Primate Behavior and Ecology Open access Peer reviewed

Pair living emerged early in placental mammals

Sam F. Walmsley, Charlotte‐Anaïs Olivier, Adrian V. Jaeggi, Lindelani Makuya and 3 more

Proceedings of the National Academy of Sciences | Jul 28, 2026

Abstract

Abstract

It is widely assumed that the first placental mammals were solitary. Here, we tested this assumption and which ecological and life history factors are related to mammalian social organization. We compiled a comprehensive database by reviewing over 14,000 primary peer-reviewed publications on the social organization of the 5,390 extant placental mammal species. We found empirical data for 726 species across 1,473 populations: 353 populations were primarily solitary, 250 populations pair living, and 870 populations group living. Notably, 367 species (51%) and 583 populations (40%) exhibited intraspecific variation in social organization. Using a Bayesian phylogenetic framework that incorporates intraspecific variation, we show that the ancestral placental mammal was mainly but not exclusively solitary, displaying intraspecific variation. Pair living was estimated to account for approximately 26% of ancestral social organization. Phylogeny explained most of the variation between species, while life history and ecology were less important. Our analysis found that the ecology and life history of extant solitary and pair-living mammals is very similar and different from those of group-living mammals. Our results revise prevailing theories of mammalian social evolution, revealing that pair living emerged early, shifting the focus of studying mammalian social evolution from the origins of pair living to the origins of group living.

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Authors

Researchers on this paper

Sam F. Walmsley

first | Dalhousie University | ORCID 0000-0001-8577-6884

Charlotte‐Anaïs Olivier

middle | Centre National de la Recherche Scientifique | ORCID 0000-0003-3382-6354

Adrian V. Jaeggi

middle | University of Zurich | ORCID 0000-0003-1695-0388

Lindelani Makuya

middle | Centre National de la Recherche Scientifique | ORCID 0000-0002-2986-1627

Jordan S. Martin

middle | Swiss Federal Institute of Aquatic Science and Technology | ORCID 0000-0001-8704-6076

Loren D. Hayes

middle | University of Tennessee at Chattanooga | ORCID 0000-0003-0713-416X

Carsten Schradin

last | Centre National de la Recherche Scientifique | ORCID 0000-0002-2706-2960

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Citation

BibTeX

@article{Walmsley2026Pair,
  title = {Pair living emerged early in placental mammals},
  author = {Sam F. Walmsley and Charlotte‐Anaïs Olivier and Adrian V. Jaeggi and Lindelani Makuya and Jordan S. Martin and Loren D. Hayes and Carsten Schradin},
  journal = {Proceedings of the National Academy of Sciences},
  year = {2026},
  doi = {10.1073/pnas.2528458123},
  url = {https://doi.org/10.1073/pnas.2528458123}
}

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