Squirrel In Action

The living squirrels are divided into five subfamilies, with about 50 genera and nearly 280 species. The oldest squirrel fossil, Hesperopetes, dates back to the Chadronian (Late Eocene, about 40–35 million years ago), and is similar to modern flying squirrels.
During the latest Eocene to the Miocene, there were a variety of squirrels which cannot be assigned with certainty to any living lineage. At least some of these probably were variants of the oldest, basal "proto-squirrels" (in the sense that they lacked the full range of living squirrels' autapomorphies). The distribution and diversity of such ancient and ancestral forms suggests that the squirrels as a group might have originated in North America.
Apart from these sometimes little-known fossil forms, the phylogeny of the living squirrels is fairly straightforward. There are three main lineages, one comprising the Ratufinae (Oriental giant squirrels). These contain a mere handful of living species in tropical Asia. The Neotropical Pygmy Squirrel of tropical South America is the sole living member of the Sciurillinae. The third lineage is by far the largest and contains all other subfamilies; it has a near-cosmopolitan distribution. This further supports the hypothesis that the common ancestor of all squirrels living and fossil lived in North America, as these three most ancient lineages seem to have radiated from there – if squirrels had originated in Eurasia for example, one would expect quite ancient lineages in Africa, but African squirrels seem to be of more recent origin.
The main group of squirrels also can be split up in three, which yields the remaining subfamilies. The Sciurinae contains the flying squirrels (Pteromyini) and the Sciurini, which among others contains the American tree squirrels; the former have often been considered a separate subfamily but are now seen as a tribe of the Sciurinae. The pine squirrels (Tamiasciurus) on the other hand are usually included with the main tree squirrel lineage, but appear to be about as distinct as the flying squirrels; hence they are sometimes considered a distinct tribe, Tamiasciurini.