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496: 401:, the canines erupt after the incisors. The crowns of the incisors are also angled in the direction of the forward tilt, and the crowns of both the incisors and canines are elongated and compressed side-to-side. The apical ridge, following along the front edges of the toothcomb teeth, is V-shaped in most lemuriforms, tapering off from the midline. As a result of this dental reconfiguration, the upper and lower incisors do not contact one another, and often the upper incisors are reduced or lost completely. 276: 405: 99: 952:
bacterial growth and tooth decay if used for exudate feeding. Supporting this, reduced interdental spacing is found in exudate feeding lemuriforms. Furthermore, the canine included in the toothcomb provides additional interdental spacing for fur combing. Even the behavior of young lemuriforms suggests that grooming plays a more important role in the use of the toothcomb than food procurement.
450:(Indriidae), the toothcomb is less procumbent and consists of four teeth instead of six. The indriid toothcomb is more robust and wider, with shorter incisors, wider spaces between the teeth (interdental spaces), and a broader apical ridge. It is unclear whether this four-toothed toothcomb consists of two pairs of incisors or one pair of incisors and one pair of canines. In 526:" ("are real combs"). More than 100 years later, the grooming function was questioned since it was difficult to observe and the interdental spaces were thought to be too small for fur. Observations later showed the teeth were used for that purpose and that immediately after grooming, hair may be found trapped in the teeth, but is removed by the sublingua later. 31: 580:, which may account for the size reduction of the poorly studied upper incisors. The toothcomb may provide pressure to stimulate glandular secretions which are then spread through the fur. Furthermore, the size reduction of the upper incisors may create a gap between the teeth (interincisal diastema) that connects the 1021:
toothcomb, homologous with that of lemuriforms. However, studies of tarsier dentition showed that only some tarsier species exhibit the characteristic striations on their incisors, yet all the species examined were reported to use their front teeth when grooming, suggesting the presence or absence of
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Indriids such as the sifakas use their toothcombs to gouge bark or dead wood (bark-prising), which is done prior to scent-marking with the gland on their chest. The more robust structure of their toothcomb is thought to help it withstand the compressive forces experienced during regular bark-prising.
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The toothcomb of treeshrews is like the lemuriform toothcomb in that it uses interdental spaces to form the comb tines, but only two of its three pairs of lower incisors are included in the toothcomb and the canines are also excluded. The lateral two incisors in the toothcomb are generally larger. In
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serrated incisors are kept clean using the front of the tongue, which is serrated to match the serrations of the incisors. Similarly, the hyracoid toothcomb consists of incisors with multiple tines, called "pectinations". In contrast to the colugos, the size and shape of the tines are more uniform.
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lemurs (particularly fork-marked lemurs and the hairy-eared dwarf lemur) and galagos, which demonstrate tooth-scraping of plant exudates, as well as sifakas, which practice bark-prising. Each of these were considered "primitive" forms among the living strepsirrhines, suggesting the first lemuriforms
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The ancestral condition of the anterior dentition on the lower jaw, based on Eocene primate fossils, suggests that earliest primates had lacked a differentiated toothcomb. Most fossil strepsirrhines lacked the stereotypic lemuriform toothcomb. Collectively, early strepsirrhine primates are known as
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In colugos, the toothcomb has a completely different structure. Instead of individual incisors and canine teeth being finely spaced to act like the teeth of a comb, the biting edge of the four incisors have become serrated with as many as 15 tines each, while the canine acts more like a molar. These
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that shaped the original lemuriform toothcomb has been a topic of considerable debate since the 1970s. Evidence can be seen as supporting a grooming function, food procurement function, or both. In the early 1900s, there was less debate. Grooming was seen as the primary function since primates lack
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following the toothcomb are usually shaped like typical canine teeth (caniniform) and assume their function. These premolars are commonly confused with canines. Normally the true canines in the lower jaw sit in front of the upper canines, and in toothcombed primates, the caniniform premolars rest
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As a homologous structure in lemuriforms, the toothcomb serves variable biological roles, despite its superficially stereotypic shape and appearance. It is primarily used as a toiletry device or grooming comb. Additionally, some species use their toothcomb for food procurement or to gouge tree
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In contrast, the grooming hypothesis emphasized that all lemuriforms use their toothcombs for grooming, and long, thin teeth are poorly suited for the mechanical stress of gouging and exudate feeding. Also the interdental spaces seen in most lemuriforms favor fur combing and would also promote
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group (containing many but not all of the descendants of the last common ancestor of the group's members) because the lemuriforms are assumed to have evolved from one of several groups of adapiforms. In terms of ecology, the evolution of the toothcomb is assumed to have required a
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on each foot to compensate. Grooming—in the form of fur-combing—is generally considered the primary function and original role of the lemuriform toothcomb, and subsequent changes in morphology across multiple lineages have altered its function and obscured its original function.
938:). If feeding ecology could have such profound effects on the shape of the anterior dentition, then convergent evolution might explain the similarities seen between the compressed lower incisors of the lemuriform toothcomb and the exudate feeding adaptations in the genus 748:
has suggested this foreshadowed the development of the lemuriform toothcomb. However, no lemuriform toothcomb has been found in the fossil record of the Eocene, and the European adapid lower jaws from that time did not resemble the derived state seen in lemuriforms.
466:(Palaeopropithecidae), the toothcomb was lost and the incisors and canines resumed a typical configuration in the front of the mouth. The aye-aye also lost its toothcomb, replacing it with continually growing (hypselodont) front teeth, similar to the incisors of 553:) does not. The toothcomb of the colugos is generally considered to function as a toothcomb, but due to the lack of striations on the teeth and no documented observations of toothcomb use during oral grooming, its use seems to be limited to food procurement. 182:. While licking the fur clean, the animal will run the toothcomb through the fur to comb it. Fine grooves or striations are usually cut into the teeth during grooming by the hair and may be seen on the sides of the teeth when viewed through a 396:
that are procumbent (tilt forward) in the front of the mouth. The procumbent lower canine teeth are the same shape as the incisors located between them, but they are more robust and curve upward and inward more-so than the incisors. In the
353:), have a similar structure sometimes referred to as the "lateral dental grooming apparatus". Living and extinct hyraxes (hyracoids) also exhibit a toothcomb, although the number of tines in the comb vary throughout the fossil record. 1005:
Based on dental microwear patterns that are characteristic of dental combing, as well as the similarities between a laterally flared anterior incisor and the appearance of the lateral canine tooth in the lemuriform toothcomb, American
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and vary 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
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may have been contemporaneous with related toothcombed primates around 50 to 48 mya, the sparse African fossil record suggests toothcomb differentiation occurred around 52 to 40 mya according to the French paleoanthropologist
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revealed fine groves or striations on the teeth in lemuriform toothcombs. These grooves were only found on the sides of the teeth on the concave surfaces between the sides, as well on the back ridge of the teeth. Between 10 and
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striations can be affected by grooming frequency, fur thickness, or interdental spacing. Dental combing (as a function of the teeth with or without a toothcomb) has also been documented in early primates from the late Eocene.
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the extinct arctocyonids, all six lower incisors were part of the toothcomb. In African antelopes, the toothcomb is strikingly similar to that of lemuriforms in that it consists of two pairs of incisors and a pair of canines.
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Jaeger, J. J.; Beard, K. C.; Chaimanee, Y.; Salem, M.; Benammi, M.; Hlal, O.; Coster, P.; Bilal, A. A.; Duringer, P.; Schuster, M.; Valentin, X.; Marandat, B.; Marivaux, L.; Métais, E.; Hammuda, O.; Brunet, M. (2010).
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exhibited similar behaviors. Also, strong selective pressure from feeding ecology placed on the anterior dentition was emphasized, based on the specialized upper anterior dentition seen in the recently extinct
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wide, these grooves indicate that hair moved repeatedly across the teeth. Inside these grooves were even finer groves, less than 1 µm, created by abrasion with the cuticular layer of the hair.
776:, have been found in Africa and date from 50 to 48 mya and were very distinct from European adapiforms. However, they lack a toothcomb. These stem lemuriforms suggest an early common ancestry with 701:
proposed that the toothcomb found in treeshrews (which he believed were primates) was an early version of the dental structure found in lemuriforms. Because he viewed the fossil lorisoids from the
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Cuozzo, F.P.; Yamashita, N. (2006). "Chapter 4: Impact of Ecology on the Teeth of Extant Lemurs: A Review of Dental Adaptations, Function, and Life History". In Gould, L.; Sauther, M.L (eds.).
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primates, most of whom do not exhibit the characteristic toothcomb, and "lemuriform" has been problematic because of its historical membership of living and fossil primates, which included the
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The minimum paleontological estimate for the divergence of lemurs and lorisoids nearly doubled when additional discoveries were made in northern Egypt during the 2000s of a stem galagid (
458:) the toothcomb is more compressed, with significantly reduced interdental spaces. All six teeth are longer, straighter, and form a more continuous apical ridge. In the recently extinct 439:
or "under-tongue", a specialized muscular structure that acts like a toothbrush to remove hair and other debris. The sublingua can extend below the end of the tongue and is tipped with
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as not having fully developed the modern lemuriform toothcomb, he implied that lemurs and lorisoids had evolved the trait independently. This view was later overturned, and the
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Rosenberger, A.L. (2010). "Chapter 14: Adaptive profile versus adaptive specialization: Fossils and gummivory in early primate evolution". In Burrows, A.M.; Nash, L.T (eds.).
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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
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Toothcombs can also be found in colugos and treeshrews, both close relatives of primates; however, the structures are different and these are considered to examples of
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teeth of the lemuriform toothcomb as canines in 1840. Canine teeth are normally used to pierce or grasp objects. With modified lower canine teeth, the first lower
988:, but excluded the lorisoids. The term "lemuriforms" is used here because it derives from one popular taxonomy that clumps the toothcombed primates into one 694:
The origins of the lemuriform toothcomb and the clade it characterizes have been the center of considerable debate for more than a century. In 1920, British
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may have been their closest relatives. The lack of a distinct toothcomb in the fossil record prior to 40 mya has created a conflict with
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730:(a trait with adaptive value for something other than what it was originally selected for), which could then be used for personal or 564:. Instead, it is used during grooming when the head sweeps upward in a distinctive motion. It is thought to comb the fur and remove 882:
is the oldest fossil primate to exhibit a distinct lemuriform toothcomb. This, as well as studies of other African adapiforms like
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Asher, R. J. (1998). "Morphological diversity of anatomical strepsirrhinism and the evolution of the lemuriform toothcomb".
741:. In some adapids, the crests of the lower incisors and canines align to form functional cropping unit, and the American 530: 183: 972:
between lemurs and lorisoids is widely accepted, their clade name is not. The term "strepsirrhine" has been used as a
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Lemuriforms are currently thought to have evolved in Africa, and the earliest known prosimian primates from Africa are
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is a secondary tongue below the primary tongue and is used to remove hair and debris from the toothcomb of lemuriforms.
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A popular hypothesis about the origins of the lemuriform clade is that they evolved from European adapiforms known as
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Tattersall, I. (2006). "Chapter 1: Origin of the Malagasy strepsirhine primates". In Gould, L.; Sauther, M.L (eds.).
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Rose, K.D.; Walker, A.; Jacobs, L.L. (1981). "Function of the mandibular tooth comb in living and extinct mammals".
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Cercamoniines were an early branch of strepsirrhine primates that occupied the northern hemisphere during the early
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from the early Eocene, which likely descended from a very early colonization of the Afro-Arabian land mass in the
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The hypothesis that the toothcomb evolved for food procurement was based on observations of recent lemuriform
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Rosenberger, A.L.; Strasser, E. (1985). "Toothcomb origins: Support for the grooming hypothesis".
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In lemuriform primates, the toothcomb has been used by scientists in the interpretation of the
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In African antelopes, the lateral dental grooming apparatus does not appear to be used during
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have more robust toothcombs to support these secondary functions. In some lemurs, such as the
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Eaglen, R.H. (1980). "Toothcomb homology and toothcomb function in extant strepsirhines".
828: 731: 648: 508: 398: 256: 202:, the toothcomb has been lost completely and replaced with another specialized dentition. 40: 838:
and others, which predict lemur–lorisoid divergence dating between 61 and 90.8 mya.
1036:), lacked a toothcomb but showed signs of dental microwear attributed to dental combing. 2791: 2555: 2538: 2373: 2343: 2326: 993: 989: 926: 856: 782: 511: 251: 186:. The toothcomb is kept clean by either the tongue or, in the case of lemuriforms, the 98: 2645:"New views on tree shrews: The role of Tupaiids in primate supraordinal relationships" 404: 2841: 2441: 1238: 1236: 1018: 914: 893: 777: 373: 295: 264: 234: 66: 2759: 225:
suggests that they evolved from an older lineage that migrated to Africa during the
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Szalay, F.S.; Seligsohn, D. (1977). "Why did the strepsirhine tooth comb evolve?".
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that tilt forward at the front of the lower jaw, followed by a canine-shaped first
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The Hunt for the Dawn Monkey: Unearthing the Origins of Monkeys, Apes, and Humans
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The primary function of the toothcomb, grooming, was first noted by the French
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The toothcomb, a special morphological arrangement of teeth in the anterior
226: 195: 187: 135: 119: 2800: 2750: 2564: 2463:, a pygmy species endemic to the montane mossy forests of central Sulawesi" 2352: 2314: 2277: 2256:
Godinot, M. (2006). "Lemuriform origins as viewed from the fossil record".
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Stevens, N. J.; O'Connor, P. M.; Roberts, E. M.; Gottfried, M. D. (2009).
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Schwartz, J.H. (2003). "Chapter 3: How close are the similarities between
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10.1002/(SICI)1096-8644(199803)105:3<355::AID-AJPA5>3.0.CO;2-Q
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and the extinct, non-toothcombed adapiforms into another, both within the
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The toothcomb of most lemuriforms includes six finely spaced teeth, four
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In lemuriform primates, the toothcomb may also play a secondary role in
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and other primates?". In Wright, P.C.; Simons, E.L.; Gursky, S (eds.).
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or earlier. One popular hypothesis is that they evolved from European
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Schmid, P. (1983). "Front Dentition of the Omomyiformes (Primates)".
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The lemuriform toothcomb, viewed from the underside of the lower jaw
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This page is about the dental structure. For the colloquialism, see
1740: 1712: 1710: 1708: 1671: 1669: 1242: 2727:"Fossil evidence for an ancient divergence of lorises and galagos" 1514: 1512: 1510: 1508: 1506: 1504: 1278: 1276: 1274: 1272: 1270: 1268: 1266: 922: 794:, is thought to be most closely related to lemuriforms. Together, 791: 644: 620: 494: 403: 311: 299: 274: 143: 127: 97: 2360:
McKenzie, A.A. (1990). "The ruminant dental grooming apparatus".
1049:. They originated in Asia and are considered a subfamily of the 1342: 1340: 1338: 780:
from outside of Europe. Based on large, procumbent lower teeth,
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differs significantly in shape from that of lemuriform primates.
1131: 1129: 1116: 1114: 1112: 1110: 1108: 1095: 1093: 1053:, extinct North American primates that resembled modern lemurs. 658:) use their toothcombs to tooth-scrape plant exudates, such as 1565: 1563: 1301: 1299: 1297: 1295: 1293: 1291: 25: 1584: 1582: 1580: 1578: 2758:
Seiffert, E.R.; Simons, E.L.; Ryan, T.M.; Attia, Y. (2005).
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primates (strepsirrhines and tarsiers) possess at least one
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Among lemurs, the toothcomb is variable in structure. Among
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and their kin. They are thought to have evolved from early
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Primates Comparative Anatomy and Taxonomy I—Strepsirhini
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to be more easily transferred to the vomeronasal organ.
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the claws needed to adequately comb the fur, although
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exhibits microscopic groves on its toothcomb, but the
2388:; Richardson, M.; Schwitzer, C.; et al. (2010). 443:, serrated points that rake between the front teeth. 380:)—also possessed an independently evolved toothcomb. 263:
between lemurs and lorisoids, and the existence of a
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Evolutionary Anthropology: Issues, News, and Reviews
2069: 2030: 1752: 1687: 1227: 1195: 676:may also use their toothcomb for food procurement. 306:primates (collectively known as lemuriforms). This 2193:"Chapter 2: Primate Classification and Diversity" 310:is a diagnostic character that helps define this 2725:Seiffert, E.R.; Simons, E.L.; Attia, Y. (2003). 2421:Mooring, M.; McKenzie, A.A.; Hart, B.L. (1996). 1946: 1843: 1518: 1282: 1072: 810:(the closest relatives) of lemuriform primates. 2821:Proceedings of the National Academy of Sciences 2771:Proceedings of the National Academy of Sciences 2093: 2081: 2054: 2042: 1934: 1807: 1728: 1716: 1675: 1660: 1483: 1346: 1135: 1120: 1099: 2814:(Primates, Prosimii) from the Eocene of Egypt" 709:between lemurs and lorisoids is now accepted. 435:The lemuriform toothcomb is kept clean by the 2507:"Evolutionary history of lorisiform primates" 1636: 900:Original function of the lemuriform toothcomb 878:) dating to 37 and 40 mya respectively. 8: 2710:. Rutgers University Press. pp. 50–96. 2505:Rasmussen, D. T.; Nekaris, K. A. I. (1998). 1569: 1495: 1471: 1459: 1447: 1370: 1305: 341:. Likewise, small- or medium-sized African 48:. A user sandbox is a subpage of the user's 2394:. Illustrated by S.D. Nash (3rd ed.). 2199:. Oxford University Press. pp. 10–30. 1588: 592:in the roof of the mouth. This would allow 584:(a cleft in the middle of the wet nose, or 2018: 1970: 1819: 1435: 1382: 1329: 2840: 2790: 2554: 2362:Zoological Journal of the Linnean Society 2342: 2143:American Journal of Physical Anthropology 2604:The Evolution of Exudativory in Primates 1624: 1540:sfn error: no target: CITEREFJones1918 ( 1421:sfn error: no target: CITEREFJones1918 ( 1399:sfn error: no target: CITEREFJones1918 ( 1212: 1164:sfn error: no target: CITEREFJones1918 ( 1084: 2327:"The sublingua and the plica fimbriata" 1982: 1958: 1922: 1886: 1855: 1831: 1795: 1783: 1764: 1612: 1600: 1065: 961: 2808:Simons, E.L.; Rasmussen, D.T. (1994). 2105: 1648: 1554: 1358: 1317: 1257: 1183: 1147: 412:, with canine-like premolars behind it 154:in different types of mammals through 2195:. In Platt, M.; Ghazanfar, A (eds.). 1910: 1898: 1699: 1535: 1416: 1394: 1159: 7: 2943: 2237:International Journal of Primatology 717:. Adapiforms are considered to be a 233:) and might have evolved from early 2707:Tarsiers: Past, Present, and Future 356:Dating to the Eocene epoch over 50 271:Homologous and analogous structures 237:from Asia. Fossil primates such as 2871:Journal of Vertebrate Paleontology 2457:Musser, G.G.; Dagosto, M. (1987). 2374:10.1111/j.1096-3642.1990.tb00564.x 2176:. University of California Press. 514:in 1829, who pointed out that the 421:Henri Marie Ducrotay de Blainville 267:of lemuriform primates in Africa. 178:The toothcomb is usually used for 24: 2031:Seiffert, Simons & Attia 2003 1228:Mooring, McKenzie & Hart 1996 1196:Mooring, McKenzie & Hart 1996 1032:(close relatives of tarsier-like 2126:(3rd ed.). Academic Press. 872:) and a stem or crown lorisoid ( 541:Among non-primates, the extinct 58:Create or edit your own sandbox 29: 2070:Rosenberger & Strasser 1985 1753:Rosenberger & Strasser 1985 1688:Rosenberger & Strasser 1985 827:. This would conflict with the 818:Although stem lemuriforms like 600:Food procurement and other uses 2924:Lemurs: Ecology and Adaptation 2607:. Springer. pp. 273–295. 2217:Lemurs: Ecology and Adaptation 1519:Rose, Walker & Jacobs 1981 1283:Rose, Walker & Jacobs 1981 859:lemur, further challenged the 855:and initially thought to be a 690:Evolutionary history of lemurs 259:studies that suggest an older 1: 81:Submit your draft for review! 2442:10.1016/0031-9384(95)02186-8 2220:. Springer. pp. 67–96. 1947:Rasmussen & Nekaris 1998 1844:Rasmussen & Nekaris 1998 1073:Rasmussen & Nekaris 1998 531:scanning electron microscopy 184:scanning electron microscope 2949:Knowledge:Featured articles 2927:. Springer. pp. 3–20. 2539:"The dental comb of lemurs" 2094:Cuozzo & Yamashita 2006 2082:Szalay & Seligsohn 1977 2055:Szalay & Seligsohn 1977 2043:Cuozzo & Yamashita 2006 1935:Simons & Rasmussen 1994 1808:Szalay & Seligsohn 1977 1729:Szalay & Seligsohn 1977 1717:Szalay & Seligsohn 1977 1676:Cuozzo & Yamashita 2006 1661:Cuozzo & Yamashita 2006 1484:Szalay & Seligsohn 1977 1347:Szalay & Seligsohn 1977 1136:Szalay & Seligsohn 1977 1121:Cuozzo & Yamashita 2006 1100:Cuozzo & Yamashita 2006 896:for lemuriforms in Africa. 833:evolutionary anthropologist 522:) had lower incisors that " 54:not an encyclopedia article 2966: 2396:Conservation International 892:, suggests a more ancient 841:In 2001, the discovery of 687: 524:sont de véritables peignes 90: 2945:Template:Featured article 2810:"A remarkable cranium of 2516:. 69 (suppl 1): 250–285. 2470:American Museum Novitates 2430:Physiology & Behavior 2122:Ankel-Simons, F. (2007). 1637:Musser & Dagosto 1987 1150:, pp. 278 & 280. 970:monophyletic relationship 707:monophyletic relationship 423:first identified the two 847:, a fossil primate from 684:Evolution in lemuriforms 572:Olfaction in lemuriforms 2833:10.1073/pnas.91.21.9946 2783:10.1073/pnas.0505310102 2760:"Additional remains of 1741:Mittermeier et al. 2010 1008:physical anthropologist 861:theory of lemur origins 764:lemuriforms, including 633:hairy-eared dwarf lemur 462:(Archaeolemuridae) and 814:Dating inconsistencies 504: 413: 296:strepsirrhine primates 284: 103: 2862:Rukwalorax jinokitana 2643:Sargis, E.J. (2004). 2197:Primate Neuroethology 2191:Cartmill, M. (2010). 1028:, a type of European 806:are considered to be 760:(65 to 55 mya). 699:Wilfrid Le Gros Clark 670:colugos in the genus 498: 407: 278: 152:evolved independently 101: 2883:10.1671/039.029.0302 2864:(gen. and sp. nov.)" 2812:Plesiopithecus teras 2537:Roberts, D. (1941). 2391:Lemurs of Madagascar 2007:Seiffert et al. 2005 1875:Seiffert et al. 2005 984:lemurs and European 884:‘Anchomomys’ milleri 800:‘Anchomomys’ milleri 773:'Anchomomys' milleri 696:palaeoanthropologist 384:Anatomical structure 339:convergent evolution 308:homologous structure 246:'Anchomomys' milleri 213:primates around the 156:convergent evolution 2893:Folia Primatologica 2777:(32): 11396–11401. 2743:10.1038/nature01489 2673:Folia Primatologica 2514:Folia Primatologica 2307:10.1038/nature09425 2301:(7319): 1095–1098. 2258:Folia Primatologica 2033:, pp. 423–424. 1985:, pp. 447–449. 1925:, pp. 453–455. 1858:, pp. 455–456. 1846:, pp. 256–258. 1798:, pp. 446–447. 1615:, pp. 236–237. 1450:, pp. 421–423. 1243:Stevens et al. 2009 1186:, pp. 279–280. 1162:, pp. 345–346. 1011:Jeffrey H. Schwartz 746:Philip D. Gingerich 551:Cynocephalus volans 399:permanent dentition 314:(related group) of 291:, is best known in 279:The toothcomb of a 207:evolution of lemurs 2952: 2664:10.1002/evan.10131 2636:10.1007/BF02389048 2543:Journal of Anatomy 2331:Journal of Anatomy 2249:10.1007/BF02692275 2170:Beard, C. (2004). 1995:Jaeger et al. 2010 906:selective pressure 643:(particularly the 505: 452:fork-marked lemurs 414: 351:Aepyceros melampus 285: 192:fork-marked lemurs 150:. The structures 104: 2934:978-0-387-34585-7 2906:10.1159/000155778 2827:(21): 9946–9950. 2762:Wadilemur elegans 2737:(6930): 421–424. 2717:978-0-8135-3236-3 2685:10.1159/000156088 2614:978-1-4419-6660-5 2580:(5798): 583–585. 2549:(Pt 2): 236–238. 2522:10.1159/000052716 2459:"The identity of 2405:978-1-934151-23-5 2382:Mittermeier, R.A. 2337:(Pt 4): 345–353. 2270:10.1159/000095391 2227:978-0-387-34585-7 2206:978-0-19-532659-8 2183:978-0-520-23369-0 2133:978-0-12-372576-9 2084:, pp. 80–81. 2045:, pp. 73–74. 1913:, pp. 51–52. 1901:, pp. 55–56. 1719:, pp. 76–77. 1678:, pp. 74–76. 1639:, pp. 33–34. 1627:, pp. 66–67. 1570:Ankel-Simons 2007 1496:Ankel-Simons 2007 1486:, pp. 78–79. 1472:Ankel-Simons 2007 1460:Ankel-Simons 2007 1448:Ankel-Simons 2007 1371:Ankel-Simons 2007 1332:, pp. 52–53. 1306:Ankel-Simons 2007 974:wastebasket taxon 590:vomeronasal organ 547:Philippine colugo 516:ring-tailed lemur 410:ring-tailed lemur 378:placental mammals 146:and some African 89: 88: 65:Other sandboxes: 63: 2957: 2938: 2917: 2886: 2868: 2854: 2844: 2818: 2804: 2794: 2768: 2754: 2721: 2696: 2667: 2649: 2639: 2618: 2597: 2586:10.1038/289583a0 2568: 2558: 2533: 2511: 2501: 2486:Osman Hill, W.C. 2481: 2467: 2453: 2436:(4–5): 965–971. 2427: 2417: 2377: 2356: 2346: 2318: 2292: 2281: 2252: 2231: 2210: 2187: 2166: 2137: 2115:Literature cited 2109: 2103: 2097: 2091: 2085: 2079: 2073: 2067: 2058: 2052: 2046: 2040: 2034: 2028: 2022: 2016: 2010: 2009:, p. 11396. 2004: 1998: 1992: 1986: 1980: 1974: 1968: 1962: 1956: 1950: 1944: 1938: 1932: 1926: 1920: 1914: 1908: 1902: 1896: 1890: 1884: 1878: 1877:, p. 11400. 1872: 1859: 1853: 1847: 1841: 1835: 1829: 1823: 1817: 1811: 1805: 1799: 1793: 1787: 1781: 1768: 1762: 1756: 1750: 1744: 1738: 1732: 1726: 1720: 1714: 1703: 1697: 1691: 1685: 1679: 1673: 1664: 1658: 1652: 1646: 1640: 1634: 1628: 1622: 1616: 1610: 1604: 1598: 1592: 1589:Rosenberger 2010 1586: 1573: 1567: 1558: 1552: 1546: 1545: 1533: 1522: 1516: 1499: 1493: 1487: 1481: 1475: 1469: 1463: 1457: 1451: 1445: 1439: 1433: 1427: 1426: 1414: 1405: 1404: 1392: 1386: 1380: 1374: 1368: 1362: 1356: 1350: 1344: 1333: 1327: 1321: 1315: 1309: 1303: 1286: 1280: 1261: 1255: 1246: 1240: 1231: 1225: 1216: 1210: 1199: 1193: 1187: 1181: 1170: 1169: 1157: 1151: 1145: 1139: 1133: 1124: 1118: 1103: 1097: 1088: 1082: 1076: 1070: 1054: 1043: 1037: 1003: 997: 966: 788:Fayum Depression 332:New World monkey 322:is found in the 298:, which include 126:(which includes 85: 84: 82: 71:Template sandbox 57: 33: 32: 26: 18:User:EngetsuGray 2965: 2964: 2960: 2959: 2958: 2956: 2955: 2954: 2941: 2935: 2920: 2889: 2866: 2857: 2816: 2807: 2766: 2757: 2724: 2718: 2699: 2670: 2647: 2642: 2621: 2615: 2600: 2571: 2536: 2509: 2504: 2484: 2465: 2461:Tarsius pumilus 2456: 2425: 2420: 2406: 2380: 2359: 2321: 2290: 2284: 2255: 2234: 2228: 2213: 2207: 2190: 2184: 2169: 2140: 2134: 2124:Primate Anatomy 2121: 2117: 2112: 2104: 2100: 2092: 2088: 2080: 2076: 2068: 2061: 2053: 2049: 2041: 2037: 2029: 2025: 2019:Tattersall 2006 2017: 2013: 2005: 2001: 1997:, p. 1095. 1993: 1989: 1981: 1977: 1973:, pp. 8–9. 1971:Tattersall 2006 1969: 1965: 1957: 1953: 1945: 1941: 1937:, p. 9946. 1933: 1929: 1921: 1917: 1909: 1905: 1897: 1893: 1885: 1881: 1873: 1862: 1854: 1850: 1842: 1838: 1830: 1826: 1820:Tattersall 2006 1818: 1814: 1806: 1802: 1794: 1790: 1782: 1771: 1763: 1759: 1751: 1747: 1739: 1735: 1727: 1723: 1715: 1706: 1698: 1694: 1686: 1682: 1674: 1667: 1659: 1655: 1651:, pp. 6–8. 1647: 1643: 1635: 1631: 1623: 1619: 1611: 1607: 1599: 1595: 1587: 1576: 1568: 1561: 1553: 1549: 1539: 1534: 1525: 1517: 1502: 1494: 1490: 1482: 1478: 1470: 1466: 1458: 1454: 1446: 1442: 1436:Osman Hill 1953 1434: 1430: 1420: 1415: 1408: 1398: 1393: 1389: 1383:Osman Hill 1953 1381: 1377: 1369: 1365: 1357: 1353: 1345: 1336: 1330:Osman Hill 1953 1328: 1324: 1316: 1312: 1304: 1289: 1281: 1264: 1256: 1249: 1241: 1234: 1226: 1219: 1211: 1202: 1194: 1190: 1182: 1173: 1163: 1158: 1154: 1146: 1142: 1134: 1127: 1119: 1106: 1098: 1091: 1083: 1079: 1071: 1067: 1063: 1058: 1057: 1044: 1040: 1013:suggested that 1004: 1000: 967: 963: 958: 902: 829:molecular clock 816: 732:social grooming 692: 686: 602: 574: 493: 484: 408:Toothcomb of a 386: 320:analogous trait 273: 257:molecular clock 229:(65 to 55  96: 80: 78: 77: 75: 74: 30: 22: 21: 20: 12: 11: 5: 2963: 2961: 2940: 2939: 2933: 2918: 2887: 2855: 2805: 2755: 2722: 2716: 2697: 2668: 2640: 2619: 2613: 2598: 2569: 2534: 2502: 2482: 2472:(2867): 1–53. 2454: 2418: 2404: 2378: 2368:(2): 117–128. 2357: 2323:Wood Jones, F. 2319: 2282: 2264:(6): 446–464. 2253: 2243:(3): 275–286. 2232: 2226: 2211: 2205: 2188: 2182: 2167: 2149:(3): 355–367. 2138: 2132: 2118: 2116: 2113: 2111: 2110: 2108:, p. 276. 2098: 2086: 2074: 2059: 2047: 2035: 2023: 2011: 1999: 1987: 1975: 1963: 1961:, p. 458. 1951: 1949:, p. 261. 1939: 1927: 1915: 1903: 1891: 1889:, p. 452. 1879: 1860: 1848: 1836: 1834:, p. 447. 1824: 1812: 1800: 1788: 1786:, p. 446. 1769: 1767:, p. 455. 1757: 1745: 1743:, p. 572. 1733: 1721: 1704: 1702:, p. 355. 1692: 1680: 1665: 1653: 1641: 1629: 1617: 1605: 1603:, p. 236. 1593: 1591:, p. 291. 1574: 1572:, p. 238. 1559: 1547: 1538:, p. 346. 1523: 1521:, p. 585. 1500: 1498:, p. 257. 1488: 1476: 1474:, p. 233. 1464: 1462:, p. 254. 1452: 1440: 1428: 1419:, p. 347. 1406: 1397:, p. 345. 1387: 1375: 1373:, p. 234. 1363: 1361:, p. 279. 1351: 1334: 1322: 1320:, p. 281. 1310: 1308:, p. 235. 1287: 1285:, p. 583. 1262: 1260:, p. 277. 1247: 1245:, p. 974. 1232: 1230:, p. 966. 1217: 1215:, p. 117. 1200: 1198:, p. 970. 1188: 1171: 1152: 1140: 1125: 1104: 1089: 1077: 1075:, p. 252. 1064: 1062: 1059: 1056: 1055: 1038: 1017:have a highly 998: 996:Strepsirrhini. 960: 959: 957: 954: 901: 898: 851:dating to the 815: 812: 804:Plesiopithecus 783:Plesiopithecus 743:paleontologist 685: 682: 601: 598: 573: 570: 512:Georges Cuvier 492: 489: 483: 480: 385: 382: 372:—two types of 345:, such as the 328:Cacajao calvus 272: 269: 252:Plesiopithecus 87: 86: 55: 36: 34: 23: 15: 14: 13: 10: 9: 6: 4: 3: 2: 2962: 2953: 2950: 2946: 2936: 2930: 2926: 2925: 2919: 2915: 2911: 2907: 2903: 2899: 2895: 2894: 2888: 2884: 2880: 2876: 2872: 2865: 2863: 2856: 2852: 2848: 2843: 2838: 2834: 2830: 2826: 2822: 2815: 2813: 2806: 2802: 2798: 2793: 2788: 2784: 2780: 2776: 2772: 2765: 2763: 2756: 2752: 2748: 2744: 2740: 2736: 2732: 2728: 2723: 2719: 2713: 2709: 2708: 2703: 2698: 2694: 2690: 2686: 2682: 2679:(1–2): 1–10. 2678: 2674: 2669: 2665: 2661: 2657: 2653: 2646: 2641: 2637: 2633: 2629: 2625: 2620: 2616: 2610: 2606: 2605: 2599: 2595: 2591: 2587: 2583: 2579: 2575: 2570: 2566: 2562: 2557: 2552: 2548: 2544: 2540: 2535: 2531: 2527: 2523: 2519: 2515: 2508: 2503: 2499: 2495: 2491: 2487: 2483: 2479: 2475: 2471: 2464: 2462: 2455: 2451: 2447: 2443: 2439: 2435: 2431: 2424: 2419: 2415: 2411: 2407: 2401: 2397: 2393: 2392: 2387: 2383: 2379: 2375: 2371: 2367: 2363: 2358: 2354: 2350: 2345: 2340: 2336: 2332: 2328: 2324: 2320: 2316: 2312: 2308: 2304: 2300: 2296: 2289: 2283: 2279: 2275: 2271: 2267: 2263: 2259: 2254: 2250: 2246: 2242: 2238: 2233: 2229: 2223: 2219: 2218: 2212: 2208: 2202: 2198: 2194: 2189: 2185: 2179: 2175: 2174: 2168: 2164: 2160: 2156: 2152: 2148: 2144: 2139: 2135: 2129: 2125: 2120: 2119: 2114: 2107: 2102: 2099: 2096:, p. 74. 2095: 2090: 2087: 2083: 2078: 2075: 2072:, p. 75. 2071: 2066: 2064: 2060: 2057:, p. 75. 2056: 2051: 2048: 2044: 2039: 2036: 2032: 2027: 2024: 2020: 2015: 2012: 2008: 2003: 2000: 1996: 1991: 1988: 1984: 1979: 1976: 1972: 1967: 1964: 1960: 1955: 1952: 1948: 1943: 1940: 1936: 1931: 1928: 1924: 1919: 1916: 1912: 1907: 1904: 1900: 1895: 1892: 1888: 1883: 1880: 1876: 1871: 1869: 1867: 1865: 1861: 1857: 1852: 1849: 1845: 1840: 1837: 1833: 1828: 1825: 1821: 1816: 1813: 1810:, p. 77. 1809: 1804: 1801: 1797: 1792: 1789: 1785: 1780: 1778: 1776: 1774: 1770: 1766: 1761: 1758: 1755:, p. 74. 1754: 1749: 1746: 1742: 1737: 1734: 1731:, p. 80. 1730: 1725: 1722: 1718: 1713: 1711: 1709: 1705: 1701: 1696: 1693: 1690:, p. 73. 1689: 1684: 1681: 1677: 1672: 1670: 1666: 1663:, p. 72. 1662: 1657: 1654: 1650: 1645: 1642: 1638: 1633: 1630: 1626: 1625:Schwartz 2003 1621: 1618: 1614: 1609: 1606: 1602: 1597: 1594: 1590: 1585: 1583: 1581: 1579: 1575: 1571: 1566: 1564: 1560: 1557:, p. 58. 1556: 1551: 1548: 1543: 1537: 1532: 1530: 1528: 1524: 1520: 1515: 1513: 1511: 1509: 1507: 1505: 1501: 1497: 1492: 1489: 1485: 1480: 1477: 1473: 1468: 1465: 1461: 1456: 1453: 1449: 1444: 1441: 1438:, p. 73. 1437: 1432: 1429: 1424: 1418: 1413: 1411: 1407: 1402: 1396: 1391: 1388: 1385:, p. 53. 1384: 1379: 1376: 1372: 1367: 1364: 1360: 1355: 1352: 1349:, p. 78. 1348: 1343: 1341: 1339: 1335: 1331: 1326: 1323: 1319: 1314: 1311: 1307: 1302: 1300: 1298: 1296: 1294: 1292: 1288: 1284: 1279: 1277: 1275: 1273: 1271: 1269: 1267: 1263: 1259: 1254: 1252: 1248: 1244: 1239: 1237: 1233: 1229: 1224: 1222: 1218: 1214: 1213:McKenzie 1990 1209: 1207: 1205: 1201: 1197: 1192: 1189: 1185: 1180: 1178: 1176: 1172: 1167: 1161: 1156: 1153: 1149: 1144: 1141: 1138:, p. 76. 1137: 1132: 1130: 1126: 1123:, p. 73. 1122: 1117: 1115: 1113: 1111: 1109: 1105: 1102:, p. 68. 1101: 1096: 1094: 1090: 1087:, p. 15. 1086: 1085:Cartmill 2010 1081: 1078: 1074: 1069: 1066: 1060: 1052: 1048: 1042: 1039: 1035: 1031: 1027: 1026: 1020: 1016: 1012: 1009: 1002: 999: 995: 991: 987: 983: 979: 975: 971: 968:Although the 965: 962: 955: 953: 949: 947: 943: 942: 937: 933: 928: 924: 919: 916: 915:grooming claw 912: 907: 899: 897: 895: 894:ghost lineage 891: 890: 885: 881: 877: 876: 871: 870: 864: 862: 858: 854: 850: 846: 845: 839: 837: 834: 831:estimates by 830: 826: 821: 813: 811: 809: 805: 801: 797: 793: 789: 785: 784: 779: 778:cercamoniines 775: 774: 769: 768: 763: 759: 755: 750: 747: 744: 740: 735: 733: 729: 725: 720: 716: 710: 708: 704: 700: 697: 691: 683: 681: 677: 675: 674: 669: 665: 661: 657: 656: 651: 650: 646: 642: 638: 634: 630: 626: 622: 618: 614: 610: 606: 599: 597: 595: 591: 587: 583: 579: 571: 569: 567: 566:ectoparasites 563: 559: 554: 552: 548: 544: 539: 537: 532: 527: 525: 521: 517: 513: 510: 502: 497: 490: 488: 481: 479: 475: 471: 469: 465: 461: 460:monkey lemurs 457: 453: 449: 444: 442: 438: 433: 430: 426: 422: 419: 411: 406: 402: 400: 395: 391: 383: 381: 379: 375: 371: 370: 365: 364: 359: 354: 352: 348: 344: 340: 335: 333: 330:), a type of 329: 325: 321: 317: 313: 309: 305: 301: 297: 294: 290: 282: 277: 270: 268: 266: 265:ghost lineage 262: 258: 254: 253: 248: 247: 242: 241: 236: 235:cercamoniines 232: 228: 224: 223:fossil record 220: 216: 212: 208: 203: 201: 197: 193: 189: 185: 181: 176: 174: 170: 166: 162: 157: 153: 149: 145: 141: 137: 133: 129: 125: 121: 117: 113: 109: 100: 94: 83: 73: 72: 68: 61: 53: 51: 47: 46: 42: 35: 28: 27: 19: 2947:is only for 2942: 2923: 2900:(1): 75–82. 2897: 2891: 2874: 2870: 2861: 2824: 2820: 2811: 2774: 2770: 2761: 2734: 2730: 2706: 2701: 2676: 2672: 2658:(2): 56–66. 2655: 2651: 2627: 2623: 2603: 2577: 2573: 2546: 2542: 2513: 2489: 2469: 2460: 2433: 2429: 2389: 2365: 2361: 2334: 2330: 2298: 2294: 2261: 2257: 2240: 2236: 2216: 2196: 2172: 2146: 2142: 2123: 2101: 2089: 2077: 2050: 2038: 2026: 2021:, p. 7. 2014: 2002: 1990: 1983:Godinot 2006 1978: 1966: 1959:Godinot 2006 1954: 1942: 1930: 1923:Godinot 2006 1918: 1906: 1894: 1887:Godinot 2006 1882: 1856:Godinot 2006 1851: 1839: 1832:Godinot 2006 1827: 1822:, p. 9. 1815: 1803: 1796:Godinot 2006 1791: 1784:Godinot 2006 1765:Godinot 2006 1760: 1748: 1736: 1724: 1695: 1683: 1656: 1644: 1632: 1620: 1613:Roberts 1941 1608: 1601:Roberts 1941 1596: 1550: 1491: 1479: 1467: 1455: 1443: 1431: 1390: 1378: 1366: 1354: 1325: 1313: 1191: 1155: 1143: 1080: 1068: 1041: 1030:microchoerid 1023: 1001: 964: 950: 939: 935: 932:koala lemurs 927:cheirogaleid 920: 903: 887: 883: 879: 873: 869:Saharagalago 867: 865: 857:cheirogaleid 842: 840: 825:Marc Godinot 819: 817: 803: 799: 795: 781: 771: 765: 751: 736: 719:paraphyletic 711: 693: 678: 673:Cynocephalus 671: 653: 647: 636: 628: 624: 616: 608: 605:Mouse lemurs 603: 575: 555: 550: 542: 540: 528: 523: 519: 506: 485: 476: 472: 464:sloth lemurs 455: 445: 434: 415: 394:canine teeth 387: 374:arctocyonids 369:Thryptacodon 367: 361: 355: 350: 336: 327: 286: 250: 244: 238: 204: 177: 175:per tooth. 165:canine teeth 115: 111: 107: 105: 67:Main sandbox 64: 38: 2386:Louis, E.E. 2106:Eaglen 1980 1649:Schmid 1983 1555:Sargis 2004 1359:Eaglen 1980 1318:Eaglen 1980 1258:Eaglen 1980 1184:Eaglen 1980 1148:Eaglen 1980 1051:notharctids 944:(a type of 936:Megaladapis 808:sister taxa 668:herbivorous 619:), and the 617:Propithecus 520:Lemur catta 441:keratinized 432:behind it. 416:The French 376:(primitive 324:bald uakari 116:dental comb 45:EngetsuGray 2877:(3): 972. 1911:Beard 2004 1899:Beard 2004 1700:Asher 1998 1536:Jones 1918 1417:Jones 1918 1395:Jones 1918 1160:Jones 1918 1061:References 1025:Necrolemur 990:infraorder 978:haplorhine 941:Callithrix 925:, such as 844:Bugtilemur 836:Anne Yoder 820:Djebelemur 796:Djebelemur 767:Djebelemur 728:exaptation 724:folivorous 715:adapiforms 688:See also: 609:Microcebus 594:pheromones 509:naturalist 261:divergence 240:Djebelemur 221:, but the 136:treeshrews 112:tooth comb 2630:: 73–84. 2498:500576914 2478:0003-0082 2414:670545286 1047:Paleogene 911:prosimian 889:Wadilemur 880:Karanisia 875:Karanisia 853:Oligocene 758:Paleocene 637:Allocebus 588:) to the 586:rhinarium 578:olfaction 529:In 1981, 501:sublingua 482:Functions 437:sublingua 429:premolars 418:anatomist 343:antelopes 289:lower jaw 227:Paleocene 211:adapiform 188:sublingua 148:antelopes 132:lorisoids 108:toothcomb 50:user page 39:the user 2801:16087891 2751:12660781 2624:Primates 2565:17104854 2488:(1953). 2353:17103846 2325:(1918). 2315:20981098 2278:17053330 1034:omomyids 1015:tarsiers 994:suborder 982:Malagasy 976:for non- 946:marmoset 849:Pakistan 754:azibiids 655:Euoticus 582:philtrum 562:browsing 543:Chriacus 534:20  491:Grooming 448:indriids 392:and two 390:incisors 363:Chriacus 316:primates 304:lorisoid 196:indriids 180:grooming 169:premolar 161:incisors 124:primates 93:Nit comb 37:This is 2851:7937923 2792:1183603 2702:Tarsius 2693:6407925 2594:7007889 2556:1252661 2530:9595692 2450:8778894 2344:1262842 2163:9545077 1019:derived 986:adapids 739:adapids 703:Miocene 641:galagos 639:), and 631:), the 613:sifakas 558:grazing 468:rodents 425:lateral 219:adapids 200:aye-aye 144:hyraxes 140:colugos 118:) is a 69:| 41:sandbox 2931:  2914:401757 2912:  2849:  2839:  2799:  2789:  2749:  2731:Nature 2714:  2691:  2611:  2592:  2574:Nature 2563:  2553:  2528:  2496:  2476:  2448:  2412:  2402:  2351:  2341:  2313:  2295:Nature 2276:  2224:  2203:  2180:  2161:  2130:  802:, and 649:Galago 645:genera 629:Phaner 487:bark. 456:Phaner 347:impala 300:lemurs 293:extant 281:colugo 249:, and 215:Eocene 128:lemurs 120:dental 110:(also 2867:(PDF) 2842:44934 2817:(PDF) 2767:(PDF) 2648:(PDF) 2510:(PDF) 2466:(PDF) 2426:(PDF) 2291:(PDF) 956:Notes 792:Egypt 625:Indri 621:indri 318:. 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Index

User:EngetsuGray
sandbox
EngetsuGray
user page
here
Main sandbox
Template sandbox
Submit your draft for review!
Nit comb

dental
primates
lemurs
lorisoids
treeshrews
colugos
hyraxes
antelopes
evolved independently
convergent evolution
incisors
canine teeth
premolar
tines
grooming
scanning electron microscope
sublingua
fork-marked lemurs
indriids
aye-aye

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