leaves. The rest of its skeleton (postcrania) was much more robust and their vertebrae had distinctly different features. Phylogeny Determining the phylogeny of subfossil lemurs has been problematic because studies of morphology, developmental biology, and molecular phylogenetics have sometimes yielded conflicting results. All studies agree that the family Daubentoniidae (including the giant aye-aye) diverged first from the other lemurs at least 60 million years ago. The relationship between the remaining families has been less clear. Morphological, developmental, and molecular studies have offered support for lumping the four sloth lemur genera of the family Palaeopropithecidae with the family Indriidae (including the indri, sifakas, and woolly lemurs). The placement of family Megaladapidae has been more controversial, with similarities in teeth and skull features suggesting a close relationship with family Lepilemuridae (sportive lemurs). Molecular data, instead, indicate a closer relationship to family Lemuridae. Likewise, a relationship between family Archaeolemuridae and family Lemuridae has been suggested, based on morphological and developmental traits, yet molar morphology, the number of teeth in the specialized toothcomb, and molecular analysis support a closer relationship with the indriid–sloth lemur clade. Other subfossil lemurs, including the giant aye-aye and Pachylemur, are more easily placed due to strong similarities with existing lemurs (the aye-aye and ruffed lemurs, respectively). Living species Subfossil sites in Madagascar have yielded the remains of more than just extinct lemurs. Extant lemur remains have also been found, and radiocarbon dating has demonstrated that both types of lemur lived at the same time. In some cases living species are locally extinct for the area in which their subfossil remains were found. Because subfossil sites are found across most of the island, with the most notable exception being the eastern rainforest, both paleocommunity composition and paleodistributions can be determined. Geographic ranges have contracted for numerous species, including the indri, greater bamboo lemur, and ruffed lemurs. For instance, subfossil remains of the indri have been found in marsh deposits near Ampasambazimba in the Central Highlands and in other deposits in both central and northern Madagascar, demonstrating a much larger range than the small region on the east coast that it currently occupies. Even the greater bamboo lemur, a critically endangered species restricted to a small portion of the south-central eastern rainforest, has undergone significant range contraction since the mid-Holocene, with subfossil remains from Ankarana Massif in the far north of Madagascar dating to 2565 BCE ± 70 years. Combined with