5024; Leu, Stephan T. ORCID: https://orcid.org/0000-0003-2547-5056; Lyon, Bruce E.; Macdonald, David W.; Madsen, Anastasia E.; Mann, Janet; Manser, Marta ORCID: https://orcid.org/0000-0001-8787-5667; Mariën, Joachim; Massawe, Apia; McDonald, Robbie A. ORCID: https://orcid.org/0000-0002-6922-3195; Morelle, Kevin; Mourier, Johann ORCID: https://orcid.org/0000-0001-9019-1717; Newman, Chris; Nussear, Kenneth ORCID: https://orcid.org/0000-0002-3849-8911; Nyaguthii, Brendah; Ogino, Mina ORCID: https://orcid.org/0000-0002-3780-2942; Ozella, Laura; Packer, Craig ORCID: https://orcid.org/0000-0002-3939-8162; Papastamatiou, Yannis P. ORCID: https://orcid.org/0000-0002-6091-6841; Paterson, Steve ORCID: https://orcid.org/0000-0002-1307-2981; Payne, Eric; Pedersen, Amy B.; Pemberton, Josephine M.; Pinter-Wollman, Noa; Planes, Serge; Raulo, Aura; Rodríguez-Muñoz, Rolando; Rudd, Lauren ORCID: https://orcid.org/0000-0002-5925-9514; Sabuni, Christopher; Sah, Pratha; Schallert, Robert J.; Sheldon, Ben C. ORCID: https://orcid.org/0000-0002-5240-7828; Shizuka, Daizaburo; Sih, Andrew; Sinn, David L.; Sluydts, Vincent; Spiegel, Orr ORCID: https://orcid.org/0000-0001-8941-3175; Telfer, Sandra; Thomason, Courtney A.; Tickler, David M.; Tregenza, Tom; VanderWaal, Kimberley ORCID: https://orcid.org/0000-0002-5764-6430; Walmsley, Sam; Walters, Eric L. ORCID: https://orcid.org/0000-0002-9414-5758; Wanelik, Klara M. ORCID: https://orcid.org/0000-0003-1485-0340; Whitehead, Hal; Wielgus, Elodie ORCID: https://orcid.org/0000-0001-8323-5128; Wilson-Aggarwal, Jared; Wohlfeil, Caroline; Bansal, Shweta. 2025 Density-dependent network structuring within and across wild animal systems. Nature Ecology & Evolution, 9 (11). 2002-2013. 10.1038/s41559-025-02843-z

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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.

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