The first non-mammalian synapsid embryo from the Triassic of South Africa

Oviparity was likely the plesiomorphic reproductive condition for non-mammalian Synapsida, the stem-mammal group. Yet, despite nearly two centuries of research, no definitive fossil eggs of late Palaeozoic or early Mesozoic synapsids have been discovered. Here, three perinate specimens of the dicynodont genus Lystrosaurus from the Early Triassic of the South African Karoo Basin are examined using high-resolution CT and synchrotron scanning. One specimen, NMQR 3636, displays a tightly curled posture suggestive of an in ovo position and completely lacks tusks. Crucially, the lower jaw symphysis remains unfused-a developmental trait found only in pre-hatching embryos of modern birds and turtles. No calcified eggshell is preserved, so the egg might have been soft and leathery. The large size of the reconstructed egg suggests a precocial, non-milk-feeding developmental strategy. As a non-cynodont synapsid, Lystrosaurus offers a rare and valuable glimpse into reproductive biology far removed from the mammalian crown group. Unlike the more derived, mammal-like cynodont Kayentatherium, whose egg size aligns with lactation, Lystrosaurus anchors the plesiomorphic condition deep within Synapsida. Its reproductive strategy may have played a crucial role in its resilience and ecological dominance following the end-Permian mass extinction.