Lemuriformes | |
---|---|
A sample of lemuriform diversity; [lower-alpha 1] 8 biological genera are depicted (from top, left to right): Lemur , Propithecus , Daubentonia , Varecia , Microcebus , Nycticebus , Loris , Otolemur . | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Primates |
Suborder: | Strepsirrhini |
Infraorder: | Lemuriformes Gray, 1821 |
Superfamilies | |
Lemuroidea |
Lemuriformes is the sole extant infraorder of primate that falls under the suborder Strepsirrhini. It includes the lemurs of Madagascar, as well as the galagos and lorisids of Africa and Asia, although a popular alternative taxonomy places the lorisoids in their own infraorder, Lorisiformes. [lower-alpha 1]
Lemuriform primates are characterized by a toothcomb, a specialized set of teeth in the front, lower part of the mouth mostly used for combing fur during grooming. [4]
Lemuriform origins are unclear and debated. American paleontologist Philip Gingerich proposed that lemuriform primates evolved from one of several genera of European adapids based on similarities between the front lower teeth of adapids and the toothcomb of extant lemuriforms; however, this view is not strongly supported due to a lack of clear transitional fossils. [5] Instead, lemuriforms may be descended from a very early branch of Asian cercamoniines or sivaladapids that immigrated to northern Africa. [6] [7]
Lemuriform phylogeny [8] [9] | ||||||||||||||||||||||||||||||||||||
|
Until discoveries of three 40-million-year-old fossil lorisoids ( Karanisia , Saharagalago , and Wadilemur ) in the El Fayum deposits of Egypt between 1997 and 2005, the oldest known lemuriforms had come from the early Miocene (~20 mya) of Kenya and Uganda. These newer finds demonstrate that lemuriform primates were present during the middle Eocene in Afro-Arabia and that the lemuriform lineage and all other strepsirrhine taxa had diverged before then. [10] [11] [12] Djebelemur from Tunisia dates to the late early or early middle Eocene (52 to 46 mya) and has been considered a cercamoniine, [13] but also may have been a stem lemuriform. [6] Azibiids from Algeria date to roughly the same time and may be a sister group of the djebelemurids. Together with Plesiopithecus from the late Eocene Egypt, the three may qualify as the stem lemuriforms from Africa. [7]
Molecular clock estimates indicate that lemurs and the lorisoids diverged in Africa during the Paleocene, approximately 62 mya. Between 47 and 54 mya, lemurs dispersed to Madagascar by rafting. [11] In isolation, the lemurs diversified and filled the niches often filled by monkeys and apes today. [14] In Africa, the lorises and galagos diverged during the Eocene, approximately 40 mya. [11] Unlike the lemurs in Madagascar, they have had to compete with monkeys and apes, as well as other mammals. [15]
Most of the academic literature provides a basic framework for primate taxonomy, usually including several potential taxonomic schemes. [16] Although most experts agree upon phylogeny, [17] many disagree about nearly every level of primate classification. [16]
Within Strepsirrhini, two common classifications include either two infraorders (Adapiformes and Lemuriformes) [18] or three infraorders (Adapiformes, Lemuriformes, Lorisiformes). [19] A less common taxonomy places the aye-aye (Daubentoniidae) in its own infraorder, Chiromyiformes. [20] In some cases, plesiadapiforms are included within the order Primates, in which case Euprimates is sometimes treated as a suborder, with Strepsirrhini becoming an infraorder, and the Lemuriformes and others become parvorders. [21] Regardless of the infraordinal taxonomy, crown strepsirrhines are composed of 10 families, three of which are extinct. [18] These three extinct families included the giant lemurs of Madagascar, [22] many of which died out within the last 1,000 years following human arrival on the island. [23]
2 infraorders [1] [2] | 3 infraorders [3] | 4 infraorders [24] |
---|---|---|
|
|
|
When Strepsirrhini is divided into two infraorders, the clade containing all toothcombed primates can be called "lemuriforms". [18] When it is divided into three infraorders, the term "lemuriforms" refers only to Madagascar's lemurs, [14] and the toothcombed primates are referred to as either "crown strepsirrhines" [25] or "extant strepsirrhines". [26] Confusion of this specific terminology with the general term "strepsirrhine", along with oversimplified anatomical comparisons and vague phylogenetic inferences, can lead to misconceptions about primate phylogeny and misunderstandings about primates from the Eocene, as seen with the media coverage of Darwinius . [27] Because the skeletons of adapiforms share strong similarities with those of lemurs and lorises, researchers have often referred to them as "primitive" strepsirrhines, [28] lemur ancestors, or a sister group to the living strepsirrhines. They are included in Strepsirrhini, [26] and are considered basal members of the clade. [29] Although their status as true primates is not questioned, the questionable relationship between adapiforms and other living and fossil primates leads to multiple classifications within Strepsirrhini. Often, adapiforms are placed in their own infraorder due to anatomical differences with lemuriforms and their unclear relationship. When shared traits with lemuriforms (which may or may not be synapomorphic) are emphasized, they are sometimes reduced to families within the infraorder Lemuriformes (or superfamily Lemuroidea). [30]
Strepsirrhini or Strepsirhini is a suborder of primates that includes the lemuriform primates, which consist of the lemurs of Madagascar, galagos ("bushbabies") and pottos from Africa, and the lorises from India and southeast Asia. Collectively they are referred to as strepsirrhines. Also belonging to the suborder are the extinct adapiform primates which thrived during the Eocene in Europe, North America, and Asia, but disappeared from most of the Northern Hemisphere as the climate cooled. Adapiforms are sometimes referred to as being "lemur-like", although the diversity of both lemurs and adapiforms does not support this comparison.
Prosimians are a group of primates that includes all living and extinct strepsirrhines, as well as the haplorhine tarsiers and their extinct relatives, the omomyiforms, i.e. all primates excluding the simians. They are considered to have characteristics that are more "primitive" than those of simians.
Haplorhini, the haplorhines or the "dry-nosed" primates, is a suborder of primates containing the tarsiers and the simians, as sister of the Strepsirrhini ("moist-nosed"). The name is sometimes spelled Haplorrhini. The simians include catarrhines, and the platyrrhines.
Tarsiiformes are a group of primates that once ranged across Europe, northern Africa, Asia, and North America, but whose extant species are all found in the islands of Southeast Asia. Tarsiers are the only living members of the infraorder; other members of Tarsiidae include the extinct Tarsius eocaenus from the Eocene, and Tarsius thailandicus from the Miocene. Two extinct genera, Xanthorhysis and Afrotarsius, are considered to be close relatives of the living tarsiers, and are generally classified within Tarsiiformes, with the former grouped within family Tarsiidae, and the latter listed as incertae sedis (undefined). Omomyids are generally considered to be extinct relatives, or even ancestors, of the living tarsiers, and are often classified within Tarsiiformes.
The simians, anthropoids, or higher primates are an infraorder of primates containing all animals traditionally called monkeys and apes. More precisely, they consist of the parvorders Platyrrhini and Catarrhini, the latter of which consists of the family Cercopithecidae and the superfamily Hominoidea.
Adapiformes is a group of early primates. Adapiforms radiated throughout much of the northern continental mass, reaching as far south as northern Africa and tropical Asia. They existed from the Eocene to the Miocene epoch. Some adapiforms resembled living lemurs.
Lorisoidea is a superfamily of nocturnal primates found throughout Africa and Asia. Members include the galagos and the lorisids. As strepsirrhines, lorisoids are related to the lemurs of Madagascar and are sometimes included in the infraorder Lemuriformes, although they are also sometimes placed in their own infraorder, LorisiformesGregory, 1915.
A toothcomb is a dental structure found in some mammals, comprising a group of front teeth arranged in a manner that facilitates grooming, similar to a hair comb. The toothcomb occurs in lemuriform primates, treeshrews, colugos, hyraxes, and some African antelopes. The structures evolved independently in different types of mammals through convergent evolution and varies both in dental composition and structure. In most mammals the comb is formed by a group of teeth with fine spaces between them. The toothcombs in most mammals include incisors only, while in lemuriform primates they include incisors and canine teeth that tilt forward at the front of the lower jaw, followed by a canine-shaped first premolar. The toothcombs of colugos and hyraxes take a different form with the individual incisors being serrated, providing multiple tines per tooth.
Bugtilemur is an extinct genus of Strepsirhine primate belonging to the adapiform family Ekgmowechashalidae.It is represented by only one species, B. mathesoni, which was found in the Chitarwata Formation of Pakistan.
Algeripithecus is an extinct genus of early fossil primate, weighing approximately 65 to 85 grams. Fossils have been found in Algeria dating from 50 to 46 million years ago.
Babakotia is an extinct genus of medium-sized lemur, or strepsirrhine primate, from Madagascar that contains a single species, Babakotia radofilai. Together with Palaeopropithecus, Archaeoindris, and Mesopropithecus, it forms the family Palaeopropithecidae, commonly known as the sloth lemurs. The name Babakotia comes from the Malagasy name for the indri, babakoto, to which it and all other sloth lemurs are closely related. Due to its mix of morphological traits that show intermediate stages between the slow-moving smaller sloth lemurs and the suspensory large sloth lemurs, it has helped determine the relationship between both groups and the closely related and extinct monkey 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.
Ekgmowechashala is an extinct genus of primate belonging to Adapiformes.
Lemurs were first classified in 1758 by Carl Linnaeus, and the taxonomy remains controversial today, with approximately 70 to 100 species and subspecies recognized, depending on how the term "species" is defined. Having undergone their own independent evolution on Madagascar, lemurs have diversified to fill many ecological 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 have become restricted to 10% of the island, or approximately 60,000 square kilometers (23,000 sq mi), and many face extinction. Concerns over lemur conservation have affected lemur taxonomy, since distinct species receive increased conservation attention compared to subspecies.
The sublingua ("under-tongue") is a muscular secondary tongue found below the primary tongue in tarsiers and living strepsirrhine primates, which includes lemurs and lorisoids. Although it is most fully developed in these primates, similar structures can be found in some other mammals, such as marsupials, treeshrews, and colugos. This "second tongue" lacks taste buds, and in lemuriforms, it is thought to be used to remove hair and other debris from the toothcomb, a specialized dental structure used to comb the fur during oral grooming.
Azibiidae is an extinct family of fossil primate from the late early or early middle Eocene from the Glib Zegdou Formation in the Gour Lazib area of Algeria. They are thought to be related to the living toothcombed primates, the lemurs and lorisoids, although paleoanthropologists such as Marc Godinot have argued that they may be early simians. It includes the genera Azibius and Algeripithecus, the latter of which was originally considered the oldest known simian, not a strepsirrhine.
Azibius is an extinct genus of fossil primate from the late early or early middle Eocene from the Glib Zegdou Formation in the Gour Lazib area of Algeria. They are thought to be related to the living toothcombed primates, the lemurs and lorisoids, although paleoanthropologists such as Marc Godinot have argued that they may be early simians. Originally described as a type of plesiadapiform, its fragmentary remains have been interpreted as a hyopsodontid, an adapid, and a macroscelidid. Less fragmentary remains discovered between 2003 and 2009 demonstrated a close relationship between Azibius and Algeripithecus, a fossil primate once thought to be the oldest known simian. Descriptions of the talus in 2011 have helped to strengthen support for the strepsirrhine status of Azibius and Algeripithecus, which would indicate that the evolutionary history of lemurs and their kin is rooted in Africa.
Djebelemur is an extinct genus of early strepsirrhine primate from the late early or early middle Eocene period from the Chambi locality in Tunisia. Although they probably lacked a toothcomb, a specialized dental structure found in living lemuriforms, they are thought to be a related stem group. The one recognized species, Djebelemur martinezi, was very small, approximately 100 g (3.5 oz).
Djebelemuridae is an extinct family of early strepsirrhine primates from Africa. It consists of five genera. The organisms in this family were exceptionally small, and were insectivores. This family dates to the early to late Eocene. Although they gave rise to the crown strepsirrhines, which includes today's lemurs and lorisoids, they lacked the toothcomb that identifies that group.
Plesiopithecus is an extinct genus of early strepsirrhine primate from the late Eocene.