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We explore how social behaviors shape direct and indirect pathogen transmission of the emerging\n                    <jats:styled-content style=\"fixed-case\">\n                      <jats:italic>Mycobacterium tuberculosis<\/jats:italic>\n                    <\/jats:styled-content>\n                    complex (MTBC) pathogen\n                    <jats:italic>M. mungi<\/jats:italic>\n                    in banded mongooses (\n                    <jats:styled-content style=\"fixed-case\">\n                      <jats:italic>Mungos mungo<\/jats:italic>\n                    <\/jats:styled-content>\n                    ). Using empirical behavioral data from a banded mongoose population in Uganda, we constructed social network models that incorporate observed intra\u2010group contacts and simulated inter\u2010group scent marking scenarios. These models show that indirect transmission can play a pivotal role, particularly when per\u2010contact transmissibility is low. Simulations predicted increased population\u2010level spread via socially directed, environmentally mediated pathways, while population\u2010level invasion potential remained relatively unchanged. These results underscore the need to incorporate all transmission modes when modeling disease spread in social networks. We further emphasize the importance of considering the temporal dynamics of indirect transmission, including the persistence of both olfactory secretion and infecting pathogens deposited in the environment. For mixed transmission systems, fully accounting for both direct and indirect transmission mechanisms may be essential for characterizing disease spread, even when direct transmission pathways dominate. Our findings suggest that pathogens that exploit socially directed environmental transmission pathways may gain an evolutionary advantage, circumventing social barriers thought to have evolved to constrain pathogen spread.\n                  <\/jats:p>","DOI":"10.1002\/ece3.73719","type":"journal-article","created":{"date-parts":[[2026,5,28]],"date-time":"2026-05-28T01:31:32Z","timestamp":1779931892000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Networks of Indirect Contact Promote Spread of an Environmentally Transmitted Pathogen in a Highly Social Species"],"prefix":"10.1002","volume":"16","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-8822-9410","authenticated-orcid":false,"given":"Jack R.","family":"Leitch","sequence":"first","affiliation":[{"name":"Department of Fish and Wildlife Conservation Virginia Tech  Blacksburg Virginia USA"},{"name":"CARACAL, Centre for Conservation of African Resources: Animals, Communities, and Land Use  Kasane Botswana"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0712-8301","authenticated-orcid":false,"given":"Neil R.","family":"Jordan","sequence":"additional","affiliation":[{"name":"Centre for Ecosystem Science, University of New South Wales, School of Biological, Earth and Environmental Sciences  Sydney New South Wales Australia"},{"name":"Taronga Wildlife Reproductive Centre Taronga Conservation Society Australia  Dubbo New South Wales Australia"},{"name":"Botswana Predator Conservation  Maun Botswana"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"John M.","family":"Drake","sequence":"additional","affiliation":[{"name":"Odum School of Ecology University of Georgia  Athens Georgia USA"},{"name":"Center for the Ecology of Infectious Diseases University of Georgia  Athens Georgia USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Michael A.","family":"Cant","sequence":"additional","affiliation":[{"name":"University of Exeter  Penryn UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7338-5341","authenticated-orcid":false,"given":"Kathleen A.","family":"Alexander","sequence":"additional","affiliation":[{"name":"Department of Fish and Wildlife Conservation Virginia Tech  Blacksburg Virginia USA"},{"name":"CARACAL, Centre for Conservation of African Resources: Animals, Communities, and Land Use  Kasane Botswana"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,5,27]]},"reference":[{"key":"e_1_2_10_2_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.anbehav.2005.10.001"},{"key":"e_1_2_10_3_1","doi-asserted-by":"publisher","DOI":"10.1128\/mBio.00281-16"},{"key":"e_1_2_10_4_1","doi-asserted-by":"publisher","DOI":"10.1079\/9781780643960.0386"},{"key":"e_1_2_10_5_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0019896"},{"key":"e_1_2_10_6_1","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.ecolsys.34.030102.151725"},{"key":"e_1_2_10_7_1","doi-asserted-by":"publisher","DOI":"10.1080\/17513758.2010.503376"},{"key":"e_1_2_10_8_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0005016"},{"key":"e_1_2_10_9_1","doi-asserted-by":"publisher","DOI":"10.1017\/S0950268810000816"},{"key":"e_1_2_10_10_1","doi-asserted-by":"publisher","DOI":"10.1006\/anbe.1999.1279"},{"key":"e_1_2_10_11_1","doi-asserted-by":"publisher","DOI":"10.1590\/S0074-02761984000400021"},{"key":"e_1_2_10_12_1","doi-asserted-by":"publisher","DOI":"10.1111\/ele.12917"},{"key":"e_1_2_10_13_1","doi-asserted-by":"publisher","DOI":"10.1098\/rspb.2009.1775"},{"key":"e_1_2_10_14_1","doi-asserted-by":"publisher","DOI":"10.1002\/ecs2.4434"},{"key":"e_1_2_10_15_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cobeha.2016.09.006"},{"key":"e_1_2_10_16_1","doi-asserted-by":"publisher","DOI":"10.1007\/s10393-014-0912-4"},{"key":"e_1_2_10_17_1","doi-asserted-by":"publisher","DOI":"10.4061\/2011\/765430"},{"key":"e_1_2_10_18_1","doi-asserted-by":"publisher","DOI":"10.1098\/rstb.2016.0086"},{"key":"e_1_2_10_19_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469-7998.1984.tb05050.x"},{"key":"e_1_2_10_20_1","doi-asserted-by":"publisher","DOI":"10.1007\/978-1-4757-4716-4_3"},{"key":"e_1_2_10_21_1","volume-title":"Analysis and Visualisation of Time Series Data on Networks With Pathpy. 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