Evolution of The Horse - History of Research

History of Research

Indigenous modern horses died out in the New World at the end of the Pleistocene, about 12, 000 years ago, and thus were absent until the Spanish brought domestic horses from Europe, beginning in 1493. Escaped horses quickly established large wild herds. In the 1760s, the early naturalist Buffon suggested this was an indication of inferiority of the New World fauna, but later reconsidered this idea. William Clark's 1807 expedition to Big Bone Lick found "leg and foot bones of the Horses", which were included with other fossils sent to Thomas Jefferson and evaluated by the anatomist Caspar Wistar, but neither commented on the significance of this find.

The first equid fossil was found in the gypsum quarries in Montmartre, Paris in the 1820s. The tooth was sent to the Paris Conservatory, where it was identified by Georges Cuvier, who identified it as a browsing equine related to the tapir. His sketch of the entire animal matched later skeletons found at the site.

During the Beagle survey expedition, the young naturalist Charles Darwin had remarkable success with fossil hunting in Patagonia. On 10 October 1833 at Santa Fe, Argentina, he was "filled with astonishment" when he found a horse's tooth in the same stratum as fossil giant armadillos, and wondered if it might have been washed down from a later layer, but concluded this was "not very probable". After the expedition returned in 1836, the anatomist Richard Owen confirmed the tooth was from an extinct species, which he subsequently named Equus curvidens, and remarked, "This evidence of the former existence of a genus, which, as regards South America, had become extinct, and has a second time been introduced into that Continent, is not one of the least interesting fruits of Mr. Darwin's palæontological discoveries."

In 1848, a study On the fossil horses of America by Joseph Leidy systematically examined Pleistocene horse fossils from various collections, including that of the Academy of Natural Sciences and concluded at least two ancient horse species had existed in North America: Equus curvidens and another, which he named Equus americanus. A decade later, however, he found the latter name had already been taken and renamed it Equus complicatus. In the same year, he visited Europe and was introduced by Owen to Darwin.

The original sequence of species believed to have evolved into the horse was based on fossils discovered in North America in the 1870s by paleontologist Othniel Charles Marsh. The sequence, from Eohippus to the modern horse (Equus), was popularized by Thomas Huxley and became one of the most widely known examples of a clear evolutionary progression. The horse's evolutionary lineage became a common feature of biology textbooks, and the sequence of transitional fossils was assembled by the American Museum of Natural History into an exhibit that emphasized the gradual, "straight-line" evolution of the horse.

Since then, as the number of equid fossils has increased, the actual evolutionary progression from Eohippus to Equus has been discovered to be much more complex and multibranched than was initially supposed. The straight, direct progression from the former to the latter has been replaced by a more elaborate model with numerous branches in different directions, of which the modern horse is only one of many. George Gaylord Simpson in 1951 first recognized the modern horse was not the "goal" of the entire lineage of equids, it is simply the only genus of the many horse lineages to survive.

Detailed fossil information on the rate and distribution of new equid species has also revealed that the progression between species was not as smooth and consistent as was once believed. Although some transitions, such as that of Dinohippus to Equus, were indeed gradual progressions, a number of others, such as that of Epihippus to Mesohippus, were relatively abrupt in geologic time, taking place over only a few million years. Both anagenesis (gradual change in an entire population's gene frequency) and cladogenesis (a population "splitting" into two distinct evolutionary branches) occurred, and many species coexisted with "ancestor" species at various times. The change in equids' traits was also not always a "straight line" from Eohippus to Equus: some traits reversed themselves at various points in the evolution of new equid species, such as size and the presence of facial fossae, and only in retrospect can certain evolutionary trends be recognized.