SREL Reprint #3799

Intrinsic and environmental drivers of pairwise cohesion in wild Canis social groups

John F. Benson1, David A. Keiter1, Peter J. Mahoney1,2, Benjamin L. Allen3,4, Lee Allen5, Francisco Álvares6,7,
Morgan L. Anderson8, Shannon M. Barber-Meyer9, Adi Barocas10,11, James C. Beasley12, Linda Behrendorff13,
Jerrold L. Belant14, Dean E. Beyer Jr14, Luigi Boitani15, Bridget L. Borg16, Stan Boutin17, Erin E. Boydston18,
Justin L. Brown19, Joseph K. Bump20, Jonathon D. Cepek21, Michael J. Chamberlain22, Yvette M. Chenaux-Ibrahim23,
Seth G. Cherry24, Duško Ćirović25, Paolo Ciucci15, H. Dean Cluff26, Susan M. Cooper27, Kevin R. Crooks28,
Daniel L. J. Dupont29, Robert N. Fisher30, Daniel Fortin31, Thomas D. Gable32, Emilio García33, Eli Geffen34,
Stanley D. Gehrt35, Michael Gillingham36, Douglas C. Heard37, Mark Hebblewhite38, Joseph W. Hinton39,
Austin T. Homkes32, Chris G. Howden40, Djuro Huber41, Pat J. Jackson42, Kyle Joly43, Allicia Kelly44,
Marcella J. Kelly45, Katrien A. Kingdon29, Abhijeet Kulkarni46, Josip Kusak41, Gerald W. Kuzyk47, Bryce C. Lake48,
Luis Llaneza49,50, José Vicente López-Bao33, Daniel R. MacNulty2, Ashley A. D. McLaren51, Philip D. McLoughlin52,
Evelyn H. Merrill17, Kenneth J. Mills53, Numi Mitchell54,55, Seth A. Moore23, Matthew A. Mumma56,
Maureen H. Murray17, Marco Musiani57, Mónia Nakamura6, Eric W. Neilson58, Lalenia M. Neufeld59,
Thomas M. Newsome60, John K. Oakleaf61, Vicente Palacios62, Marlo M. Perdicas63, Thomas Perry64,
Tyler R. Petroelje65, Cyrenea B. Piper61, Christina M. Prokopenko29, Laura R. Prugh66, Seth P. D. Riley19,
Helena Rio-Maior6, Gretchen H. Roffler67, Dale Rollins68, Håkan Sand69, Fiona K. A. Schmiegelow59, Dale R. Seip70,
Mathew S. Sorum43, Colleen C. St. Clair17, Robin Steenweg71, Michael W. Strohbach72, Jack Tatler73,
Maria Thaker74, Connor A. Thompson75, Julie W. Turner29, Abi T. Vanak76,77, Eric Vander Wal29,
Petter Wabakken78, Scott E. Walter79, Sarah C. Webster12, Tyler J. Wheeldon51, Camilla Wikenros69,
Steve K. Windels80, Julie K. Young2, Sana Zabihi-Seissan29, Barbara Zimmermann78, Brent R. Patterson51

1School of Natural Resources, University of Nebraska-Lincoln, Lincoln, Nebraska, USA
2Department of Wildland Resources and Ecology Center, Utah State University, Logan, Utah, USA
3Institute for Life Sciences and the Environment, University of Southern Queensland, Toowoomba,
Queensland, Australia
4Centre for African Conservation Ecology, Nelson Mandela University, Port Elizabeth, South Africa
5Department of Agriculture and Fisheries, Queensland Government, Toowoomba, Australia
6CIBIO/InBIO Research Center in Biodiversity and Genetic Resources, University of Porto, Vairão, Portugal
7BIOPOLIS Program in Genomics, Biodiversity and Land Planning, CIBIO, Campus de Vairão, Vairão, Portugal
8British Columbia Ministry of Forests Lands, and Natural Resource Operations, Prince George, British Columbia,
Canada
9Pacific Whale Foundation, Wailuku, Hawaii, USA
10Hula Research Center, Department of Animal Sciences, Tel-Hai College, Qiryat Shemona, Israel
11MIGAL-Galilee Research Institute, Kiryat Shmona, Israel
12University of Georgia Savannah River Ecology Laboratory, Warnell School of Forestry and Natural
Resources, Aiken, South Carolina, USA
13Department of Environment and Science, K’gari (Fraser Island), Fraser Coast, Queensland, Australia
14Department of Fisheries and Wildlife, Michigan State University, East Lansing, Michigan, USA
15Department of Biology and Biotechnologies, University of Rome La Sapienza, Roma, Italy
16Denali National Park and Preserve, National Park Service, Denali Park, Alaska, USA
17Department of Biological Sciences, University of Alberta, Edmonton, Alberta, Canada
18S.P.E.C.I.E.S., Ventura, California, USA
19Santa Monica Mountains National Recreation Area, National Park Service, Thousand Oaks,
California, USA
20Department of Fisheries, Wildlife, and Conservation Biology, University of Minnesota, Twin Cities, Saint Paul,
Minnesota, USA
21Cleveland Metroparks, Strongsville, Ohio, USA
22Warnell School of Forestry and Natural Resources, University of Georgia, Athens, Georgia, USA
23Grand Portage Band of Lake Superior Chippewa, Grand Portage, Minnesota, USA
24Parks Canada Agency, Radium Hot Springs, British Columbia, Canada
25Faculty of Biology, University of Belgrade, Belgrade, Serbia
26Environment and Natural Resources, Government of the Northwest Territories, Yellowknife,
Northwest Territories, Canada
27Texas A&M AgriLife Research, Uvalde, Texas, USA
28Department of Fish, Wildlife, and Conservation Biology, Colorado State University, Fort Collins,
Colorado, USA
29Department of Biology, Memorial University of Newfoundland and Labrador, St. John’s,
Newfoundland and Labrador, Canada
30Western Ecological Research Center, U.S. Geological Survey, San Diego, California, USA
31Department of Biology, Université Laval, Quebec City, Canada
32Department of Fisheries, Wildlife, and Conservation Biology, University of Minnesota, St. Paul,
Minnesota, USA
33Biodiversity Research Institute (CSIC – Oviedo University – Principality of Asturias), Oviedo University,
Mieres, Spain
34School of Zoology, Tel Aviv University, Tel Aviv, Israel
35School of Environment and Natural Resources, The Ohio State University, Columbus, Ohio, USA
36Ecosystem Science and Management, University of Northern British Columbia, Prince George,
British Columbia, Canada
37Tithonus Wildlife Research, Prince George, British Columbia, Canada
38Wildlife Biology Program, WA Franke College of Forestry and Conservation, University of Montana,
Missoula, Montana, USA
39Wolf Conservation Center, South Salem, New York, USA
40Tricky Solutions, Epping, New South Wales, Australia
41Faculty of Veterinary Medicine, University of Zagreb, Zagreb, Croatia
42Nevada Department of Wildlife, Reno, Nevada, USA
43Yukon-Charley Rivers National Preserve, National Park Service, Fairbanks, Alaska, USA
44Department of Environment and Natural Resources, Government of the Northwest Territories, Fort Smith,
Northwest Territories, Canada
45Department of Fish and Wildlife Conservation, Virginia Polytechnic and State University, Blacksburg,
Virginia, USA
46Ashoka Trust for Research in Ecology and the Environment, Bangalore, India
47Ministry of Environment, Government of Saskatchewan, Meadow Lake, Saskatchewan, Canada
48Yukon Flats National Wildlife Refuge, U.S. Fish and Wildlife Service, Fairbanks, Alaska, USA
49A.RE.NA Asesores en Recursos Naturales S.L., Lugo, Spain
50Área de Zooloxía, Departamento de Bioloxía, Facultade de Ciencias, Universidade da Coruña,
Campus da Zapateira, A Coruña, Spain
51Wildlife Research and Monitoring Section, Ministry of Natural Resources and Forestry, Peterborough,
Ontario, Canada
52Department of Biology, University of Saskatchewan, Saskatoon, Saskatchewan, Canada
53Wyoming Game and Fish Department, Pinedale, Wyoming, USA
54The Conservation Agency, Jamestown, Rhode Island, USA
55Natural Resources Science, University of Rhode Island, Kingston, Rhode Island, USA
56Idaho Department of Fish and Game, Boise, Idaho, USA
57Dipartimento di Scienze Biologiche, Geologiche e Ambientali (BiGeA), Università di Bologna, Bologna, Italy
58Canadian Forest Service, Edmonton, Alberta, Canada
59Department of Renewable Resources, University of Alberta, Edmonton, Alberta, Canada
60School of Life and Environmental Sciences, The University of Sydney, Sydney, New South Wales,
Australia
61U.S. Fish and Wildlife Service, Albuquerque, New Mexico, USA
62Association for the Conservation of Nature in Human Environments, Paterna, Spain
63Summit Metro Parks, Akron, Ohio, USA
64Prince Albert National Park, Parks Canada, Waskesiu Lake, Saskatchewan, Canada
65Wildlife Division, Michigan Department of Natural Resources, Marquette, Michigan, USA
66School of Environmental and Forest Sciences, University of Washington, Seattle, Washington, USA
67Division of Wildlife Conservation, Alaska Department of Fish and Game, Douglas, Alaska, USA
68Rolling Plains Quail Research Foundation, Roby, Texas, USA
69Grimsö Wildlife Research Station, Department of Ecology, Swedish University of Agricultural Sciences,
Uppsala, Sweden
70British Columbia Ministry of Environment, Prince George, British Columbia, Canada
71Canadian Wildlife Service, Environment and Climate Change Canada, Kelowna, British Columbia, Canada
72Technische Universität Braunschweig, Landscape Ecology and Environmental Systems Analysis,
Institute of Geoecology, Braunschweig, Germany
73School of Biological Sciences, The University of Adelaide, North Terrace, South Australia, Australia
74Centre for Ecological Sciences, Indian Institute of Science, Bengaluru, India
75Environmental and Life Sciences Graduate Program, Trent University, Peterborough, Ontario, Canada
76Wellcome Trust/Department of Biotechnology (DBT) India Alliance Program, Kaushik Society, Hyderabad,
India
77School of Life Sciences, University of KwaZulu-Natal, Durban, South Africa
78Faculty of Applied Ecology, Agricultural Sciences and Biotechnology,
Inland Norway University of Applied Sciences, Koppang, Norway
79Kickapoo Valley Reserve, La Farge, Wisconsin, USA
80Voyageurs National Park, US National Park Service, International Falls, Minnesota, USA

Abstract: Animals within social groups respond to costs and benefits of sociality by adjusting the proportion of time they spend in close proximity to other individuals in the group (cohesion). Variation in cohesion between individuals, in turn, shapes important group-level processes such as subgroup formation and fission–fusion dynamics. Although critical to animal sociality, a comprehensive understanding of the factors influencing cohesion remains a gap in our knowledge of cooperative behavior in animals. We tracked 574 individuals from six species within the genus Canis in 15 countries on four continents with GPS telemetry to estimate the time that pairs of individuals within social groups spent in close proximity and test hypotheses regarding drivers of cohesion. Pairs of social canids (Canis spp.) varied widely in the proportion of time they spent together (5%–100%) during seasonal monitoring periods relative to both intrinsic characteristics and environmental conditions. The majority of our data came from three species of wolves (gray wolves, eastern wolves, and red wolves) and coyotes. For these species, cohesion within social groups was greatest between breeding pairs and varied seasonally as the nature of cooperative activities changed relative to annual life history patterns. Across species, wolves were more cohesive than coyotes. For wolves, pairs were less cohesive in larger groups, and when suitable, small prey was present reflecting the constraints of food resources and intragroup competition on social associations. Pair cohesion in wolves declined with increased anthropogenic modification of the landscape and greater climatic variability, underscoring challenges for conserving social top predators in a changing world. We show that pairwise cohesion in social groups varies strongly both within and across Canis species, as individuals respond to changing ecological context defined by resources, competition, and anthropogenic disturbance. Our work highlights that cohesion is a highly plastic component of animal sociality that holds significant promise for elucidating ecological and evolutionary mechanisms underlying cooperative behavior.

Keywords: animal sociality, Canis, cohesion, cooperative behavior, coyotes, group size, human footprint, wolves

SREL Reprint #3799

Benson, J. F., D. A. Keiter, P. J. Mahoney, B. L. Allen, L. Allen, F. Álvares, M. L. Anderson, S. M. Barber-Meyer, A. Barocas, J. C. Beasley, L. Behrendorff, J. L. Belant, D. E. Jr. Beyer, L. Boitani, B. L. Borg, S. Boutin, E. E. Boydston, J. L. Brown, J. K. Bump, J. D. Cepek, M. J. Chamberlain, Y. M. Chenaux-Ibrahim, S. G. Cherry, D. Ćirović, P. Ciucci, H. D. Cluff, S. M. Cooper, K. R. Crooks, D. L. J. Dupont, R. N. Fisher, D. Fortin, T. D. Gable, E. Garcia, E. Geffen, S. D. Gehrt, M. Gillingham, D. C. Heard, M. Hebblewhite, J. W. Hinton, A. T. Homkes, C. G. Howden, D. Huber, P. J. Jackson, K. Joly, A. Kelly, M. J. Kelly, K. A. Kingdon, A. Kulkarni, J. Kusak, G. W. Kuzyk, B. C. Lake, L. Llaneza, J. V. López-Bao, D. R. MacNulty, A. A. D. McLaren, P. D. McLoughlin, E. H. Merrill, K. J. Mills, N. Mitchell, S. A. Moore, M. A. Mumma, M. H. Murray, M. Musiani, M. Nakamura, E. W. Neilson, L. M. Neufeld, T. M. Newsome, J. K. Oakleaf, V. Palacios, M. M. Perdicas, T. Perry, T. R. Petroelje, C. B. Piper, C. M. Prokopenko, L. R. Prugh, S. P. D. Riley, H. Rio-Maior, G. H. Roffler, D. Rollins, H. Sand, F. K. A. Schmiegelow, D. R. Seip, M. S. Sorum, C. C. St. Clair, R. Steenweg, M. W. Strohbach, J. Tatler, M. Thaker, C. A. Thompson, J. W. Turner, A. T. Vanak, E. Vander Wal, P. Wabakken, S. E. Walter, S. C. Webster, T. J. Wheeldon, C. Wikenros, S. K. Windels, J. K. Young, S. Zabihi-Seissan, B. Zimmermann, and B. R. Patterson. 2024. Intrinsic and environmental drivers of pairwise cohesion in wild Canis social groups. Ecology(e4492).

This information was provided by the University of Georgia's Savannah River Ecology Laboratory (srel.uga.edu).