Density-dependent network structuring within and across wild animal systems

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

doi:10.1101/2024.06.28.601262

Density-dependent network structuring within and across wild animal systems

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

School of Environmental & Natural Sciences

Research output: Contribution to journal › Article › peer-review

Abstract

High population density should drive individuals to more frequently share space and interact, producing better-connected spatial and social networks. Despite this widely-held assumption, it remains unconfirmed how local density generally drives individuals’ positions within wild animal networks. We analysed 34 datasets of simultaneous spatial and social behaviour in >55,000 individual animals, spanning 28 species of fish, reptiles, birds, mammals, and insects. >80% of systems exhibited strongly positive relationships between local density and network centrality, providing broad empirical evidence that local density increases connectedness at the individual level. However, >75% of density-connectedness relationships were nonlinear, and density’s importance declined at higher values in >70% of systems, signifying saturating effects. Density’s effect was much stronger and less saturating for spatial than social networks, suggesting population density drives individuals to become disproportionately spatially connected rather than socially. These findings reveal fundamental trends underlying societal structuring, with widespread behavioural, ecological, and evolutionary implications.

Original language	English
Journal	Nature Ecology and Evolution
DOIs	https://doi.org/10.1101/2024.06.28.601262
Publication status	Published - 16 Jul 2025

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2024.06.28.601262v2.full
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Albery, G. F., Becker, D. J., Firth, J. A., De Moor, D., Ravindran, S., Silk, M., Sweeny, A. R., Vander Wal, E., Webber, Q., Allen, B., Babayan, S. A., Barve, S., Begon, M., Birtles, R. J., Block, T. A., Block, B. A., Bradley, J. E., Budischak, S., Buesching, C., ... Bansal, S. (2025). Density-dependent network structuring within and across wild animal systems. Nature Ecology and Evolution. https://doi.org/10.1101/2024.06.28.601262

@article{d6e7e7f2ee2240038929dcde291fa965,

title = "Density-dependent network structuring within and across wild animal systems",

abstract = "High population density should drive individuals to more frequently share space and interact, producing better-connected spatial and social networks. Despite this widely-held assumption, it remains unconfirmed how local density generally drives individuals{\textquoteright} positions within wild animal networks. We analysed 34 datasets of simultaneous spatial and social behaviour in >55,000 individual animals, spanning 28 species of fish, reptiles, birds, mammals, and insects. >80\% of systems exhibited strongly positive relationships between local density and network centrality, providing broad empirical evidence that local density increases connectedness at the individual level. However, >75\% of density-connectedness relationships were nonlinear, and density{\textquoteright}s importance declined at higher values in >70\% of systems, signifying saturating effects. Density{\textquoteright}s effect was much stronger and less saturating for spatial than social networks, suggesting population density drives individuals to become disproportionately spatially connected rather than socially. These findings reveal fundamental trends underlying societal structuring, with widespread behavioural, ecological, and evolutionary implications.",

author = "Albery, \{Gregory F\} and Becker, \{Daniel J\} and Firth, \{Josh A\} and \{De Moor\}, Delphine and Sanjana Ravindran and Matthew Silk and Sweeny, \{Amy R\} and \{Vander Wal\}, Eric and Quinn Webber and Bryony Allen and Babayan, \{Simon A\} and Sahas Barve and Mike Begon and Birtles, \{Richard J\} and Block, \{Theadora A\} and Block, \{Barbara A\} and Bradley, \{Janette E\} and Sarah Budischak and Christina Buesching and Burthe, \{Sarah J\} and Carlisle, \{Aaron B\} and Caselle, \{Jennifer E\} and Ciro Cattuto and Chaine, \{Alexis S\} and Chapple, \{Taylor K\} and Cheney, \{Barbara J\} and Timothy Clutton-Brock and Melissa Collier and Curnick, \{David J\} and Delahay, \{Richard J\} and Farine, \{Damien R\} and Andy Fenton and Francesco Ferretti and Laura Feyrer and Helen Fielding and Vivienne Foroughirad and Celine Frere and Gardner, \{Michael G\} and Eli Geffen and Godfrey, \{Stephanie S\} and Graham, \{Andrea L\} and Hammond, \{Phil S\} and Maik Henrich and Marco Heurich and Paul Hopwood and Amiyaal Ilany and Jackson, \{Joseph A\} and Nicola Jackson and Jacoby, \{David M P\} and Ann-Marie Jacoby and Milo{\v s} Je{\v z}ek and Lucinda Kirkpatrick and Alisa Klamm and Klarevas-Irby, \{James A\} and Sarah Knowles and Lee Koren and Ewa Krzyszczyk and Kusch, \{Jillian M\} and Xavier Lambin and Lane, \{Jeffrey E\} and Herwig Leirs and Leu, \{Stephan T\} and Lyon, \{Bruce E\} and Macdonald, \{David W\} and Madsen, \{Anastasia E\} and Janet Mann and Marta Manser and Joachim Mari{\"e}n and Apia Massawe and McDonald, \{Robbie A\} and Kevin Morelle and Johann Mourier and Chris Newman and Kenneth Nussear and Brendah Nyaguthii and Mina Ogino and Laura Ozella and Craig Packer and Papastamatiou, \{Yannis P\} and Steve Paterson and Eric Payne and Pedersen, \{Amy B\} and Pemberton, \{Josephine M\} and Noa Pinter-Wollman and Serge Planes and Aura Raulo and Rolando Rodr{\'i}guez-Mu{\~n}oz and Lauren Rudd and Christopher Sabuni and Pratha Sah and Schallert, \{Robert J\} and Sheldon, \{Ben C\} and Daizaburo Shizuka and Andrew Sih and Sinn, \{David L\} and Vincent Sluydts and Orr Spiegel and Sandra Telfer and Thomason, \{Courtney A\} and Tickler, \{David M\} and Tom Tregenza and Kimberley VanderWaal and Sam Walmsley and Walters, \{Eric L\} and Wanelik, \{Klara M\} and Hal Whitehead and Elodie Wielgus and Jared Wilson-Aggarwal and Caroline Wohlfeil and Shweta Bansal",

year = "2025",

month = jul,

day = "16",

doi = "10.1101/2024.06.28.601262",

language = "English",

journal = "Nature Ecology and Evolution",

issn = "2397-334X",

publisher = "Nature Publishing Group",

}

Albery, GF, Becker, DJ, Firth, JA, De Moor, D, Ravindran, S, Silk, M, Sweeny, AR, Vander Wal, E, Webber, Q, Allen, B, Babayan, SA, Barve, S, Begon, M, Birtles, RJ, Block, TA, Block, BA, Bradley, JE, Budischak, S, Buesching, C, Burthe, SJ, Carlisle, AB, Caselle, JE, Cattuto, C, Chaine, AS, Chapple, TK, Cheney, BJ, Clutton-Brock, T, Collier, M, Curnick, DJ, Delahay, RJ, Farine, DR, Fenton, A, Ferretti, F, Feyrer, L, Fielding, H, Foroughirad, V, Frere, C, Gardner, MG, Geffen, E, Godfrey, SS, Graham, AL, Hammond, PS, Henrich, M, Heurich, M, Hopwood, P, Ilany, A, Jackson, JA, Jackson, N, Jacoby, DMP, Jacoby, A-M, Ježek, M, Kirkpatrick, L, Klamm, A, Klarevas-Irby, JA, Knowles, S, Koren, L, Krzyszczyk, E, Kusch, JM, Lambin, X, Lane, JE, Leirs, H, Leu, ST, Lyon, BE, Macdonald, DW, Madsen, AE, Mann, J, Manser, M, Mariën, J, Massawe, A, McDonald, RA, Morelle, K, Mourier, J, Newman, C, Nussear, K, Nyaguthii, B, Ogino, M, Ozella, L, Packer, C, Papastamatiou, YP, Paterson, S, Payne, E, Pedersen, AB, Pemberton, JM, Pinter-Wollman, N, Planes, S, Raulo, A, Rodríguez-Muñoz, R, Rudd, L, Sabuni, C, Sah, P, Schallert, RJ, Sheldon, BC, Shizuka, D, Sih, A, Sinn, DL, Sluydts, V, Spiegel, O, Telfer, S, Thomason, CA, Tickler, DM, Tregenza, T, VanderWaal, K, Walmsley, S, Walters, EL, Wanelik, KM, Whitehead, H, Wielgus, E, Wilson-Aggarwal, J, Wohlfeil, C & Bansal, S 2025, 'Density-dependent network structuring within and across wild animal systems', Nature Ecology and Evolution. https://doi.org/10.1101/2024.06.28.601262

TY - JOUR

T1 - Density-dependent network structuring within and across wild animal systems

AU - Albery, Gregory F

AU - Becker, Daniel J

AU - Firth, Josh A

AU - De Moor, Delphine

AU - Ravindran, Sanjana

AU - Silk, Matthew

AU - Sweeny, Amy R

AU - Vander Wal, Eric

AU - Webber, Quinn

AU - Allen, Bryony

AU - Babayan, Simon A

AU - Barve, Sahas

AU - Begon, Mike

AU - Birtles, Richard J

AU - Block, Theadora A

AU - Block, Barbara A

AU - Bradley, Janette E

AU - Budischak, Sarah

AU - Buesching, Christina

AU - Burthe, Sarah J

AU - Carlisle, Aaron B

AU - Caselle, Jennifer E

AU - Cattuto, Ciro

AU - Chaine, Alexis S

AU - Chapple, Taylor K

AU - Cheney, Barbara J

AU - Clutton-Brock, Timothy

AU - Collier, Melissa

AU - Curnick, David J

AU - Delahay, Richard J

AU - Farine, Damien R

AU - Fenton, Andy

AU - Ferretti, Francesco

AU - Feyrer, Laura

AU - Fielding, Helen

AU - Foroughirad, Vivienne

AU - Frere, Celine

AU - Gardner, Michael G

AU - Geffen, Eli

AU - Godfrey, Stephanie S

AU - Graham, Andrea L

AU - Hammond, Phil S

AU - Henrich, Maik

AU - Heurich, Marco

AU - Hopwood, Paul

AU - Ilany, Amiyaal

AU - Jackson, Joseph A

AU - Jackson, Nicola

AU - Jacoby, David M P

AU - Jacoby, Ann-Marie

AU - Ježek, Miloš

AU - Kirkpatrick, Lucinda

AU - Klamm, Alisa

AU - Klarevas-Irby, James A

AU - Knowles, Sarah

AU - Koren, Lee

AU - Krzyszczyk, Ewa

AU - Kusch, Jillian M

AU - Lambin, Xavier

AU - Lane, Jeffrey E

AU - Leirs, Herwig

AU - Leu, Stephan T

AU - Lyon, Bruce E

AU - Macdonald, David W

AU - Madsen, Anastasia E

AU - Mann, Janet

AU - Manser, Marta

AU - Mariën, Joachim

AU - Massawe, Apia

AU - McDonald, Robbie A

AU - Morelle, Kevin

AU - Mourier, Johann

AU - Newman, Chris

AU - Nussear, Kenneth

AU - Nyaguthii, Brendah

AU - Ogino, Mina

AU - Ozella, Laura

AU - Packer, Craig

AU - Papastamatiou, Yannis P

AU - Paterson, Steve

AU - Payne, Eric

AU - Pedersen, Amy B

AU - Pemberton, Josephine M

AU - Pinter-Wollman, Noa

AU - Planes, Serge

AU - Raulo, Aura

AU - Rodríguez-Muñoz, Rolando

AU - Rudd, Lauren

AU - Sabuni, Christopher

AU - Sah, Pratha

AU - Schallert, Robert J

AU - Sheldon, Ben C

AU - Shizuka, Daizaburo

AU - Sih, Andrew

AU - Sinn, David L

AU - Sluydts, Vincent

AU - Spiegel, Orr

AU - Telfer, Sandra

AU - Thomason, Courtney A

AU - Tickler, David M

AU - Tregenza, Tom

AU - VanderWaal, Kimberley

AU - Walmsley, Sam

AU - Walters, Eric L

AU - Wanelik, Klara M

AU - Whitehead, Hal

AU - Wielgus, Elodie

AU - Wilson-Aggarwal, Jared

AU - Wohlfeil, Caroline

AU - Bansal, Shweta

PY - 2025/7/16

Y1 - 2025/7/16

N2 - High population density should drive individuals to more frequently share space and interact, producing better-connected spatial and social networks. Despite this widely-held assumption, it remains unconfirmed how local density generally drives individuals’ positions within wild animal networks. We analysed 34 datasets of simultaneous spatial and social behaviour in >55,000 individual animals, spanning 28 species of fish, reptiles, birds, mammals, and insects. >80% of systems exhibited strongly positive relationships between local density and network centrality, providing broad empirical evidence that local density increases connectedness at the individual level. However, >75% of density-connectedness relationships were nonlinear, and density’s importance declined at higher values in >70% of systems, signifying saturating effects. Density’s effect was much stronger and less saturating for spatial than social networks, suggesting population density drives individuals to become disproportionately spatially connected rather than socially. These findings reveal fundamental trends underlying societal structuring, with widespread behavioural, ecological, and evolutionary implications.

AB - High population density should drive individuals to more frequently share space and interact, producing better-connected spatial and social networks. Despite this widely-held assumption, it remains unconfirmed how local density generally drives individuals’ positions within wild animal networks. We analysed 34 datasets of simultaneous spatial and social behaviour in >55,000 individual animals, spanning 28 species of fish, reptiles, birds, mammals, and insects. >80% of systems exhibited strongly positive relationships between local density and network centrality, providing broad empirical evidence that local density increases connectedness at the individual level. However, >75% of density-connectedness relationships were nonlinear, and density’s importance declined at higher values in >70% of systems, signifying saturating effects. Density’s effect was much stronger and less saturating for spatial than social networks, suggesting population density drives individuals to become disproportionately spatially connected rather than socially. These findings reveal fundamental trends underlying societal structuring, with widespread behavioural, ecological, and evolutionary implications.

U2 - 10.1101/2024.06.28.601262

DO - 10.1101/2024.06.28.601262

M3 - Article

C2 - 40908382

SN - 2397-334X

JO - Nature Ecology and Evolution

JF - Nature Ecology and Evolution

ER -

Density-dependent network structuring within and across wild animal systems

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