new ideas about the origins, diversity, and distribution of these animals. The number of Malagasy subfossil sites containing subfossil lemurs has increased significantly since the mid-20th century. At that time, subfossil lemurs had only been found in the center, south, and southwest of the island. Since then, only the eastern rainforests have not been represented, and paleodistributions are now known for both extinct and living species around most of the island. Large quantities of subfossil lemur remains have been found in caves, marshes, and streambank sites in drier regions. The subfossil sites are clustered together geographically and are recent in age, mostly dating between 2,500 and 1,000 years old, with a few spanning back into the last glaciation, which ended 10,000 years ago. Extinction At least 17 species of giant subfossil lemur vanished during the Holocene, with all or most extinctions happening after the colonization of Madagascar by humans around 2,000 years ago. Madagascar's megafauna included not only giant lemurs, but also elephant birds, giant tortoises, several species of Malagasy hippopotamuses, Cryptoprocta spelea (a "giant fossa"), large crocodiles (Voay robustus), and Plesiorycteropus, a unique digging mammal, all of which died out during the same period. Madagascar's megafaunal extinctions were among the most severe for any continent or large island, with all endemic wildlife over disappearing, totaling approximately 25 species. The most severely impacted lemurs were generally large and diurnal, particularly the clade containing the living indriids and extinct sloth lemurs. Although only the indriids are alive today and represent only a small percentage of the living lemur species, this clade collectively contained the majority of the extinct giant lemur species. By region, the Central Highlands lost the greatest number of lemur species. It has lost nearly all of its woodland habitat, but some lemur species still survive in isolated forest patches. Lemur diversity is tightly linked with plant diversity, which in turn decreases with increased forest fragmentation. In extreme cases, treeless sites such as the town of Ampasambazimba from the central region no longer support any of the lemur species represented in their subfossil record. Other locations no longer have giant subfossil lemurs, yet they still maintain forested habitat that could support them. Even though the giant lemurs have disappeared from these locations, while the smaller species survive in the forest patches that remain, the subfossil remains indicate that the living species used to be more widespread and coexisted with