A newly published paper postulates that ungulate evolution has been shaped by two major ecological shifts. Tectonic shifts and global climate change have been the drivers of major biota turnover amongst large herbivorous mammals.
Ungulates are hoofed mammals. This extremely diverse clade is subdivided into two orders, the Artiodactyla (even-toed) mammals that walk on two of their five toes and the Perissodactyla (odd-toed) mammals that walk on one or three toes. Molecular data led to a reclassification at the turn of the century with the establishment of a larger clade the Euungulata.
- Typical artiodactyls – ruminants such as cattle, goats, sheep, llamas, camels, hippos, giraffes, deer, pigs.
- Typical perissodactyls – rhinos, horses, tapirs.
It should be noted that the odd-toed ungulates were much more diverse in the past. Brontotheres and knuckle-walking chalicotheres are extinct members of the Perissodactyla.

A wonderful prehistoric animal model. A 1:20 scale replica of the chalicothere Moropus – an extinct perissodactyl.
The picture above shows a typical chalicothere. It is a 1:20 scale replica of Moropus from CollectA. To view the range of CollectA scale prehistoric animal models: CollectA Deluxe Prehistoric Life.
Large Herbivores Shaping the Landscape
Large herbivores have shaped the Earth’s landscapes for the last forty million years. A new study led by researchers from the University of Gothenburg in collaboration with scientists from Spain and the Museum für Naturkunde Berlin examines how these animals reacted to dramatic environmental changes. The study, published in “Nature Communications” demonstrates that ecosystems managed to remain stable despite the extinction of many families.
Ungulates and other large mammals such as elephants are key ecosystem engineers. The decline of large herbivores threatens entire habitats. These large herbivores shape landscapes and promote biodiversity. Current extinction rates, often referred to as the sixth mass extinction could lead to a loss of key taxa.
Dr Fernando Blanco, a visiting scientist at the Museum für Naturkunde Berlin led the research. The fossils of over three thousand large herbivores from the past sixty million years to more recent times were studied.
Dr Blanco commented:
“We found that these ecosystems have remained surprisingly stable over long periods of time, even though species were added and others became extinct. Twice in the last sixty million years, however, the environmental pressure was so great that the entire system was globally reorganised.”
The End of the Tethys
The team discovered that the first major reorganisation of ecosystems took place around twenty-one million years ago. This was during the Miocene Epoch. The Tethys Sea closed and this created a landbridge between Eurasia and Africa. A mass migration of species occurred. The ancestors of many extant ungulates moved into new habitats. This had a profound effect on ungulate evolution.
A second major reorganisation of ecosystems occurred approximately ten million years ago. This was during the Tortonian stage of the Miocene. Global cooling led to a massive reduction in forests and the spread of grasslands. This led to a dramatic increase in grazing species and a gradual disappearance of many forest-dwelling species. The researchers postulate that this was the beginning of a sustained decline in the functional diversity of large, herbivorous mammals. This has led to a decrease in their influence over the planet’s ecosystems. Despite the extinctions, the researchers found that the basic ecological structure of herbivore communities remained remarkably stable. Ecosystems remained stable even when many iconic taxa died out such as mastodons, mammoths and giant rhinoceroses.

A typical artiodactyl – a Cape buffalo (Syncerus caffer) photographed in Tanzania. Scientists reflect on ungulate evolution and the impact of large herbivores on the Earth’s ecosystems. Picture credit: Juan López Cantalapiedra.
Picture credit: Juan López Cantalapiedra
Co-author of the study, Dr Ignacio A. Lazagabaster from CENIEH (Centro National de Investigacion Sobre la Evolucion Humana, Spain) commented:
“It’s like a football team changing players during a game without the line-up changing significantly. New species came into play and the communities changed, but the new players fulfilled similar ecological tasks – as a result, the overall structure remained stable.”
The Resilience of Large Herbivores is Not Guaranteed
The research team remarks on the resilience of large mammals to survive glaciation, global cooling and other environmental crises. However, many large taxa are extremely vulnerable today.
Fellow co-author of the study Dr Juan L. Cantalapiedra from the Spanish MNCN (Museo Nacional de Ciencias Naturales) stated:
“Our results show how enormously adaptable ecosystems can be. But there are limits. If we continue to lose species and their ecological roles on such a massive scale as in the present, we could soon reach a third global tipping point – and we humans are actively contributing to this.”
Everything Dinosaur acknowledges the assistance of the Museum für Naturkunde Berlin in the compilation of this article.
The scientific paper: “Two major ecological shifts shaped 60 million years of ungulate faunal evolution” by Fernando Blanco, Ignacio A. Lazagabaster, Óscar Sanisidro, Faysal Bibi, Nicola S. Heckeberg, María Ríos, Bastien Mennecart, María Teresa Alberdi, Jose Luis Prado, Juha Saarinen, Daniele Silvestro, Johannes Müller, Joaquín Calatayud and Juan L. Cantalapiedra published in Nature Communications.
For models of extinct mammals and other prehistoric animals: Prehistoric Animal Models and Toys.
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