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Strepsirrhini 1
Strepsirrhini
Strepsirrhines
Temporal range: Early EoceneRecent
Ring-tailed lemur (Lemur catta)
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Suborder: Strepsirrhini. Geoffroy, 1812
Infraorders
AdapiformesLemuriformes(See text)
Range of living strepsirrhine primates (green) and Eocene-Miocene fossil sites (red)
Strepsirrhini or Strepsirhini ( i/strpsrani/; STREP-s-RY-nee) 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. Also belonging to the suborder are the extinct adapiform primates, a
diverse and widespread group that thrived during the Eocene (56 to 34 million years ago [mya]) in Europe, North
America, and Asia, but disappeared from most of the Northern Hemisphere as the climate cooled. The last of the
adapiforms died out at the end of the Miocene (~7 mya). Adapiforms are sometimes referred to as being
"lemur-like", although the diversity of both lemurs and adapiforms do not support this comparison. The two leading
taxonomic classifications for the suborder divide living strepsirrhine primates into either two superfamilies(Lemuroidea and Lorisoidea) within the infraorder Lemuriformes or two infraorders, Lemuriformes and
Lorisiformes. The suborder represents a related group, and replaced the widely used and now obsolete suborder
http://en.wikipedia.org/w/index.php?title=Miocenehttp://en.wikipedia.org/w/index.php?title=Miocenehttp://en.wikipedia.org/w/index.php?title=Miocenehttp://en.wikipedia.org/w/index.php?title=Miocenehttp://en.wikipedia.org/w/index.php?title=Miocenehttp://en.wikipedia.org/w/index.php?title=Taxonomy_%28biology%29http://en.wikipedia.org/w/index.php?title=Taxonomic_rankhttp://en.wikipedia.org/w/index.php?title=Taxonomic_rankhttp://en.wikipedia.org/w/index.php?title=Taxonomic_rankhttp://en.wikipedia.org/w/index.php?title=Taxonomy_%28biology%29http://en.wikipedia.org/w/index.php?title=Miocenehttp://en.wikipedia.org/w/index.php?title=Northern_Hemispherehttp://en.wikipedia.org/w/index.php?title=Mya_%28unit%29http://en.wikipedia.org/w/index.php?title=Eocenehttp://en.wikipedia.org/w/index.php?title=Adapiformeshttp://en.wikipedia.org/w/index.php?title=Lorishttp://en.wikipedia.org/w/index.php?title=Pottohttp://en.wikipedia.org/w/index.php?title=Galagohttp://en.wikipedia.org/w/index.php?title=Madagascarhttp://en.wikipedia.org/w/index.php?title=Lemurhttp://en.wikipedia.org/w/index.php?title=Lemuriformeshttp://en.wikipedia.org/w/index.php?title=Primatehttp://en.wikipedia.org/w/index.php?title=Taxonomic_rankhttp://en.wikipedia.org/wiki/Pronunciation_respelling_keyhttp://en.wikipedia.org/w/index.php?title=Help:IPA_for_Englishhttp://en.wikipedia.org/w/index.php?title=Help:IPA_for_English%23Keyhttp://en.wikipedia.org/w/index.php?title=Help:IPA_for_Englishhttp://en.wikipedia.org/w/index.php?title=File:En-us-Strepsirrhini.ogghttp://en.wikipedia.org/w/index.php?title=File:Speakerlink-new.svghttp://en.wikipedia.org/w/index.php?title=File%3AExtant_strepsirrhine_range_with_fossil_sites.pnghttp://en.wikipedia.org/w/index.php?title=Lemuriformeshttp://en.wikipedia.org/w/index.php?title=Adapiformeshttp://en.wikipedia.org/w/index.php?title=Order_%28biology%29http://en.wikipedia.org/w/index.php?title=%C3%89tienne_Geoffroy_Saint-Hilairehttp://en.wikipedia.org/w/index.php?title=Primatehttp://en.wikipedia.org/w/index.php?title=Mammalhttp://en.wikipedia.org/w/index.php?title=Chordatehttp://en.wikipedia.org/w/index.php?title=Animalhttp://en.wikipedia.org/w/index.php?title=Biological_classificationhttp://en.wikipedia.org/w/index.php?title=Ring-tailed_lemurhttp://en.wikipedia.org/w/index.php?title=File%3ARingtailedlemur_-_24937.jpghttp://en.wikipedia.org/w/index.php?title=Eocene -
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Strepsirrhini 2
Prosimii ("prosimians"), which included strepsirrhines and tarsiers, a grouping based primarily on shared anatomical
traits. Today, Strepsirrhini excludes the tarsiers, which are now grouped in the other major primate suborder,
Haplorhini, along with the monkeys and apes (simians or anthropoids). Strepsirrhines are often inappropriately
referred to as "living fossils". Instead, they have evolved for millions of years under natural selection, and have
diversified to fill many ecological niches. Some of their traits may be derived from ancestral primates, while others
are unique to strepsirrhines.
Strepsirrhines are defined by their wet nose or rhinarium. They also have a smaller brain than comparably sized
simians, large olfactory lobes for smell, a vomeronasal organ to detect pheromones, and a bicornuate uterus with an
epitheliochorial placenta. Their eyes contain a reflective layer to improve their night vision, and their eye sockets
include a ring of bone around the eye, but they lack a wall of thin bone behind it. Strepsirrhine primates produce
their own vitamin C, whereas haplorhine primates must obtain it from their diets. 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. Often, the toothcomb is incorrectly used to characterize all strepsirrhines. Instead, it is unique
to lemuriforms and is not seen among adapiforms. Lemuriforms groom orally, and also possess a grooming claw on
the second toe of each foot for scratching in areas that are inaccessible to the mouth and tongue. It is unclear whether
adapiforms possessed grooming claws.The taxonomy of strepsirrhines is controversial and has a complicated history. Confused taxonomic terminology and
oversimplified anatomical comparisons have created misconceptions about primate and strepsirrhine phylogeny,
illustrated by the media attention surrounding the single "Ida" fossil in 2009. Strepsirrhines diverged from the
haplorhine primates near the beginning of the primate radiation between 55 and 90 mya. Older divergence dates are
based on genetic analysis estimates, while younger dates are based on the scarce fossil record. Lemuriform primates
may have evolved from either cercamoniines or sivaladapids, both of which were adapiforms that may have
originated in Asia. They were once thought to have evolved from adapids, a more specialized and younger branch of
adapiform primarily from Europe. Lemurs rafted to Madagascar between 47 and 54 mya, whereas the lorises split
from the African galagos around 40 mya and later colonized Asia. Both living and extinct strepsirrhines are
behaviorally diverse, although all are primarily arboreal (tree-dwelling). Most living lemuriforms are nocturnal,while most extinct adapiforms were diurnal. Both living and extinct groups primarily fed on fruit, leaves, and insects.
Many of today's strepsirrhines are endangered due to habitat destruction, hunting for bushmeat, and live capture for
the exotic pet trade.
Etymology
The taxonomic name Strepsirrhini derives from the Greek (strepsis or "a turning [inward])" and
(rhinos or "nose"), which refers to the appearance of the sinuous (comma-shaped) nostrils on the rhinarium or wet
nose.[1][2] The name was first used by French naturalist tienne Geoffroy Saint-Hilaire in 1812 as a subordinal rank
comparable to Platyrrhini (New World monkeys) and Catarrhini (Old World monkeys).[3]
In his description, hementioned "Les narines terminales et sinueuses" ("Nostrils terminal and winding").[4]
When British zoologist Reginald Innes Pocock revived Strepsirrhini and defined Haplorhini in 1918, he omitted the
second "r" from both ("Strepsirhini" and "Haplorhini" instead of "Strepsirrhini" and "Haplorrhini"), [5][6] although he
did not remove the second "r" from Platyrrhini or Catarrhini, both of which were also named by . Geoffroy in 1812.
Following Pocock, many researchers continued to spell Strepsirrhini with a single "r" until primatologists Paulina
Jenkins and Prue Napier pointed out the error in 1987.[7]
http://en.wikipedia.org/w/index.php?title=Primatologyhttp://en.wikipedia.org/w/index.php?title=Primatologyhttp://en.wikipedia.org/w/index.php?title=Haplorhinihttp://en.wikipedia.org/w/index.php?title=Reginald_Innes_Pocockhttp://en.wikipedia.org/w/index.php?title=Zoologyhttp://en.wikipedia.org/w/index.php?title=Species_descriptionhttp://en.wikipedia.org/w/index.php?title=Old_World_monkeyhttp://en.wikipedia.org/w/index.php?title=New_World_monkeyhttp://en.wikipedia.org/w/index.php?title=Order_%28biology%29http://en.wikipedia.org/w/index.php?title=%C3%89tienne_Geoffroy_Saint-Hilairehttp://en.wikipedia.org/w/index.php?title=Natural_historyhttp://en.wikipedia.org/w/index.php?title=Rhinariumhttp://en.wikipedia.org/w/index.php?title=Ancient_Greekhttp://en.wikipedia.org/w/index.php?title=Taxonomy_%28biology%29http://en.wikipedia.org/w/index.php?title=Exotic_pethttp://en.wikipedia.org/w/index.php?title=Bushmeathttp://en.wikipedia.org/w/index.php?title=Habitat_destructionhttp://en.wikipedia.org/w/index.php?title=Endangered_specieshttp://en.wikipedia.org/w/index.php?title=Diurnalityhttp://en.wikipedia.org/w/index.php?title=Nocturnalityhttp://en.wikipedia.org/w/index.php?title=Oceanic_dispersalhttp://en.wikipedia.org/w/index.php?title=Adapidaehttp://en.wikipedia.org/w/index.php?title=Sivaladapidaehttp://en.wikipedia.org/w/index.php?title=Cercamoniinaehttp://en.wikipedia.org/w/index.php?title=Fossilhttp://en.wikipedia.org/w/index.php?title=Molecular_clockhttp://en.wikipedia.org/w/index.php?title=Adaptive_radiationhttp://en.wikipedia.org/w/index.php?title=Darwiniushttp://en.wikipedia.org/w/index.php?title=Phylogeneticshttp://en.wikipedia.org/w/index.php?title=Grooming_clawhttp://en.wikipedia.org/w/index.php?title=Personal_groominghttp://en.wikipedia.org/w/index.php?title=Toothcombhttp://en.wikipedia.org/w/index.php?title=Vitamin_Chttp://en.wikipedia.org/w/index.php?title=Postorbital_barhttp://en.wikipedia.org/w/index.php?title=Orbit_%28anatomy%29http://en.wikipedia.org/w/index.php?title=Night_visionhttp://en.wikipedia.org/w/index.php?title=Tapetum_lucidumhttp://en.wikipedia.org/w/index.php?title=Placentationhttp://en.wikipedia.org/w/index.php?title=Bicornuate_uterushttp://en.wikipedia.org/w/index.php?title=Pheromonehttp://en.wikipedia.org/w/index.php?title=Vomeronasal_organhttp://en.wikipedia.org/w/index.php?title=Olfactory_bulbhttp://en.wikipedia.org/w/index.php?title=Rhinariumhttp://en.wikipedia.org/w/index.php?title=Symplesiomorphyhttp://en.wikipedia.org/w/index.php?title=Ecological_nichehttp://en.wikipedia.org/w/index.php?title=Natural_selectionhttp://en.wikipedia.org/w/index.php?title=Evolutionhttp://en.wikipedia.org/w/index.php?title=Living_fossilhttp://en.wikipedia.org/w/index.php?title=Simianhttp://en.wikipedia.org/w/index.php?title=Haplorhinihttp://en.wikipedia.org/w/index.php?title=Anatomyhttp://en.wikipedia.org/w/index.php?title=Tarsierhttp://en.wikipedia.org/w/index.php?title=Prosimian -
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Strepsirrhini 3
Evolutionary history
Primate phylogeny
Euarchonta
Scandentia (treeshrews)
Primatomorpha
Dermoptera (colugos)
Primates
Plesiadapiformes
Euprimates
Haplorhini
Simians
Tarsiers
Omomyiformes
Strepsirrhini
Adapiformes
LemuriformesLorisoids
Lemurs
Strepsirrhines and haplorhines diverged shortly after the emergence of the first true primates (euprimates). The relationship betweeneuprimates, treeshrews, colugos, and plesiadapiforms is less certain. Sometimes plesiadapiforms are grouped with the euprimatesunder the order Primates, colugos are grouped with primates under Primatomorpha, and all four are grouped under Euarchonta. [8]
Strepsirrhines include the extinct adapiforms and the lemuriform primates, which include lemurs and lorisoids
(lorises, pottos, and galagos).[9] The lemuriforms, and particularly the lemurs of Madagascar, are often portrayed
inappropriately as "living fossils" or as examples of "basal", or "inferior" primates. [10][11][12] These views have
historically hindered the understanding of mammalian evolution and the evolution of strepsirrhine traits,[10] such as
their reliance on smell (olfaction), characteristics of their skeletal anatomy, and their brain size, which is relatively
small. In the case of lemurs, natural selection has driven this isolated population of primates to diversify significantly
and fill a rich variety of ecological niches, despite their smaller and less complex brains compared to simians.[11][12]
The origin of the earliest primates, from which both the strepsirrhines and haplorhines (simians and tarsiers) evolved,
is a mystery. Both their place of origin and the group from which they evolved are uncertain. Although the fossil
record demonstrating their initial radiation across the Northern Hemisphere is very detailed,[13] the fossil record from
the tropicswhere primates most likely evolvedis very poor, particularly around the time that primates and other
major clades (groups consisting of an ancestor and all its descendants) of eutherian mammals were first appearing.
Consequently, geneticists and primatologists have used genetic analyses to determine the relatedness between
primate lineages and the amount of time since they diverged. Using this molecular clock, divergence dates for the
major primate lineages have suggested that primates evolved more than 8090 mya, nearly 40 million years before
the first primates appear in the fossil record.[14]
http://en.wikipedia.org/w/index.php?title=Molecular_clockhttp://en.wikipedia.org/w/index.php?title=Genetic_divergencehttp://en.wikipedia.org/w/index.php?title=Geneticshttp://en.wikipedia.org/w/index.php?title=Eutheriahttp://en.wikipedia.org/w/index.php?title=Cladehttp://en.wikipedia.org/w/index.php?title=Evolutionary_radiationhttp://en.wikipedia.org/w/index.php?title=Fossilhttp://en.wikipedia.org/w/index.php?title=Fossilhttp://en.wikipedia.org/w/index.php?title=Ecological_nichehttp://en.wikipedia.org/w/index.php?title=Natural_selectionhttp://en.wikipedia.org/w/index.php?title=Olfactionhttp://en.wikipedia.org/w/index.php?title=Evolution_of_mammalshttp://en.wikipedia.org/w/index.php?title=Basal_%28phylogenetics%29http://en.wikipedia.org/w/index.php?title=Living_fossilhttp://en.wikipedia.org/w/index.php?title=Galagohttp://en.wikipedia.org/w/index.php?title=Pottohttp://en.wikipedia.org/w/index.php?title=Lorishttp://en.wikipedia.org/w/index.php?title=Lemuriformeshttp://en.wikipedia.org/w/index.php?title=Euarchontahttp://en.wikipedia.org/w/index.php?title=Primatomorphahttp://en.wikipedia.org/w/index.php?title=Order_%28biology%29http://en.wikipedia.org/w/index.php?title=Plesiadapiformeshttp://en.wikipedia.org/w/index.php?title=Colugohttp://en.wikipedia.org/w/index.php?title=Treeshrewhttp://en.wikipedia.org/w/index.php?title=Lemurhttp://en.wikipedia.org/w/index.php?title=Lorisoideahttp://en.wikipedia.org/w/index.php?title=Adapiformeshttp://en.wikipedia.org/w/index.php?title=Omomyiformeshttp://en.wikipedia.org/w/index.php?title=Tarsierhttp://en.wikipedia.org/w/index.php?title=Simianhttp://en.wikipedia.org/w/index.php?title=Haplorhinihttp://en.wikipedia.org/w/index.php?title=Plesiadapiformeshttp://en.wikipedia.org/w/index.php?title=Primatehttp://en.wikipedia.org/w/index.php?title=Colugohttp://en.wikipedia.org/w/index.php?title=Primatomorphahttp://en.wikipedia.org/w/index.php?title=Treeshrewhttp://en.wikipedia.org/w/index.php?title=Euarchonta -
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Strepsirrhini 4
Early primates possessed adaptations for arboreal
locomotion that enabled maneuvering along fine
branches, as seen in this slender loris.
The early primates include both nocturnal and diurnal small-bodied
species,[15] and all were arboreal, with hands and feet specially adapted
for maneuvering on small branches.[16] Plesiadapiforms from the early
Paleocene are sometimes considered "archaic primates" because their
teeth resembled those of early primates, and because they possessed
arboreal adaptations, such as a divergent hallux (big toe). Althoughplesiadapiforms were closely related to primates, they may represent a
paraphyletic group from which primates may or may not have directly
evolved,[17] and some genera may have been more closely related to
colugos, also known as dermopterans or "flying lemurs", which are
thought to be closely related to primates.[18]
The first true primates (euprimates) do not appear in the fossil record
until the early Eocene (~55 mya), at which point they radiated across the Northern Hemisphere during a brief period
of rapid global warming known as the PaleoceneEocene Thermal Maximum.[19] These first primates included
Cantius, Donrussellia, Altanius, and Teilhardina on the northern continents,[20] as well as the more questionable
(and fragmentary) fossilAltiatlasius from Paleocene Africa.[21] These earliest fossil primates are often divided into
two groups, adapiforms (sometimes called adapids, or adapoids) and omomyiforms (or omomyidssuspected
relatives of tarsiers). Both appeared suddenly in the fossil record without transitional forms to indicate ancestry, [22]
and both groups were rich in diversity and were widespread throughout the Eocene. Although few fossils of extant
primate groupslemuriforms, tarsiers, or simiansare known from the Early to Middle Eocene,[23][24][25] evidence
from genetics and newer fossil finds suggest they may have been present during this early adaptive radiation.[19] The
divergence between strepsirrhines, simians, and tarsiers likely followed almost immediately after primates first
evolved.[21]
Adapiform evolution
Notharctus, a type of North
American adapiform, resembled
lemurs but it did not give rise to
them.
Adapiforms are extinct strepsirrhines that shared many anatomical similarities
with lemuriforms.[26] They are sometimes referred to as lemur-like primates,
although the diversity of both lemurs and adapiforms do not support this
analogy.[27] Like living strepsirrhines, adapiforms were extremely diverse,[10]
with at least 30 genera and 80 species known from the fossil record as of the
early 2000s.[28][29] They diversified across Laurasia during the Eocene,[26] some
reaching North America via a land bridge.[30] They were among the most
common mammals found in the fossil beds from that time. A few rare species
have also been found in northern Africa.[26] The most basal of the adapiforms
include the genera Cantius from North America and Europe and Donrussellia
from Europe. The latter bears the most ancestral traits, so it is often considered a
sister group or stem group of the other adapiforms.[31]
Adapiforms are often divided into three major groups:
Adapids were most commonly found in Europe, although the oldest
specimens (Adapoides from middle Eocene China) indicate that they most
likely evolved in Asia and immigrated. They died out in Europe during the
Grande Coupure, part of a significant extinction event at the end of the
Eocene.[32]
http://en.wikipedia.org/w/index.php?title=Extinction_eventhttp://en.wikipedia.org/w/index.php?title=Eocene%E2%80%93Oligocene_extinction_eventhttp://en.wikipedia.org/w/index.php?title=Adapoideshttp://en.wikipedia.org/w/index.php?title=Adapidaehttp://en.wikipedia.org/w/index.php?title=Stem_grouphttp://en.wikipedia.org/w/index.php?title=Sister_grouphttp://en.wikipedia.org/w/index.php?title=Symplesiomorphyhttp://en.wikipedia.org/w/index.php?title=Land_bridgehttp://en.wikipedia.org/w/index.php?title=Laurasiahttp://en.wikipedia.org/w/index.php?title=File%3ANotharctus_osborni.jpghttp://en.wikipedia.org/w/index.php?title=Notharctushttp://en.wikipedia.org/w/index.php?title=Adaptive_radiationhttp://en.wikipedia.org/w/index.php?title=Extant_taxonhttp://en.wikipedia.org/w/index.php?title=Transitional_fossilhttp://en.wikipedia.org/w/index.php?title=Omomyiformeshttp://en.wikipedia.org/w/index.php?title=List_of_fossil_primateshttp://en.wikipedia.org/w/index.php?title=Altiatlasiushttp://en.wikipedia.org/w/index.php?title=Teilhardinahttp://en.wikipedia.org/w/index.php?title=Altaniushttp://en.wikipedia.org/w/index.php?title=Donrusselliahttp://en.wikipedia.org/w/index.php?title=Cantiushttp://en.wikipedia.org/w/index.php?title=Paleocene%E2%80%93Eocene_Thermal_Maximumhttp://en.wikipedia.org/w/index.php?title=Global_warminghttp://en.wikipedia.org/w/index.php?title=Eocenehttp://en.wikipedia.org/w/index.php?title=Colugohttp://en.wikipedia.org/w/index.php?title=Genushttp://en.wikipedia.org/w/index.php?title=Paraphylyhttp://en.wikipedia.org/w/index.php?title=Halluxhttp://en.wikipedia.org/w/index.php?title=Paleocenehttp://en.wikipedia.org/w/index.php?title=Plesiadapiformeshttp://en.wikipedia.org/w/index.php?title=Diurnalityhttp://en.wikipedia.org/w/index.php?title=Nocturnalityhttp://en.wikipedia.org/w/index.php?title=File%3ALoris_lydekkerianus_nordicus_003.jpghttp://en.wikipedia.org/w/index.php?title=Slender_lorishttp://en.wikipedia.org/w/index.php?title=Arboreal_locomotionhttp://en.wikipedia.org/w/index.php?title=Arboreal_locomotion -
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Strepsirrhini 5
Notharctids, which most closely resembled some of Madagascar's lemurs, come from Europe and North America.
The European branch is often referred to as cercamoniines.[33] The North American branch thrived during the
Eocene, but did not survive into the Oligocene.[34][35] Like the adapids, the European branch were also extinct by
the end of the Eocene.[30]
Sivaladapids of southern and eastern Asia are best known from the Miocene, and the only adapiforms to survive
past the Eocene/Oligocene boundary (~34 mya). Their relationship to the other adapiforms remains unclear.[36]
They had vanished before the end of the Miocene (~7 mya).[26]
The relationship between adapiform and lemuriform primates has not been clearly demonstrated, so the position of
adapiforms as a paraphyletic stem group is questionable. Both molecular clock data and new fossil finds suggest that
the lemuriform divergence from the other primates and the subsequent lemur-lorisoid split both predate the
appearance of adapiforms in the early Eocene.[22] Yet new calibration methods may reconcile the discrepancies
between the molecular clock and the fossil record, favoring more recent divergence dates. [37] The fossil record
suggests that the strepsirrhine adapiforms and the haplorhine omomyiforms had been evolving independently before
the early Eocene, although their most basal members share enough dental similarities to suggest that they diverged
during the Paleocene (6555 mya).[26]
Lemuriform evolution
The earliest primates, including early strepsirrhines,diversified across the northern continents during the early
Eocene during a period of global warming.
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.[38] Instead, lemuriforms may be
descended from a very early branch of Asian cercamoniinesor sivaladapids that immigrated to northern Africa.[25][39]
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.[40][41][42]Djebelemurfrom Tunisia dates to the late early or early middle Eocene (52 to 46 mya) and has been
considered a cercamoniine,[43] but also may have been a stem lemuriform.[25] 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.[39]
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. [41] In isolation, the
lemurs diversified and filled the niches often filled by monkeys and apes today. [44] In Africa, the lorises and galagos
diverged during the Eocene, approximately 40 mya.[41] Unlike the lemurs in Madagascar, they have had to compete
with monkeys and apes, as well as other mammals.[11]
http://en.wikipedia.org/w/index.php?title=Oceanic_dispersalhttp://en.wikipedia.org/w/index.php?title=Plesiopithecushttp://en.wikipedia.org/w/index.php?title=Djebelemuridaehttp://en.wikipedia.org/w/index.php?title=Algeriahttp://en.wikipedia.org/w/index.php?title=Azibiidaehttp://en.wikipedia.org/w/index.php?title=Tunisiahttp://en.wikipedia.org/w/index.php?title=Djebelemurhttp://en.wikipedia.org/w/index.php?title=Ugandahttp://en.wikipedia.org/w/index.php?title=Kenyahttp://en.wikipedia.org/w/index.php?title=Faiyum_Oasishttp://en.wikipedia.org/w/index.php?title=Faiyum_Oasishttp://en.wikipedia.org/w/index.php?title=Wadilemurhttp://en.wikipedia.org/w/index.php?title=Saharagalagohttp://en.wikipedia.org/w/index.php?title=Karanisiahttp://en.wikipedia.org/w/index.php?title=Toothcombhttp://en.wikipedia.org/w/index.php?title=Philip_D._Gingerichhttp://en.wikipedia.org/w/index.php?title=Paleontologyhttp://en.wikipedia.org/w/index.php?title=File%3ABlakey_50moll.jpghttp://en.wikipedia.org/w/index.php?title=Eocenehttp://en.wikipedia.org/w/index.php?title=Miocenehttp://en.wikipedia.org/w/index.php?title=Sivaladapidaehttp://en.wikipedia.org/w/index.php?title=Oligocenehttp://en.wikipedia.org/w/index.php?title=Cercamoniinaehttp://en.wikipedia.org/w/index.php?title=Notharctidae -
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Strepsirrhini 6
Taxonomic classification
The suborder Strepsirrhini was
proposed by . Geoffroy in 1812.
Strepsirrhine primates were first grouped under the genus Lemur by Swedish
taxonomist Carl Linnaeus in the 10th edition of Systema Naturae published in
1758. At the time, only three species were recognized, one of which (a colugo) is
no longer recognized as a primate.[45] In 1785, Dutch naturalist Pieter Boddaert
divided the genusLemurinto two genera:Prosimia for the lemurs, colugos, andtarsiers and Tardigradus for the lorises.[46] Ten years later, . Geoffroy and
Georges Cuvier grouped the tarsiers and galagos due to similarities in their
hindlimb morphology, a view supported by German zoologist Johann Karl
Wilhelm Illiger, who placed them in the family Macrotarsi while placing the
lemurs and tarsiers in the family Prosimia (Prosimii) in 1811. The use of the
tarsier-galago classification continued for many years until 1898, when Dutch
zoologist Ambrosius Hubrecht demonstrated two different types of placentation
(formation of a placenta) in the two groups.[47]
English comparative anatomist William Henry Flower created the suborderLemuroidea in 1883 to distinguish these primates from the simians, which were grouped under English biologist St.
George Jackson Mivart's suborder Anthropoidea (=Simiiformes). According to Flower, the suborder Lemuroidea
contained the families Lemuridae (lemurs, lorises, and galagos), Chiromyidae (aye-aye), and Tarsiidae (tarsiers).
Lemuroidea was later replaced by Illiger's suborder Prosimii.[47] Many years earlier, in 1812, . Geoffroy first
named the suborder Strepsirrhini, in which he included the tarsiers.[48] This taxonomy went unnoticed until 1918,
when Pocock compared the structure of the nose and reinstated the use of the suborder Strepsirrhini, while also
moving the tarsiers and the simians into a new suborder, Haplorhini.[5][47] It was not until 1953, when British
anatomist William Charles Osman Hill wrote an entire volume on strepsirrhine anatomy, that Pocock's taxonomic
suggestion became noticed and more widely used. Since then, primate taxonomy has shifted between
Strepsirrhini-Haplorhini and Prosimii-Anthropoidea multiple times.[47]
Most of the academic literature provides a basic framework for primate taxonomy, usually including several
potential taxonomic schemes.[49] Although most experts agree upon phylogeny,[50] many disagree about nearly every
level of primate classification.[49]
Competing strepsirrhine taxonomic nomenclature
2 infraorders[51]
3 infraorders[52]
http://en.wikipedia.org/w/index.php?title=Phylogeneticshttp://en.wikipedia.org/w/index.php?title=William_Charles_Osman_Hillhttp://en.wikipedia.org/w/index.php?title=Aye-ayehttp://en.wikipedia.org/w/index.php?title=St._George_Jackson_Mivarthttp://en.wikipedia.org/w/index.php?title=St._George_Jackson_Mivarthttp://en.wikipedia.org/w/index.php?title=Biologyhttp://en.wikipedia.org/w/index.php?title=Taxonomic_rankhttp://en.wikipedia.org/w/index.php?title=William_Henry_Flowerhttp://en.wikipedia.org/w/index.php?title=Comparative_anatomyhttp://en.wikipedia.org/w/index.php?title=Placentahttp://en.wikipedia.org/w/index.php?title=Placentationhttp://en.wikipedia.org/w/index.php?title=Ambrosius_Hubrechthttp://en.wikipedia.org/w/index.php?title=Family_%28biology%29http://en.wikipedia.org/w/index.php?title=Johann_Karl_Wilhelm_Illigerhttp://en.wikipedia.org/w/index.php?title=Johann_Karl_Wilhelm_Illigerhttp://en.wikipedia.org/w/index.php?title=Morphology_%28biology%29http://en.wikipedia.org/w/index.php?title=Pieter_Boddaerthttp://en.wikipedia.org/w/index.php?title=10th_edition_of_Systema_Naturaehttp://en.wikipedia.org/w/index.php?title=Carl_Linnaeushttp://en.wikipedia.org/w/index.php?title=File%3AGeoffroy72.jpghttp://en.wikipedia.org/w/index.php?title=%C3%89tienne_Geoffroy_Saint-Hilaire -
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Strepsirrhini 7
Suborder Strepsirrhini
Infraorder Adapiformes
Superfamily Adapoidea
Family Adapidae
Family Notharctidae
Family Sivaladapidae
Infraorder Lemuriformes
Superfamily Lemuroidea
Family Archaeolemuridae
Family Cheirogaleidae
Family Daubentoniidae
Family Indriidae
Family Lemuridae
Family Lepilemuridae
Family Megaladapidae
Family Palaeopropithecidae
Superfamily Lorisoidea
Family Lorisidae
Family Galagidae
Suborder Strepsirrhini
Infraorder Adapiformes
Superfamily Adapoidea
Family Adapidae
Family Notharctidae
Family Sivaladapidae
Infraorder Lemuriformes
Superfamily Lemuroidea
Family Archaeolemuridae
Family Cheirogaleidae
Family Daubentoniidae
Family Indriidae
Family Lemuridae
Family Lepilemuridae
Family Megaladapidae
Family Palaeopropithecidae
Infraorder Lorisiformes
Superfamily Lorisoidea
Family Lorisidae
Family Galagidae
Subordinal controversies
The most commonly recurring debate in primatology during the 1970s, 1980s, and early 2000s concerned the
phylogenetic position of tarsiers compared to both simians and the other prosimians. [53][54] Tarsiers are most often
placed in either the suborder Haplorhini with the simians or in the suborder Prosimii with the strepsirrhines.[44]
Prosimii is one of the two traditional primate suborders and is based on evolutionary grades (groups united by
anatomical traits) rather than phylogenetic clades, while the Strepsirrhini-Haplorrhini taxonomy was based on
evolutionary relationships.[2] Yet both systems persist because the Prosimii-Anthropoidea taxonomy is familiar andfrequently seen in the research literature and textbooks.[14]
Strepsirrhines are traditionally characterized by several symplesiomorphic (ancestral) traits not shared with the
simians, particularly the rhinarium.[55][56][44][57] Other symplesiomorphies include long snouts, convoluted
maxilloturbinals, relatively large olfactory bulbs, and smaller brains. The toothcomb is a synapomorphy (shared,
derived trait) seen among lemuriforms,[44] although it is frequently and incorrectly used to define the strepsirrhine
clade.[58][44] However, one group of strepsirrhines lacks the toothcombthe adapiforms. The toothcomb is therefore
the primary hallmark of the lemuriforms,[31] although at least one family only retains it in modified form. [59]
Strepsirrhine primates are also united in possessing an epitheliochorial placenta.[44] Unlike the tarsiers and simians,
strepsirrhines are capable of producing their own vitamin C and do not need it supplied in their diet. [60] Further
genetic evidence for the relationship between tarsiers and simians as a haplorhine clade is the shared possession of
three SINE markers.[61]
Because of their historically mixed assemblages which included tarsiers and close relatives of primates, both
Prosimii and Strepsirrhini have been considered wastebasket taxa for "lower primates".[62][63] Regardless, the
strepsirrhine and haplorrhine clades are generally accepted and viewed as the preferred taxonomic division. Yet,
tarsiers still closely resemble both strepsirrhines and simians in different ways,[14] and since the early split between
strepsirrhines, tarsiers, and simians is ancient and hard to resolve,[64] a third taxonomic arrangement with three
suborders is sometimes used: Prosimii, Tarsiiformes, and Anthropoidea.[9] More often, the term "prosimian" is no
longer used in official taxonomy,[65] but is still used to illustrate the behavioral ecology of tarsiers relative to the
other primates.[57]
http://en.wikipedia.org/w/index.php?title=Wastebasket_taxonhttp://en.wikipedia.org/w/index.php?title=Retrotransposon%23SINEshttp://en.wikipedia.org/w/index.php?title=Vitamin_Chttp://en.wikipedia.org/w/index.php?title=Biosynthesishttp://en.wikipedia.org/w/index.php?title=Placentation%23Placentation_in_mammalshttp://en.wikipedia.org/w/index.php?title=Synapomorphyhttp://en.wikipedia.org/w/index.php?title=Olfactory_bulbhttp://en.wikipedia.org/w/index.php?title=Nasal_conchahttp://en.wikipedia.org/w/index.php?title=Snouthttp://en.wikipedia.org/w/index.php?title=Symplesiomorphyhttp://en.wikipedia.org/w/index.php?title=Evolutionary_gradehttp://en.wikipedia.org/w/index.php?title=Galagidaehttp://en.wikipedia.org/w/index.php?title=Lorisidaehttp://en.wikipedia.org/w/index.php?title=Lorisoideahttp://en.wikipedia.org/w/index.php?title=Sloth_lemurhttp://en.wikipedia.org/w/index.php?title=Megaladapishttp://en.wikipedia.org/w/index.php?title=Sportive_lemurhttp://en.wikipedia.org/w/index.php?title=Lemuridaehttp://en.wikipedia.org/w/index.php?title=Indriidaehttp://en.wikipedia.org/w/index.php?title=Aye-ayehttp://en.wikipedia.org/w/index.php?title=Cheirogaleidaehttp://en.wikipedia.org/w/index.php?title=Monkey_lemurhttp://en.wikipedia.org/w/index.php?title=Lemurhttp://en.wikipedia.org/w/index.php?title=Sivaladapidaehttp://en.wikipedia.org/w/index.php?title=Notharctidaehttp://en.wikipedia.org/w/index.php?title=Adapidaehttp://en.wikipedia.org/w/index.php?title=Adapiformes -
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Strepsirrhini 8
In addition to the controversy over tarsiers, the debate over the origins of simians once called the strepsirrhine clade
into question. Arguments for an evolutionary link between adapiforms and simians made by paleontologists
Gingerich, Elwyn Simons, Tab Rasmussen, and others could have potentially excluded adapiforms from
Strepsirrhini.[66][67] In 1975, Gingerich proposed a new suborder, Simiolemuriformes,[68] to suggest that
strepsirrhines are more closely related to simians than tarsiers. [69] However, no clear relationship between the two
had been demonstrated by the early 2000s.[67] The idea reemerged briefly in 2009 during the media attention
surrounding Darwinius masillae (dubbed "Ida"), a cercamoniine from Germany that was touted as a "missing link
between humans and earlier primates" (simians and adapiforms).[70] However, the cladistic analysis was flawed and
the phylogenetic inferences and terminology were vague.[70] Although the authors noted that Darwinius was not a
"fossil lemur", they did emphasize the absence of a toothcomb,[71] which adapiforms did not possess.[31]
Infraordinal classification and clade terminology
Strepsirrhini phylogeny[39]
Adapiformes
stem lemuriforms
lemuriforms orcrown strepsirrhines
lemuroid cladeDaubentoniidae
other lemurs
lorisoid cladelorises
galagos
Within Strepsirrhini, two common classifications include either two infraorders (Adapiformes and Lemuriformes)[72]
or three infraorders (Adapiformes, Lemuriformes, Lorisiformes).[73] A less common taxonomy places the aye-aye
(Daubentoniidae) in its own infraorder, Chiromyiformes.[74] 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.[9] Regardless of the infraordinal taxonomy,
Strepsirrhini is composed of three ranked superfamilies and 14 families, seven of which are extinct. [72] Three ofthese extinct families included the recently extinct giant lemurs of Madagascar,[75] many of which died out within
the last 1,000 years following human arrival on the island.[76]
http://en.wikipedia.org/w/index.php?title=Subfossil_lemurhttp://en.wikipedia.org/w/index.php?title=Crown_grouphttp://en.wikipedia.org/w/index.php?title=Stem_grouphttp://en.wikipedia.org/w/index.php?title=Transitional_fossilhttp://en.wikipedia.org/w/index.php?title=Darwiniushttp://en.wikipedia.org/w/index.php?title=Elwyn_Simons -
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Strepsirrhini 9
The media attention over "Ida"
played upon confused terminology
and misconceptions about
strepsirrhines.
When Strepsirrhini is divided into two infraorders, the clade containing all
toothcombed primates can be called "lemuriforms".[72] When it is divided into
three infraorders, the term "lemuriforms" refers only to Madagascar's lemurs,[44]
and the toothcombed primates are referred to as either "crown strepsirrhines"[36]
or "extant strepsirrhines".[2] Confusion of this specific terminology with the
general term "strepsirrhine", along with oversimplified anatomical comparisonsand vague phylogenetic inferences, can lead to misconceptions about primate
phylogeny and misunderstandings about primates from the Eocene, as seen with
the media coverage ofDarwinius.[70] Because the skeletons of adapiforms share
strong similarities with those of lemurs and lorises, researchers have often
referred to them as primitive strepsirrhines,[77] lemur ancestors, or a sister group
to the living strepsirrhines. They are included in Strepsirrhini,[2] and are
considered basal members of the clade.[78] 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 anatomicaldifferences 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).[26]
The first fossil primate described was the adapiform Adapis parisiensis by French naturalist Georges Cuvier in
1821,[43] who compared it to a hyrax ("le Daman"), then considered a member of a now obsolete group called
pachyderms.[79] It was not recognized as a primate until it was reevaluated in the early 1870s. [28][43] Originally,
adapiforms were all included under the family Adapidae, which was divided into two or three subfamilies: Adapinae,
Notharctinae, and sometimes Sivaladapinae. All North American adapiforms were lumped under Notharctinae, while
the Old World forms were usually assigned to Adapinae. Around the 1990s, two distinct groups of European"adapids" began to emerge, based on differences in the postcranial skeleton and the teeth. One of these two European
forms was identified as cercamoniines, which were allied with the notharctids found mostly in North America, while
the other group falls into the traditional adapid classification.[31] The three major adapiform divisions are now
typically regarded as three families within Adapiformes (Notharctidae, Adapidae and Sivaladapidae), but other
divisions ranging from one to five families are used as well.[28]
Anatomy and physiology
Strepsirrhines have a reflective layer in the eye,
called a tapetum lucidum that helps them see
better at night.
All lemuriforms possess a specialized dental structure called a
toothcomb,[2][80] with the exception of the aye-aye, in which the
structure has been modified into two continually growing
(hypselodont) incisors (or canine teeth), similar to those of rodents.
The toothcomb consists of either two or four procumbent lower
incisors and procumbent lower canine teeth followed by a
canine-shaped premolar.[81] It is used to comb the fur during oral
grooming. Shed hairs that accumulate between the teeth of the
toothcomb are removed by the sublingua or "under-tongue".[82]
Lemuriforms also possess a grooming claw on the second digit of each
foot for scratching.[2][80][82] Adapiforms did not possess a toothcomb.
Instead, their lower incisors varied in orientationfrom somewhat
http://en.wikipedia.org/w/index.php?title=Sublinguahttp://en.wikipedia.org/w/index.php?title=Sublinguahttp://en.wikipedia.org/w/index.php?title=Grooming_clawhttp://en.wikipedia.org/w/index.php?title=Grooming_clawhttp://en.wikipedia.org/w/index.php?title=Tapetum_lucidumhttp://en.wikipedia.org/w/index.php?title=Grooming_clawhttp://en.wikipedia.org/w/index.php?title=Sublinguahttp://en.wikipedia.org/w/index.php?title=Premolarhttp://en.wiktionary.org/wiki/procumbenthttp://en.wikipedia.org/w/index.php?title=Rodenthttp://en.wikipedia.org/w/index.php?title=Canine_toothhttp://en.wikipedia.org/w/index.php?title=Incisorhttp://en.wikipedia.org/w/index.php?title=File%3AGarnettsGalago_CincinnatiZoo.jpghttp://en.wikipedia.org/w/index.php?title=Tapetum_lucidumhttp://en.wikipedia.org/w/index.php?title=Postcraniahttp://en.wikipedia.org/w/index.php?title=Pachydermatahttp://en.wikipedia.org/w/index.php?title=Hyraxhttp://en.wikipedia.org/w/index.php?title=Georges_Cuvierhttp://en.wikipedia.org/w/index.php?title=Adapishttp://en.wikipedia.org/w/index.php?title=File%3ADarwinius_masillae_PMO_214.214.jpghttp://en.wikipedia.org/w/index.php?title=Darwinius -
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Strepsirrhini 10
procumbent to somewhat verticaland the lower canines were projected upwards and were often prominent.
Adapiforms may have had a grooming claw,[31] but there is little evidence of this.[83]
Like all primates, strepsirrhine orbits (eye sockets) have a postorbital bar, a protective ring of bone created by a
connection between the frontal and zygomatic bones.[84] Both living and extinct strepsirrhines lack a thin wall of
bone behind the eye, referred to as postorbital closure, which is only seen in haplorhine primates.[84][85] Although
the eyes of strepsirrhines point forward, giving stereoscopic vision,[84]
the orbits do not face fully forward.[85]
Among living strepsirrhines, most or all species are thought to possess a reflective layer behind the retina of the eye,
called a tapetum lucidum, which improves vision in low light,[86][80] but they lack a fovea, which improves day
vision. This differs from tarsiers, which lack a tapetum lucidum but possess a fovea.[87]
Strepsirrhines are characterized by a typically
longer snout and wet nose compared to
haplorhine primates.
Strepsirrhine primates have a brain relatively comparable to or slightly
larger in size than most mammals.[10][44] Compared to simians,
however, they have a relatively small brain-to-body size ratio.[85]
Strepsirrhines are also traditionally noted for their unfused mandibular
symphysis (two halves of the lower jaw),[85] however, fusion of the
mandibular symphysis was common in adapiforms,[88] notably
Notharctus.[89] Also, several extinct giant lemurs exhibited a fusedmandibular symphysis.[90]
Many nocturnal species have large, independently movable ears,[91][92]
although there are significant differences in sizes and shapes of the ear
between species.[93] The structure of the middle and inner ear of
strepsirrhines differs between the lemurs and lorisoids. In lemurs, the
tympanic cavity, which surrounds the middle ear, is expanded. This leaves the ectotympanic ring, which supports the
eardrum, free within the auditory bulla.[94] This trait is also seen in adapiforms.[88] In lorisoids, however, the
tympanic cavity is smaller and the ectotympanic ring becomes attached to the edge of the auditory bulla. The
tympanic cavity in lorisoids also has two accessory air spaces, which are not present in lemurs. Both lorisoids and
cheirogaleid lemurs have replaced the internal carotid artery with an enlarged ascending pharyngeal artery.[94]
Strepsirrhines also possess distinctive features in their tarsus (ankle bones) that differentiate them from haplorhines,
such as a sloping talo-fibular facet (the face where the talus bone and fibula meet) and a difference in the location of
the position of the flexor fibularis tendon on the talus.[95] These differences give strepsirrhines the ability to make
more complex rotations of the ankle and indicate that their feet are habitually inverted, or turned inward, an
adaptation for grasping vertical supports.[96]
Sexual dichromatism (different coloration patterns between males and females) can be seen in most brown lemur
species,[97] but otherwise lemurs show very little if any difference in body size or weight between sexes. This lack of
sexual dimorphism is not characteristic of all strepsirrhines.[98] Some adapiforms were sexually dimorphic, with
males bearing a larger sagittal crest (a ridge of bone on the top of the skull to which jaw muscles attach) and canineteeth.[99] Lorisoids exhibit some sexual dimorphism,[98] but males are typically no more than 20 percent larger than
females.[100]
http://en.wikipedia.org/w/index.php?title=Sagittal_cresthttp://en.wikipedia.org/w/index.php?title=Sexual_dimorphismhttp://en.wikipedia.org/w/index.php?title=Flexor_hallucis_longus_musclehttp://en.wikipedia.org/w/index.php?title=Fibulahttp://en.wikipedia.org/w/index.php?title=Talus_bonehttp://en.wikipedia.org/w/index.php?title=Tarsus_%28skeleton%29http://en.wikipedia.org/w/index.php?title=Ascending_pharyngeal_arteryhttp://en.wikipedia.org/w/index.php?title=Internal_carotid_arteryhttp://en.wikipedia.org/w/index.php?title=Cheirogaleidaehttp://en.wikipedia.org/w/index.php?title=Auditory_bullahttp://en.wikipedia.org/w/index.php?title=Eardrumhttp://en.wikipedia.org/w/index.php?title=Ectotympanic_ringhttp://en.wikipedia.org/w/index.php?title=Tympanic_cavityhttp://en.wikipedia.org/w/index.php?title=Inner_earhttp://en.wikipedia.org/w/index.php?title=Middle_earhttp://en.wikipedia.org/w/index.php?title=Symphysis_mentihttp://en.wikipedia.org/w/index.php?title=Symphysis_mentihttp://en.wikipedia.org/w/index.php?title=File%3AEulemur_mongoz_%28male_-_face%29.jpghttp://en.wikipedia.org/w/index.php?title=Haplorhinihttp://en.wikipedia.org/w/index.php?title=Foveahttp://en.wikipedia.org/w/index.php?title=Tapetum_lucidumhttp://en.wikipedia.org/w/index.php?title=Retinahttp://en.wikipedia.org/w/index.php?title=Stereopsishttp://en.wikipedia.org/w/index.php?title=Zygomatic_bonehttp://en.wikipedia.org/w/index.php?title=Frontal_bonehttp://en.wikipedia.org/w/index.php?title=Postorbital_bar -
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Strepsirrhini 11
Rhinarium and olfaction
The noses of five prosimian primates: A) dwarf
lemur, B) greater galago, C) lesser galago, D)
aye-aye, and E) a tarsier.A through D possess a rhinarium and are
strepsirrhines, whereas E does not and is a
haplorhine.
Strepsirrhines have a long snout that ends in a moist and
touch-sensitive rhinarium, similar to that of dogs and many other
mammals. The rhinarium is surrounded by vibrissae that are also
sensitive to touch. Convoluted maxilloturbinals on the inside of their
nose filter, warm, and moisten the incoming air, while olfactoryreceptors of the main olfactory system lining the ethmoturbinals detect
airborne smells.[44][101] The olfactory bulbs of lemurs are comparable
in size to those of other arboreal mammals.[44]
The surface of the rhinarium does not have any olfactory receptors, so
it is not used for smell in terms of detecting volatile substances.
Instead, it has sensitive touch receptors (Merkel cells). The rhinarium,
upper lip, and gums are tightly connected by a fold of mucous
membrane called the philtrum, which runs from the tip of the nose to
the mouth.[102] The upper lip is constrained by this connection and has
fewer nerves to control movement, which leaves it less mobile than the
upper lips of simians.[103][104] The philtrum creates a gap (diastema)
between the roots of the first two upper incisors.[102][105]
The strepsirrhine rhinarium can collect relatively non-volatile, fluid-based chemicals (traditionally categorized as
pheromones) and transmit them to the vomeronasal organ (VNO),[106] which is located below and in front of the
nasal cavity, above the mouth.[107] The VNO is an encased duct-like structure made of cartilage[108] and is isolated
from the air passing through the nasal cavity.[109] The VNO is connected to the mouth through nasopalatine ducts
(also known as the incisive foramen), which pass through the hard palate at the top, front of the mouth. [107] Fluids
traveling from the rhinarium to the mouth and then up the nasopalatine ducts to the VNO are detected, and
information is relayed to the accessory olfactory bulb, which is relatively large in strepsirrhines.[110] From theaccessory olfactory bulb, information is sent to the amygdala, which handles emotions, and then to the
hypothalamus, which handles basic body functions and metabolic processes. This neural pathway differs from that
used by the main olfactory system.[111]
All lemuriforms have a VNO, as do tarsiers and some New World monkeys. [112] Adapiforms exhibit the gap
between the upper incisors, which indicates the presence of a VNO, but there is some disagreement over whether or
not they possessed a rhinarium.[113]
Reproductive physiology
Extant strepsirrhines have an epitheliochorial placenta,[44]
where the maternal blood does not come in direct contactwith the fetal chorion like it does in the hemochorial placenta of haplorhines. The strepsirrhine uterus has two
distinct chambers (bicornuate).[103] Despite having similar gestation periods to comparably sized haplorhines, fetal
growth rates are generally slower in strepsirrhines, which results in newborn offspring that are as little as one-third
the size of haplorhine newborns.[103][114] Extant strepsirrhines also have a lower basal metabolic rate, which elevates
in females during gestation, putting greater demands on the mother.[115]
Most primates have two mammary glands,[116] but the number and positions vary between species within
strepsirrhines.[117] Lorises have two pairs,[118] while others, like the ring-tailed lemur, have one pair on the chest
(pectoral).[119] The aye-aye also has two mammary glands, but they are located near the groin (inguinal). [120] In
females, the clitoris is sometimes enlarged and pendulous, resembling the male penis, which can make gender
identification difficult for human observers.[121] The clitoris may also have a bony structure in it, similar to thebaculum (penis bone) in males.[122] Most male primates have a baculum, but it is typically larger in strepsirrhines
http://en.wikipedia.org/w/index.php?title=Baculumhttp://en.wikipedia.org/w/index.php?title=Baculumhttp://en.wikipedia.org/w/index.php?title=Baculumhttp://en.wikipedia.org/w/index.php?title=Pseudo-penishttp://en.wikipedia.org/w/index.php?title=Clitorishttp://en.wikipedia.org/w/index.php?title=Mammary_glandhttp://en.wikipedia.org/w/index.php?title=Basal_metabolic_ratehttp://en.wikipedia.org/w/index.php?title=Gestation_periodhttp://en.wikipedia.org/w/index.php?title=Uterushttp://en.wikipedia.org/w/index.php?title=Placentation%23Placentation_in_mammalshttp://en.wikipedia.org/w/index.php?title=Chorionhttp://en.wikipedia.org/w/index.php?title=Hypothalamushttp://en.wikipedia.org/w/index.php?title=Amygdalahttp://en.wikipedia.org/w/index.php?title=Olfaction%23Accessory_olfactory_systemhttp://en.wikipedia.org/w/index.php?title=Hard_palatehttp://en.wikipedia.org/w/index.php?title=Incisive_foramenhttp://en.wikipedia.org/w/index.php?title=Cartilagehttp://en.wikipedia.org/w/index.php?title=Duct_%28anatomy%29http://en.wikipedia.org/w/index.php?title=Nasal_cavityhttp://en.wikipedia.org/w/index.php?title=Vomeronasal_organhttp://en.wikipedia.org/w/index.php?title=Pheromonehttp://en.wikipedia.org/w/index.php?title=Diastema_%28dentistry%29http://en.wikipedia.org/w/index.php?title=Philtrumhttp://en.wikipedia.org/w/index.php?title=Mucous_membranehttp://en.wikipedia.org/w/index.php?title=Mucous_membranehttp://en.wikipedia.org/w/index.php?title=Merkel_cellhttp://en.wikipedia.org/w/index.php?title=Volatility_%28chemistry%29http://en.wikipedia.org/w/index.php?title=Nasal_conchahttp://en.wikipedia.org/w/index.php?title=Olfactory_systemhttp://en.wikipedia.org/w/index.php?title=Olfactory_receptorhttp://en.wikipedia.org/w/index.php?title=Olfactory_receptorhttp://en.wikipedia.org/w/index.php?title=Whiskershttp://en.wikipedia.org/w/index.php?title=Rhinariumhttp://en.wikipedia.org/w/index.php?title=File%3AProsimian_noses.jpghttp://en.wikipedia.org/w/index.php?title=Tarsierhttp://en.wikipedia.org/w/index.php?title=Aye-ayehttp://en.wikipedia.org/w/index.php?title=Lesser_bushbabyhttp://en.wikipedia.org/w/index.php?title=Greater_galagohttp://en.wikipedia.org/w/index.php?title=Dwarf_lemurhttp://en.wikipedia.org/w/index.php?title=Dwarf_lemur -
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Strepsirrhini 12
and usually forked at the tip.[123]
Behavior
Like other primates, infants often
cling to their mother's fur.
Approximately three-quarters of all extant strepsirrhine species are nocturnal,
sleeping in nests made from dead leaves or tree hollows during the day. [124] All
of the lorisoids from continental Africa and Asia are nocturnal, a circumstancethat minimizes their competition with the simian primates of the region, which
are diurnal. The lemurs of Madagascar, living in the absence of simians, are more
variable in their activity cycles. The aye-aye, mouse lemurs, woolly lemurs, and
sportive lemurs are nocturnal, while ring-tailed lemurs and most of their kin,
sifakas, and indri are diurnal.[78] Yet some or all of the brown lemurs (Eulemur)
are cathemeral, which means they may be active during the day or night,
depending on factors such as temperature and predation.[125] Many extant
strepsirrhines are well adapted for nocturnal activity due to their relatively large
eyes; large, movable ears; sensitive tactile hairs; strong sense of smell; and the
tapetum lucidum behind the retina.[92] Among the adapiforms, most are
considered diurnal, with the exception ofPronycticebus and Godinotia from
Middle Eocene Europe, both of which had large orbits that suggest
nocturnality.[78][88]
Reproduction in most strepsirrhine species tends to be seasonal, particularly in lemurs. Key factors that affect
seasonal reproduction include the length of the wet season, subsequent food availability, and the maturation time of
the species.[126] Like other primates, strepsirrhines are relatively slow breeders compared to other mammals. Their
gestation period and interbirth intervals are usually long, and the young develop slowly, just like in haplorhine
primates.[103][114] Unlike simians, some strepsirrhines produce two or three offspring, although some produce only a
single offspring. Those that produce multiple offspring tend to build nests for their young. These two traits arethought to be plesiomorphic (ancestral) for primates.[103] The young are precocial (relatively mature and mobile) at
birth, but not as coordinated as ungulates (hoofed mammals).[44] Infant care by the mother is relatively prolonged
compared to many other mammals, and in some cases, the infants cling to the mother's fur with their hands and
feet.[103]
Despite their relatively smaller brains compared to other primates, lemurs have demonstrated levels of technical
intelligence in problem solving that are comparable to those of seen in simians. However, their social intelligence
differs, often emphasizing within-group competition over cooperation, which may be due to adaptations for their
unpredictable environment.[127] Although lemurs have not been observed using objects as tools in the wild, they can
be trained to use objects as tools in captivity and demonstrate a basic understanding about the functional properties
of the objects they are using.[128]
Social systems and communication
The nocturnal strepsirrhines have been traditionally described as "solitary", although this term is no longer favored
by the researchers who study them.[129] Many are considered "solitary foragers", but many exhibit complex and
diverse social organization, often overlapping home ranges, initiating social contact at night, and sharing sleeping
sites during the day. Even the mating systems are variable, as seen in wooly lemurs, which live in monogamous
breeding pairs.[130] Because of this social diversity among these solitary but social primates, whose level of social
interaction is comparable to that of diurnal simians,[129] alternative classifications have been proposed to emphasize
their gregarious, dispersed, or solitary nature.
[131]
http://en.wikipedia.org/w/index.php?title=Breeding_pairhttp://en.wikipedia.org/w/index.php?title=Solitary_but_socialhttp://en.wikipedia.org/w/index.php?title=Breeding_pairhttp://en.wikipedia.org/w/index.php?title=Solitary_but_socialhttp://en.wikipedia.org/w/index.php?title=Solitary_but_socialhttp://en.wikipedia.org/w/index.php?title=Breeding_pairhttp://en.wikipedia.org/w/index.php?title=Monogamous_pairing_in_animalshttp://en.wikipedia.org/w/index.php?title=Home_rangehttp://en.wikipedia.org/w/index.php?title=Tool_use_by_animalshttp://en.wikipedia.org/w/index.php?title=Ungulatehttp://en.wikipedia.org/w/index.php?title=Precocialhttp://en.wikipedia.org/w/index.php?title=Symplesiomorphyhttp://en.wikipedia.org/w/index.php?title=Godinotiahttp://en.wikipedia.org/w/index.php?title=Pronycticebushttp://en.wikipedia.org/w/index.php?title=Predationhttp://en.wikipedia.org/w/index.php?title=Cathemeralhttp://en.wikipedia.org/w/index.php?title=True_lemurhttp://en.wikipedia.org/w/index.php?title=Indrihttp://en.wikipedia.org/w/index.php?title=Sifakahttp://en.wikipedia.org/w/index.php?title=Ring-tailed_lemurhttp://en.wikipedia.org/w/index.php?title=Sportive_lemurhttp://en.wikipedia.org/w/index.php?title=Woolly_lemurhttp://en.wikipedia.org/w/index.php?title=Mouse_lemurhttp://en.wikipedia.org/w/index.php?title=File%3ACaptive_N._bengalensis_from_Laos_with_6-week_baby.JPG -
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Strepsirrhini 13
Among extant strepsirrhines, only the diurnal and cathemeral lemurs have evolved to live in multi-male/multi-female
groups, comparable to most living simians.[130][132] This social trait, seen in two extant lemur families (Indriidae and
Lemuridae), is thought to have evolved independently.[133] Group sizes are smaller in social lemurs than in simians,
and despite the similarities, the community structures differ.[134] Female dominance, which is rare in simians, is
fairly common in lemurs.[135] Strepsirrhines spend a considerable amount of time grooming each other
(allogrooming).[136] When lemuriform primates groom, they lick the fur and then comb it with their toothcomb.
They also use their grooming claw to scratch places they cannot reach with their mouth.[82]
Like New World monkeys, strepsirrhines rely on scent marking for much of their communication. This involves
smearing secretions from epidermal scent glands on tree branches, along with urine and feces. In some cases,
strepsirrhines may anoint themselves with urine (urine washing). Body postures and gestures may be used, although
the long snout, non-mobile lips, and reduced facial enervation restrict the use of facial expressions in strepsirrhines.
Short-range calls, long-range calls, and alarm calls are also used. [137] Nocturnal species are more constrained by the
lack of light, so their communication systems differ from those of diurnal species, often using long-range calls to
claim their territory.[138]
Locomotion
Dwarf lemurs (top) are arboreal quadrupeds while galagos (bottom) favor vertical clinging and leaping.
Living strepsirrhines are predominantly arboreal, with only the ring-tailed lemur spending considerable time on the
ground.[139] Most species move around quadrupedally (on four legs) in the trees, including five genera of smaller,
nocturnal lemurs.[124][139] Galagos, indriids, sportive lemurs, and bamboo lemurs leap from vertical surfaces,[139]
and the indriids are highly specialized for vertical clinging and leaping. [124] Lorises are slow-moving, deliberate
climbers.[139]
Analyses of extinct adapiforms postcranial skeletons suggest a variety of locomotor behavior.[139] The Europeanadapids Adapis, Palaeolemur, and Leptadapis shared adaptations for slow climbing like the lorises, although they
may have been quadrupedal runners like small New World monkeys. Both Notharctus and Smilodectes from North
America andEuropolemurfrom Europe exhibit limb proportions and joint surfaces comparable to vertical clinging
and leaping lemurs, but were not as specialized as indriids for vertical clinging, suggesting that they ran along
branches and did not leap as much.[88][139] Notharctids Cantius andPronycticebus appear to have been agile arboreal
quadrupeds, with adaptations comparable to the brown lemurs.[139]
http://en.wikipedia.org/w/index.php?title=Europolemurhttp://en.wikipedia.org/w/index.php?title=Smilodecteshttp://en.wikipedia.org/w/index.php?title=Notharctushttp://en.wikipedia.org/w/index.php?title=Leptadapishttp://en.wikipedia.org/w/index.php?title=Palaeolemurhttp://en.wikipedia.org/w/index.php?title=Vertical_clinging_and_leapinghttp://en.wikipedia.org/w/index.php?title=Bamboo_lemurhttp://en.wikipedia.org/w/index.php?title=Quadrupedalismhttp://en.wikipedia.org/w/index.php?title=Vertical_clinging_and_leapinghttp://en.wikipedia.org/w/index.php?title=Galagohttp://en.wikipedia.org/w/index.php?title=Dwarf_lemurhttp://en.wikipedia.org/w/index.php?title=File:Galago_senegalensis.jpghttp://en.wikipedia.org/w/index.php?title=File:Fat-tailed_Dwarf_Lemur%2C_Kirindy%2C_Madagascar.jpghttp://en.wikipedia.org/w/index.php?title=Territory_%28animal%29http://en.wikipedia.org/w/index.php?title=Alarm_signalhttp://en.wikipedia.org/w/index.php?title=Urine_washinghttp://en.wikipedia.org/w/index.php?title=Feceshttp://en.wikipedia.org/w/index.php?title=Urinehttp://en.wikipedia.org/w/index.php?title=Epidermis_%28skin%29http://en.wikipedia.org/w/index.php?title=Scent_markinghttp://en.wikipedia.org/w/index.php?title=Social_groominghttp://en.wikipedia.org/w/index.php?title=Lemuridaehttp://en.wikipedia.org/w/index.php?title=Indriidaehttp://en.wikipedia.org/w/index.php?title=Multi-male_grouphttp://en.wikipedia.org/w/index.php?title=Multi-male_group -
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Strepsirrhini 14
Diet
Primates primarily feed on fruits (including seeds), leaves (including flowers), and animal prey (arthropods, small
vertebrates, and eggs). Diets vary markedly between strepsirrhine species. Like other leaf-eating (folivorous)
primates, some strepsirrhines can digest cellulose and hemicellulose.[140] Some strepsirrhines, such as the galagos,
slender lorises, and angwantibos are primarily insectivorous. Other species, such as fork-marked lemurs and
needle-clawed bushbabies, specialize on tree gum, while indriids, sportive lemurs, and bamboo lemurs are folivores.Many strepsirrhines are frugivores (fruit eaters), and others, like the ring-tailed lemur and mouse lemurs, are
omnivores, eating a mix of fruit, leaves, and animal matter.[141]
Among the adapiforms, frugivory seems to have been the most common diet, particularly for medium-sized to large
species, such as Cantius, Pelycodus and Cercamonius.[141] Folivory was also common among the medium and
large-sized adapiforms, including Smilodectes,Notharctus,Adapis andLeptadapis. Sharp cusps on the teeth of some
of the smaller adapiforms, such asAnchomomys andDonrussellia, indicate that they were either partly or primarily
insectivorous.[88][141]
Distribution and habitat
Before their extinction, adapiform primates were primarily found across North America, Asia, and Europe, with a
few species in Africa. They flourished during the Eocene when those regions were more tropical in nature, and they
disappeared when the climate became cooler and drier.[30] Today, the lemuriforms are confined in the tropics,[142]
ranging between 28 S to 26 N latitude.[10] Lorises are found both in equatorial Africa and Southeast Asia, while
the galagos are limited to the forests and woodlands of sub-Saharan Africa.[11][94] Lemurs are endemic to
Madagascar, although much of their diversity and habitat has been lost due to recent human activity.[11][44]
As with nearly all primates, strepsirrhines typically reside in tropical rainforests. These habitats allow strepsirrhines
and other primates to evolve diverse communities of sympatric species. In the eastern rainforests of Madagascar, as
many as 11 or 12 species share the same forests, and prior to human arrival, some forests had nearly double that
diversity.[142] Several species of lemur are found in drier, seasonal forests, including the spiny forest on the southerntip of the island, although the lemur communities in these regions are not as rich.[143]
Conservation
Strepsirrhines are threatened by deforestation in
tropical regions.
Like all other primates, strepsirrhines face an elevated risk of
extinction due to human activity, particularly deforestation in tropical
regions. Much of their habitat has been converted for human use, such
as agriculture and pasture.[16] The threats facing strepsirrhine primates
fall into three main categories: habitat destruction, hunting (for
bushmeat or traditional medicine), and live capture for export or localexotic pet trade. Although hunting is often prohibited, the laws
protecting them are rarely enforced. In Madagascar, local taboos
known as fady sometimes help protect lemur species, although some
are still hunted for traditional medicine.[144]
In 2012, the International Union for Conservation of Nature (IUCN) announced that lemurs were the "most
endangered mammals", due largely to elevated illegal logging and hunting following a political crisis in 2009. [] In
Southeast Asia, slow lorises are threatened by the exotic pet trade and traditional medicine, in addition to habitat
destruction.[145][146] Both lemurs and slow lorises are protected from commercial international trade under CITES
Appendix I.[]
http://en.wikipedia.org/w/index.php?title=CITEShttp://en.wikipedia.org/w/index.php?title=CITEShttp://en.wikipedia.org/w/index.php?title=Slow_lorishttp://en.wikipedia.org/w/index.php?title=2009_Malagasy_political_crisishttp://en.wikipedia.org/w/index.php?title=Illegal_logging_in_Madagascarhttp://en.wikipedia.org/w/index.php?title=International_Union_for_Conservation_of_Naturehttp://en.wikipedia.org/w/index.php?title=Fady_%28taboo%29http://en.wikipedia.org/w/index.php?title=Exotic_pethttp://en.wikipedia.org/w/index.php?title=Traditional_medicinehttp://en.wikipedia.org/w/index.php?title=Bushmeathttp://en.wikipedia.org/w/index.php?title=Habitat_destructionhttp://en.wikipedia.org/w/index.php?title=Deforestationhttp://en.wikipedia.org/w/index.php?title=File%3AKoh_Kong_logging.JPGhttp://en.wikipedia.org/w/index.php?title=Deforestationhttp://en.wikipedia.org/w/index.php?title=Madagascar_spiny_thicketshttp://en.wikipedia.org/w/index.php?title=Sympatryhttp://en.wikipedia.org/w/index.php?title=Tropical_rainforesthttp://en.wikipedia.org/w/index.php?title=Endemismhttp://en.wikipedia.org/w/index.php?title=Anchomomyshttp://en.wikipedia.org/w/index.php?title=Cusp_%28dentistry%29http://en.wikipedia.org/w/index.php?title=Cercamoniushttp://en.wikipedia.org/w/index.php?title=Pelycodushttp://en.wikipedia.org/w/index.php?title=Omnivorehttp://en.wikipedia.org/w/index.php?title=Frugivorehttp://en.wikipedia.org/w/index.php?title=Needle-clawed_bushbabyhttp://en.wikipedia.org/w/index.php?title=Fork-marked_lemurhttp://en.wikipedia.org/w/index.php?title=Insectivorehttp://en.wikipedia.org/w/index.php?title=Angwantibohttp://en.wikipedia.org/w/index.php?title=Slender_lorishttp://en.wikipedia.org/w/index.php?title=Hemicellulosehttp://en.wikipedia.org/w/index.php?title=Cellulosehttp://en.wikipedia.org/w/index.php?title=Folivorehttp://en.wikipedia.org/w/index.php?title=Arthropod -
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Notes
[1][1] Ankel-Simons 2007, p. 394.
[2] Vaughan, Ryan & Czaplewski 2011, p. 169.
[3][3] Osman Hill 1953, p. 39.
[4][4] Geoffroy Saint-Hilaire 1812, p. 156.
[5][5] Groves 2008, p. 166.
[6][6] Pocock 1918, p. 51.[7] Jenkins & Napier 1987, p. 1.
[8] Rose 2006, pp. 157 & 165167.
[9][9] Rose 2006, p. 166.
[10] Whitten & Brockman 2001, p. 322.
[11][11] Hartwig 2011, p. 29.
[12] Fichtel & Kappeler 2009, p. 397.
[13] Rose 2006, pp. 168169.
[14][14] Hartwig 2011, p. 22.
[15][15] Hartwig 2011, p. 24.
[16] Cartmill & Smith 2011, p. 84.
[17][17] Rose 2006, p. 169.
[18][18] Rose 2006, p. 165.
[19][19] Rose 2006, p. 343.
[20] Miller, Gunnell & Martin 2005, p. 67.
[21] Hartwig 2011, pp. 2829.
[22] Rose 2006, pp. 178179.
[23][23] Beard 2002, p. 133.
[24] Simons 2003, pp. 1517.
[25][25] Godinot 2006, p. 446.
[26][26] Rose 2006, p. 179.
[27] Covert 2002, pp. 1516.
[28][28] Gebo 2002, p. 21.
[29][29] Covert 2002, p. 14.
[30][30] Gebo 2002, p. 35.
[31][31] Rose 2006, p. 182.[32] Rose 2006, pp. 185186.
[33] Rose 2006, pp. 182185.
[34][34] Godinot 1998, p. 239.
[35] Gunnell, Rose & Rasmussen 2008, p. 257.
[36][36] Rose 2006, p. 186.
[37] Steiper & Seiffert 2012, p. 6006.
[38] Rose 2006, p. 182 & 186.
[39] Tabuce et al. 2009, pp. 40914092.
[40] Hartwig 2011, p. 2425.
[41] Vaughan, Ryan & Czaplewski 2011, pp. 170171.
[42][42] Rose 2006, p. 187.
[43][43] Rose 2006, p. 185.[44] Cartmill & Smith 2011, p. 89.
[45] Groves 2008, pp. 8990 & 96.
[46][46] Groves 2008, p. 103.
[47] Schwartz 2003, pp. 5354.
[48] Groves 2008, pp. 113114.
[49] Hartwig 2011, pp. 20 & 22.
[50][50] Sussman 2003, p. 45.
[51][51] Cartmill 2010, p. 15.
[52] Hartwig 2011, pp. 2021.
[53][53] Schwartz 2003, p. 54.
[54][54] Ankel-Simons 2007, p. 96.
[55] Since they are thought to be close relatives of tarsiers, omomyiforms are classified as haplorhines. However, the spacing of the roots of their
upper incisors suggests that they may have had a rhinarium, like the strepsirrhines.Rose 2006,
p. 182.
-
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Strepsirrhini 16
[56][56] Beard 1988, p. 92.
[57][57] Hartwig 2011, p. 28.
[58] Frequent mentions of a "strepsirrhine toothcomb" or references to Strepsirrhini as being "toothcombed primates" can be found in the
literature.Vaughan, Ryan & Czaplewski 2011, p. 169.
[59][59] Ankel-Simons 2007, p. 257.
[60][60] Schwartz 2003, p. 71.
[61][61] Ankel-Simons 2007, p. 32.
[62] Cartmill 2010, pp. 1617.[63] Rasmussen & Nekaris 1998, p. 252.
[64] Ankel-Simons 2007, pp. 3132.
[65][65] Groves 1998, p. 13.
[66][66] Rose 2006, p. 167.
[67] Beard 2002, pp. 146147.
[68][68] Gingerich 1975, p. 164.
[69] Yoder 2003, pp. 161162.
[70][70] Williams et al. 2010, p. 567.
[71][71] Franzen et al. 2009, p. e5723.
[72] Cartmill & Smith 2011, p. 90.
[73][73] Hartwig 2011, p. 20.
[74][74] Groves 2005, p. 121.[75] Godfrey & Jungers 2002, p. 106.
[76] Godfrey & Jungers 2002, p. 97.
[77][77] Rose 2006, p. 181.
[78][78] Covert 2002, p. 18.
[79] Dunkel, Zijlstra & Groves 2011/2012, p. 68.
[80] Mittermeier, Rylands & Konstant 1999, p. 4.
[81] Cuozzo & Yamashita 2006, p. 73.
[82][82] Osman Hill 1953, p. 96.
[83][83] Gebo 2002, p. 39.
[84] Rose 2006, pp. 166167.
[85][85] Tattersall 2006, p. 7.
[86][86] Ankel-Simons 2007, p. 458.
[87] Mittermeier, Rylands & Konstant 1999, p. 6.
[88][88] Rose 2006, p. 180.
[89][89] Rose 2006, p. 184.
[90] Godfrey & Jungers 2002, pp. 106 & 112.
[91][91] Ankel-Simons 2007, p. 429.
[92] Mittermeier, Rylands & Konstant 1999, pp. 45.
[93][93] Ankel-Simons 2007, p. 431.
[94] Cartmill & Smith 2011, p. 91.
[95] Dagosto 1988, pp. 4748.
[96][96] Dagosto 1988, p. 49.
[97][97] Ankel-Simons 2007, p. 68.
[98] Whitten & Brockman 2001, p. 323.
[99] Rose 2006, pp. 168 & 180.[100][100] Plavcan 2004, p. 239.
[101][101] Ankel-Simons 2007, p. 400.
[102] Ankel-Simons 2007, pp. 392394.
[103] Mittermeier, Rylands & Konstant 1999, p. 2425.
[104][104] Ankel-Simons 2007, p. 396.
[105][105] Beard 1988, p. 85.
[106] Ankel-Simons 2007, pp. 392393 & 400401.
[107] Ankel-Simons 2007, pp. 392393.
[108][108] Ankel-Simons 2007, p. 402.
[109][109] Ankel-Simons 2007, p. 410.
[110][110] Ankel-Simons 2007, p. 401.
[111] Ankel-Simons 2007, p. 410
411.
[112] Mittermeier, Rylands & Konstant 1999, p. 26.
[113] Rose 2006, pp. 181182.
-
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Strepsirrhini 17
[114] Whitten & Brockman 2001, pp. 325326.
[115] Whitten & Brockman 2001, pp. 330331.
[116] Cartmill & Smith 2011, p. 88.
[117][117] Osman Hill 1953, p. 81.
[118][118] Osman Hill 1953, p. 93.
[119][119] Osman Hill 1953, p. 391.
[120][120] Osman Hill 1953, p. 672.
[121][121] Ankel-Simons 2007, p. 523.[122][122] Ankel-Simons 2007, p. 522.
[123][123] Ankel-Simons 2007, p. 521.
[124] Mittermeier, Rylands & Konstant 1999, pp. 46.
[125] Mittermeier, Rylands & Konstant 1999, p. 5.
[126] Whitten & Brockman 2001, pp. 325 & 335.
[127] Fichtel & Kappeler 2009, pp. 395396.
[128] Fichtel & Kappeler 2009, pp. 401402.
[129] Nekaris & Bearder 2011, p. 51.
[130] Mittermeier, Rylands & Konstant 1999, pp. 2627.
[131] Gould, Sauther & Cameron 2011, p. 74.
[132] Fichtel & Kappeler 2009, pp. 395 & 397.
[133] Fichtel & Kappeler 2009, p. 398.[134] Mittermeier, Rylands & Konstant 1999, pp. 1516.
[135] Overdorff & Tecot 2006, p. 247.
[136] Cloninger & Kedia 2011, p. 86.
[137] Mittermeier, Rylands & Konstant 1999, pp. 2526.
[138] Mittermeier, Rylands & Konstant 1999, pp. 5 & 26.
[139] Covert 2002, pp. 1617.
[140] Mittermeier, Rylands & Konstant 1999, pp. 2224.
[141][141] Covert 2002, p. 16.
[142] Mittermeier, Rylands & Konstant 1999, p. 14.
[143] Mittermeier, Rylands & Konstant 1999, p. 19.
[144] Mittermeier, Rylands & Konstant 1999, pp. 3234.
[145] Fitch-Snyder & Livingstone 2008.
[146][146] Nekaris et al. 2010, p. 878.
References
Literature cited
Ankel-Simons, F. (2007).Primate Anatomy (3rd ed.). Academic Press. ISBN 978-0-12-372576-9.
Beard, K. C. (1988). "The phylogenetic significance of strepsirhinism in Paleogene primates" (http://link.
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