Pair living emerged early in placental mammals

S Sam F. Walmsley (Department of Biology, Dalhousie University) C Charlotte-Anaïs Olivier (Institut Pluridisciplinaire Hubert Curien, CNRS, Department of Ethology and Evolutionary Physiology, University of Strasbourg) A Adrian V. Jaeggi (Human Ecology Group, Institute of Evolutionary Medicine, University of Zürich) L Lindelani Makuya (Institut Pluridisciplinaire Hubert Curien, CNRS, Department of Ethology and Evolutionary Physiology, University of Strasbourg) J Jordan S. Martin (Human Ecology Group, Institute of Evolutionary Medicine, University of Zürich) L Loren D. Hayes (Department of Biology, Geology, and Environmental Science, University of Tennessee at Chattanooga) C Carsten Schradin (Institut Pluridisciplinaire Hubert Curien, CNRS, Department of Ethology and Evolutionary Physiology, University of Strasbourg)

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.

Article Details

Volume / Issue Vol. 123, Issue 31
Published August 04, 2026
ISSN 0027-8424
Publisher National Academy of Sciences

Authors (7)

S

Sam F. Walmsley

Department of Biology, Dalhousie University

C

Charlotte-Anaïs Olivier

Institut Pluridisciplinaire Hubert Curien, CNRS, Department of Ethology and Evolutionary Physiology, University of Strasbourg

A

Adrian V. Jaeggi

Human Ecology Group, Institute of Evolutionary Medicine, University of Zürich

L

Lindelani Makuya

Institut Pluridisciplinaire Hubert Curien, CNRS, Department of Ethology and Evolutionary Physiology, University of Strasbourg

J

Jordan S. Martin

Human Ecology Group, Institute of Evolutionary Medicine, University of Zürich

L

Loren D. Hayes

Department of Biology, Geology, and Environmental Science, University of Tennessee at Chattanooga

C

Carsten Schradin

Institut Pluridisciplinaire Hubert Curien, CNRS, Department of Ethology and Evolutionary Physiology, University of Strasbourg