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Proterocheris is another ancient fossil turtle that lived at the same time as Proganochelys. Proterocheris has many features that suggest that it is a side-necked turtle. If this is true, the two major taxonomic groups of living turtles, suborders Pleurodira (side-necked turtles) and Cryptodira (hidden necks), had their origins some 230 million years ago during the Triassic Period (251.9 million to 201.3 million years ago) at the latest, making turtles an extremely ancient group. Proterocheris and two later-appearing Triassic genera are likely not true side-necks but turtles that share some pleurodire characteristics. Unquestionable pleurodires do not appear until the Early Cretaceous (about 145 million to 100.5 million years ago), and the first modern side-neck families do not appear until the Late Cretaceous (100.5 million to 66 million years ago).In tracing back the history of the other turtle suborder, Cryptodira, Kayentachelys aprix of the Late Jurassic (some 150 million years ago) is almost assuredly a cryptodire; it is also the oldest known North American turtle. Other cryptodires are known from the Late Jurassic, although they are not representative of existing families. The largest known turtles are Archelon ischyros, a sea turtle species that lived during the Late Cretaceous and reached a length of about 3.5 metres (12 feet), and Stupendymys geographicus, a side-necked freshwater turtle that lived during the Miocene and whose shell alone measured 2.4 metres (about 8 feet) long. Softshell turtles (family Trionychidae) are the first modern turtles found in the fossil record, appearing in the Cretaceous Period. The oldest sea turtle (Santanachelys gaffneyi) is known from the mid-Cretaceous. It is a member of the Protostegidae, a likely sister group of modern leatherback sea turtles. S. gaffneyi had a streamlined shell of about 1.5 metres (5 feet) and forelimbs well along the evolutionary path to becoming flippers.

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Turtles have adapted to a remarkable variety of environments, but the greatest number of species occur in southeastern North America and South Asia. In both areas, most species are aquatic, living in bodies of water ranging from small ponds and bogs to large lakes and rivers. A few are strictly terrestrial (tortoises), and others divide their time between land and water. Although turtles as a group are broadly distributed, each species has a preferred habitat and is seldom found elsewhere. For example, both the gopher tortoise (Gopherus polyphemus) and the Eastern box turtle (Terrapene carolina) live in the southern United States and are equally terrestrial, but they are not usually found together, as the box turtle prefers moist forest and the gopher tortoise open woodlands on sand ridges. The eastern mud turtle (Kinosternon subrubrum) is commonly considered an aquatic turtle, yet it spends the summer months in dormancy, estivating beneath vegetation in woodlands adjacent to its pond and stream habitats. The alligator snapping turtle (Macrochelys temmincki) lives in the deep, slow-moving streams and backwaters of the U.S. Gulf Coast. Map turtles (Graptemys), on the other hand, select the faster-flowing waters of those same streams. The saltwater terrapin (Malaclemys terrapin) lives in brackish coastal estuaries and marshes from Cape Cod, Massachusetts, to Padre Island, Texas. In some instances, juvenile sea turtles share these estuaries; larger and older sea turtles swim offshore in coastal waters to the mid-continental shelf.