Evolution of lemurs

Lemurs, primates belonging to the suborder Strepsirrhini which branched off from other primates less than 63 million years ago, evolved on the island of Madagascar, for at least 40 million years. They share some traits with the most basal primates, and thus are often confused as being ancestral to modern monkeys, apes, and humans. Instead, they merely resemble ancestral primates.

Lemurs are thought to have evolved during the Eocene or earlier, sharing a closest common ancestor with lorises, pottos, and galagos (lorisoids). Fossils from Africa and some tests of nuclear DNA suggest that lemurs made their way to Madagascar between 40 and 52 mya. Other mitochondrial and nuclear DNA sequence comparisons offer an alternative date range of 62 to 65 mya. An ancestral lemur population is thought to have inadvertently rafted to the island on a floating mat of vegetation, although hypotheses for land bridges and island hopping have also been proposed. The timing and number of hypothesized colonizations has traditionally hinged on the phylogenetic affinities of the aye-aye, the most basal member of the lemur clade.

Having undergone their own independent evolution on Madagascar, lemurs have diversified to fill many niches normally filled by other types of mammals. They include the smallest primates in the world, and once included some of the largest. Since the arrival of humans approximately 2,000 years ago, lemurs are now restricted to 10% of the island, or approximately 60,000 square kilometers (23,000 square miles), with many facing extinction.

Lemurs are primates belonging to the suborder Strepsirrhini. Like other strepsirrhine primates, such as lorises, pottos, and galagos, they share ancestral traits with early primates. In this regard, lemurs are popularly confused with ancestral primates; however, lemurs did not give rise to monkeys and apes, but evolved independently on Madagascar.

Primates first evolved sometime between the Middle Cretaceous and the early Paleocene periods on either the supercontinent of Laurasia or in Africa. According to molecular clock studies, the last common ancestor of all primates dates to around 79.6 mya, although the earliest known fossil primates are only 54–55 million years old. The closest relatives of primates are the extinct plesiadapiforms, the modern colugos (commonly and inaccurately named "flying lemurs"), and treeshrews. Some of the earliest known true primates are represented by the fossil groups Omomyidae, Eosimiidae, and Adapiformes.

The relationship between known fossil primate families remains unclear. A conservative estimate for the divergence of haplorhines (tarsiers, monkeys, apes, and humans) and strepsirrhines is 58 to 63 mya. A consensus is emerging that places omomyids as a sister group to tarsiers, eosimiids as a stem group to simians (non-tarsier haplorhines), and Djebelemur, an African genus likely to be related to an early Asian branch of cercamoniine adapiforms, as a stem group to modern strepsirrhines, including lemurs. In 2009, a highly publicized and scientifically criticized publication proclaimed that a 47-million-year-old adapiform fossil, Darwinius masillae, demonstrated both adapiform and simian traits, making it a transitional form between the prosimian and simian lineages. Media sources inaccurately dubbed the fossil as a "missing link" between lemurs and humans.

Lemurs were traditionally thought to have evolved during the Eocene (55 to 37 mya) based on the fossil record, although molecular tests suggest the Paleocene (66 to 56 mya) or later. Until recently, they were thought to have descended directly from the diverse group of adapiforms due to several shared postcranial traits, as well as long snouts and small brains. Although adapiforms also had lemur-like auditory bullae, a prosimian characteristic, they had smaller brains and longer snouts than lemurs. There are also several other morphological differences. Most noticeably, adapiforms lack a key derived trait, the toothcomb, and possibly the toilet-claw, found not only in extant (living) strepsirrhines but also in tarsiers. Unlike lemurs, adapiforms exhibited a fused mandibular symphysis (a characteristic of simians) and also possessed four premolars, instead of three or two.

Comparative studies of the cytochrome b gene, which are frequently used to determine phylogenetic relationships among mammals—particularly within families and genera—have been used to show that lemurs share common ancestry with lorisoids. This conclusion is also corroborated by the shared strepsirrhine toothcomb, an unusual trait that is unlikely to have evolved twice. If adapiforms were the ancestors of the living strepsirrhines, then the last common ancestor of modern strepsirrhines would have to predate the early Eocene, a view supported by molecular phylogenetic studies by Anne D. Yoder and Ziheng Yang in 2004, which showed that lemurs split from lorises approximately 62 to 65 mya. These dates were confirmed by more extensive tests by Julie Horvath et al. in 2008. These molecular studies also showed that lemuroids diversified before the modern lorisoids. Using a more limited data set and only nuclear genes, another study in 2005 by Céline Poux et al. dated the split between lemurs and lorises at 60 mya, lemur diversification at 50 mya, and the lemur colonization of Madagascar somewhere between these two approximate dates. However, the 2003 discovery of fossil lorisoids at the Fayum Depression in Egypt pushed the date of lorisoid divergence back to the Eocene, matching the divergence dates predicted by Yoder and Horvath.