strepsirrhini or strepsirhini ( / ˌstrɛpsəˈraɪni / ; strep-sə-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 .

collectively they are referred to as strepsirrhines .

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 does 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 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 from africa 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 .

strepsirrhini

* strepsirrhini on wikipedia . wikipedia * strepsirrhini on wikispecies . wikispecies * â strepsirrhini on wikimedia commons . wikimedia commons * strepsirrhini at the tree of life web project
github - laura - dietz / strepsirrhini : strepsirrhini , a modular composable toolkit in scala for retrieval , reranking , and expansion with and without entity annotations .
strepsirrhini ( strepsirrhines : primates with moist noses . ) ( also known as prosimians )
catalogue of primates in the british museum ( natural history ) . part iv : suborder strepsirrhini
tree of life web project . 1999 . strepsirrhini . version 01 january 1999 ( temporary ) .
strepsirrhine n . any member of the clade strepsirrhini , one of the two suborders of primates .
strepsi - tools : basic utilities for sequence and map processing , used by all other strepsirrhini components .
strepsirrhini , a modular composable toolkit in scala for retrieval , reranking , and expansion with and without entity annotations .
most species of prosimians are placed in the taxon strepsirrhini , which is one of the two suborders of primates .
grzegorz jagodziński added the polish common name
lemurowe
to
strepsirrhini é . geoffroy saint - hilaire , 1812
.
) and significantly ( p < 0 . 05 ) exceed the average genetic divergence across all of strepsirrhini ( nodes 176–178 ,
] . the mean date estimate for the basal primate split , strepsirrhini and haplorrhini at 63 . 7 mya ( additional file
this tree diagrams the branches of primates . notice that lemurs are found listed under strepsirrhini . within the order primates , members of strepsirrhini are closely related to new world monkeys , humans , great apes , gibbons , old world monkeys , tarsiers .
comparative morphological configuration of the cardiac nervous system in lorises and galagos ( infraorder lorisiformes , strepsirrhini , primates ) with evolutionary perspective .
phylogenetic relationship ) , lemurs and lorises are hence allocated to the suborder strepsirrhini , while tarsiers and simians are allocated to the suborder haplorhini .
) bayesian posterior mean estimates for the origins of crown strepsirrhini and crown lemuriformes differ by only 6 . 5 ma , whereas the estimates of roos
the living primates are classified in the order primates , which is commonly divided into two suborders : strepsirrhini and haplorhini . strepsirrhini means primates with moist noses and includes the lemuriform primates ( lemurs of madagascar , galagos and pottos from africa , and the lorises from india and southeast asia ) .
comparative morphological configuration of the cardiac nervous system in lorises and galagos ( infraorder lorisiformes , strepsirrhini , primates ) wit . . . - pubmed - ncbi
primates , linnaeus , 1758 ; strepsirrhini , geoffroy , 1812 ; lorisiformes , gregory , 1915 ; galagidae , gray , 1825 ; wadilemur , simons , 1997 .
we used the abr method to estimate the auditory sensitivities of 11 species ( n = 30 individuals ) in the suborder strepsirrhini ( electronic supplementary material , table s1 ) .
this tree diagrams the branches of strepsirrhini . notice that ring - tailed lemurs are found listed under lemuridae . within strepsirrhini , the members of lemuridae are closely related to a number of other families , including indridae ( avahi , sifakas , and indri ) , palaeopropithecidae ( giant sloth lemurs ) , cheirogaleidae ( dwarf lemurs and mouse lemurs ) , etc .
the suborder strepsirrhini is composed of seven families and includes four families of creatures typically called lemurs , and three families with the lorises , pottos , galagos , and aye - aye .
our results place the tarsiers as sister group to strepsirrhini , with a posterior probability of 1 . 0 . the divergence date estimate for the split between tarsiers and strepsirrhines at 58 . 6 mya is only slightly younger than the primate - euarchonta split ( 67 . 6 mya ) and the strepsirrhini - haplorrhini split ( 63 . 7 mya ) . we concur with eizirik
the suborder prosimii is now infrequently used in taxonomic classification . most species , with the exception of the tarsiers , formerly grouped in the suborder prosimii are now classified in the suborder strepsirrhini .
jenkins , p . d . , 1987 : catalogue of primates in the british museum ( natural history ) part iv : suborder strepsirrhini . british museum ( natural history ) , london .
l . catta is the only species of the genus lemur , but they share the family lemuridae with many others . their closest relatives are the gentle lemurs : the golden bamboo lemur , the grey gentle lemur and the broad - nosed gentle lemur . they are classified under the order of primates , and of the suborder strepsirrhini . strepsirrhini split from other primates approximately 63 million years ago .
strepsirrhini * a taxonomic suborder within the order primatesâ â the clade of primates other than the tarsiers and true simians . hyponyms * ( suborder ) : adapiformes ( extinct ) , chiromyiformes , lemuriformes , lorisiformes ( infraorders )
suborder : strepsirrhini infraorder : lorisiformes family : lorisidae genus : nycticebus species : n . bengalensis , n . coucang , n . javanicus , n . pygmaeus subspecies : n . c . coucang , n . c . menagensis
galagos are members of the taxon strepsirrhini , which is one of the two suborders of primates , and one that also includes the prosimians known commonly as lemurs , lorises , pottos , and aye - aye . members of strepsirrhini are known for their
wet noses ,
referring to the presence of the rhinarium , a wet , naked surface around the nostrils of the nose , such as found in most mammals , but lacking in the other primate suborder , the haplorrhini . the strepsirrhini ( non - tarsier prosimians ) also are called
curly - nosed
primates while the haplorrhini ( tarsiers , monkeys and apes ) are known as
simple - nosed
primates .
prosimian is an informal grouping of primates consisting of those belonging to the primates suborder strepsirrhini and the tarsiers of the suborder haplorrhini . at one point , all prosimians comprised the taxon prosimii , but this grouping was determined to be polyphyletic .
strepsirrhini species , height of the animal in the tree , direct distance from the observer to the animal , perpendicular distance from the animal to the transect line , the distance along transect and gps coordinates will be recorded for each sighting .
this file contains dental measurements of afradapis longicristatus gen . et sp . nov . , full results of phylogenetic analyses , morphological character support for nodes within strepsirrhini , the morphological character matrix , and the constraint tree used in the phylogenetic analysis .
of the order primates was reorganized so as to make taxa equivalent , as far as possible , to clades . in this classification , the order is divided into the suborders strepsirrhini and haplorrhini , and these in turn are divided into infraorders .
early classification schemes broke the primate order into the suborders prosimii ( prosimians ) and anthropoidea ( simians— monkeys and apes ) . however the prosimian tarsiers have been shown to be more closely related to the simians , and so tarseiers has been moved into the anthropoidea , which is now renamed as haplorrhini and prosimii renamed as strepsirrhini . thus , present day taxonomies have the primates divided into the suborder strepsirrhini , with all prosimians except the tarsiers , and suborder haplorrhini , with the tarsiers , monkeys , and apes .
of the primates has had to be revised . the two suborders recognized today are strepsirrhini ( lemurs and lorises ) and haplorrhini ( tarsiers , monkeys , and apes , including humans ) . the present classification was adopted in the early 1970s , when the logic of
comments : mckenna and bell ( 1997 ) divide the strepsirrhini into superfamilies daubentonioidea , lemuroidea , loroidea ( including cheirogaleidae ) and indroidea . evidence that cheirogaleidae is not related to lorises , and that indriidae is sister - group to lemuridae , is given by groves ( 2001c )
early classification schemes broke the primate order into the suborders prosimii ( wikipedia : prosimians ) and anthropoidea ( wikipedia : simians - monkeys and wikipedia : apes ) . however , the prosimian wikipedia : tarsiers have been shown to be more closely related to the wikipedia : simians , and so it has been joined with the simians . prosimii are now renamed strepsirrhini , and anthropoidea are now renamed haplorrhini . other classifications split strepsirrhini directly into four superfamilies : daubentonioidea , lemuroidea , loroidea ( including cheirogaleidae ) and indroidea . however , significant evidence suggests that cheirogaleidae is not related to the lorises , and that indridae is sister - group to lemuridae .
the suborder strepsirrhini is composed of seven families split into two groups . the first group is the infraorder wikipedia : lemuriformes , five families of primates typically called wikipedia : lemurs . the other two families are split with the wikipedia : lorises , wikipedia : pottos and the wikipedia : galagos in the infraorder wikipedia : lorisiformes .
prosimians include the lemurs , lorises , galagos , and the aye - aye of strepsirrhini and the tarsiers of haplorrhini . most prosimians are small , nocturnal , have a claw for grooming , and have a well - developed sense of smell . however , the members of the strepsirrhini are characterized by a moist nose and eyes with a tapetum lucidum ( shiny , reflective layer useful for night vision ) , whereas the tarsiers are part of the
dry - nosed
primates and lack a tapetum lucidum ( despite being nocturnal ) . all strepsirrhines except the aye - ayes also have a toothcomb—a comblike projection of their incisors and canine teeth that is used for grooming—while the tarsiers lack a toothcomb .
delpero , m . , j . c . masters , p . cervella , s . crovella , g . ardito and y . rumpler . 2001 . phylogenetic relationships among the malagasy lemuriforms ( primates : strepsirrhini ) as indicated by mitochondrial sequence data from the 12s rrna gene . zool . j . linnean soc . 132 : 83 - 103 .
lemur , ( suborder strepsirrhini ) , generally , any primitive primate except the tarsier ; more specifically , any of the indigenous primates of madagascar . in the broad sense , the term lemur applies not only to the typical lemurs ( family lemuridae ) but also to the avahis , sifakas , indri , and aye - aye of madagascar , in addition to the…
stereoscopic vision ( forward facing eyes ) , nails , post orbital bar , grasping big toe , body size tends to be larger , maturation / gestation / life span tends to be longer , tend to have fewer offspring , most are socially complex . derived ( most primates do not have a rhinarium ( haplorhini ) , primitive ( some primates have a rhinarium ( strepsirrhini )
, a trait characterized as a wet nose , generally present in mammals . in his catalog of features attributed to the strepsirrhini , he lists
les narines terminales et sinueuses
( translated
sinuous , twisted or curly nostrils
) . strepsirrhines are often characterized by their wet nose , but the etymology of their group name refers to the sinuous openings of the rhinarium ' s nostrils .
here we describe a rare and enigmatic primate species from bq - 2 that , unlike other primates from the eocene of africa , does not fit unambiguously into either anthropoidea or strepsirrhini on the basis of the limited dental evidence available . the absence of clear relatives on northern continents suggests that this genus might be a highly specialized member of an ancient and previously undocumented clade of endemic african primates .
a long continuous tarsiiformes branch ( node 142 ) , marked by 25 synapomorphic indels , is consistent with a relict lineage of ancient origin . the sequence phylogeny unambiguously supports tarsiers as a sister lineage ( albeit distant ) to simiiformes ( bs = 85 mp ; 98 ml ; 0 . 99 pp ) to form haplorrhini ( node 143 ) . a few indels ( table s5 ) define alternate evolutionary topologies , such as tarsiers aligned with strepsirrhini ( 1 indel , zfx ) or scandentia ( 1 indel , dctn2 ) , compared with those that support an ancestral grouping of tarsiiformes + strepsirrhini + dermoptera + scandentia ( 2 indels , plcb4 , pola1 ) . these incongruent alternatives suggest further investigation of more complex rare genomic changes as cladistic markers of ancient speciation is needed [ 17 ] , [ 18 ] .
as for morphological similarities between tarsiers and either one of the other two groups , resemblances to both can be found . again , the issue in deciding whether prosimians or anthropoids is more similar to tarsiers cannot be unequivocally disentangled . however , the dogma of the nose - based cladistic grouping ( strepsirrhini vs . haplorhini ) has become widely accepted among new world primatologists . therefore , the following statement is surprising :
within strepsirrhini , two common classifications include either two infraorders ( adapiformes and lemuriformes ) [ 15 ] or three infraorders ( adapiformes , lemuriformes , lorisiformes ) . [ 16 ] a less common taxonomy places the aye - aye ( daubentoniidae ) in its own infraorder , chiromyiformes . [ 17 ] 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 . [ 18 ] regardless of the infraordinal taxonomy , crown strepsirrhines are composed of 10 families , three of which are extinct . [ 15 ] these three extinct families included the giant lemurs of madagascar , [ 19 ] many of which died out within the last 1 , 000 years following human arrival on the island . [ 20 ]
in modern classifications , the order primates comprises two suborders : strepsirrhini ( wet - nosed primates ) and haplorrhini ( dry - nosed primates ) . this has not always been the case . one of the foremost debates in primate systematics has long concerned the position of tarsiers . traditionally viewed as being more closely associated with lemurs and lorises , tarsiers were placed within a suborder prosimii , under the gradistic view of primate taxonomy [
consider them to be a third suborder of the strepsirrhini because of their small size and galago - like behavior . others emphasize the fact that they lack the wet noses characteristic of lemurs and lorises . subsequently , they place them in the suborder haplorrhini along with monkeys , apes , and humans . the second of these two alternatives is used in this tutorial with the understanding that this issue has not yet been adequately resolved .
one of the most distinguishing characteristic of these primates is their
wet noses .
the name strepsirrhini is derived from greek meaning having a
curved or bent nose
( a terpsimbrotos compound of strepho or
bend
and rhis or
nose
) . the wet nose refers to the rhinarium , which is the wet , naked surface around the nostrils of the nose , such as found in most mammals . primates are phylogenetically divided into strepsirrhini , or
curly - nosed
primates with rhinariums , and haplorrhini , or
simple - nosed
primates that have a more mobile , continuous , dry upper lip instead of the rhinarium . mammals with rhinariums tend to have a stronger sense of olfaction , and the loss of the rhinarium in the haplorrhine primates is related to their decreased reliance on olfaction , being associated with other derived characteristics such as a reduced number of turbinates .
to provide information on evolutionary morphological differences and similarities between tarsiformes ( tarsiers ) and other primates ( strepsirrhini and platyrrhini ) , we examined the previously undescribed extrinsic cardiac nervous system ( ecns ) and surrounding structures in three philippine tarsiers ( tarsius syrichta ) . we found that the complicated ecns morphology shows little variation in contrast to significant anatomical diversity of the surrounding somatic structures in tarsiers as reported in our previous primate studies . further , our comparative morphological data on the ecns show the following aspects of evolutionary anatomical significance ; ( 1 ) the traits of the general relationship between the superior cervical ganglion and the spinal nerves are common to the tarsii and strepsirrhini , ( 2 ) the traits of the general composition of the cervicothoracic ganglion , its communicating branches to spinal nerves , and its cardiac nerve are common to the tarsii and platyrrhini , and ( 3 ) the traits of the general relationship between the middle cervical ganglion and the spinal nerves and its cardiac nerve are common among strepsirrhini , tarsii , and platyrrhini . therefore , the general anatomical characteristics of the ecns in tarsiers retain a primitive state for some traits and exhibit a derived state for others . these characteristics of the ecns also reflect an evolutionary history without the modification and specialization as seen in the surrounding somatic structures . it may play a key role in unifying future studies on molecular phylogeny and updating evolutionary anatomical information .
ring - tailed lemurs are prosimians . prosimians are a paraphyletic group that includes all strepsirrhini and tarsii , the most
ancestral
of the primates . prosimians once dominated the trees in many continents , but eventually were replaced by monkeys , who were stronger in the trees . they migrated to islands off the coast of africa and asia where the absence of monkeys allowed them to flourish into many species . in the wild , lemurs are only found on the island of madagascar .
when strepsirrhini is divided into two infraorders , the clade containing all toothcombed primates can be called
lemuriforms
. [ 15 ] when it is divided into three infraorders , the term
lemuriforms
refers only to madagascar ' s lemurs , [ 11 ] and the toothcombed primates are referred to as either
crown strepsirrhines
[ 24 ] or
extant strepsirrhines
. [ 25 ] confusion of this specific terminology with the general term
strepsirrhine
, along with oversimplified anatomical comparisons and vague phylogenetic inferences , can lead to misconceptions about primate phylogeny and misunderstandings about primates from the eocene , as seen with the media coverage of darwinius . [ 26 ] because the skeletons of adapiforms share strong similarities with those of lemurs and lorises , researchers have often referred to them as primitive strepsirrhines , [ 27 ] lemur ancestors , or a sister group to the living strepsirrhines . they are included in strepsirrhini , [ 25 ] and are considered basal members of the clade . [ 28 ] although their status as true primates is not questioned , the questionable relationship between adapiforms and other living and fossil primates leads to multiple classifications within strepsirrhini . often , adapiforms are placed in their own infraorder due to anatomical differences with lemuriforms and their unclear relationship . when shared traits with lemuriforms ( which may or may not be synapomorphic ) are emphasized , they are sometimes reduced to families within the infraorder lemuriformes ( or superfamily lemuroidea ) . [ 29 ]
the wikipedia : clade strepsirrhini ( wikipedia : new latin , from greek strepsis ,
a turning
and the stem of rhis ,
nose
) [ 1 ] is one of the two wikipedia : suborders of wikipedia : primates . wikipedia : madagascar ' s only primates ( apart from wikipedia : humans ) are strepsirrhines , and others can be found in southeast wikipedia : asia and wikipedia : africa . the scientist given credit for the name , wikipedia : étienne geoffroy saint - hilaire , named it for the retention of the wikipedia : rhinarium , a trait characterized as a wet nose , generally present in mammals . in his catalog of features attributed to the strepsirrhini , he lists
les narines terminales et sinueuses
( translated
sinuous , twisted or curley nostrils
) . [ 2 ] [ note 1 ] strepsirrhines are often characterized by their wet nose , but the etymology of their group name refers to the sinuous openings of the rhinarium ' s nostrils .
aided by samples of rare taxa , the phylogeny expands upon recent findings [ 19 ] - [ 21 ] to better resolve long - standing questions on the evolution of lorisiformes and the two endemic madagascar infraorders of chiromyiformes and lemuriformes . our data affirm the ancient split of strepsirrhini , approximately 68 . 7 mya ( node 144 ) , into the progenitors of lemuriformes / chiromyiformes ( origin 58 . 6 mya , node 174 ) and lorisiformes ( origin 40 . 3 mya , node 184 ) .
the strepsirrhini suborder contains the following infraorders : lemuriformes ( lemurs ) , chiromyiformes ( the aye - aye lemur ) , and lorisiformes ( galagos , lorises , and pottos ) . distinctive characteristics of strepsirrhines include : a
wet nose
( i . e . rhinarium ) , large olfactory lobes , a vomeral nasal organ for detecting pheromones , a reflective tapetum lucidum layer in the eye which provides better nocturnal vision , a postorbital bar bone that runs around the eye socket , and a tooth comb .
to estimate the structure and tempo of primate evolutionary history , we employed bayesian phylogenetic methods to analyze data supermatrices comprising 7 mitochondrial genes ( 6 , 138 nucleotides ) from 219 species across 67 genera and 3 nuclear genes ( 2 , 157 nucleotides ) from 26 genera . many taxa were only partially represented , with an average of 3 . 95 and 5 . 43 mitochondrial genes per species and per genus , respectively , and 2 . 23 nuclear genes per genus . our analyses of mitochondrial dna place tarsiiformes as the sister group of strepsirrhini . within haplorrhini , we find support for the primary divergence of pitheciidae in platyrrhini , and our results suggest a sister grouping of african and non - african colobines within colobinae and of cercopithecini and papionini within cercopthecinae . date estimates for nodes within each family and genus are presented , with estimates for key splits including : strepsirrhini - haplorrhini 64 million years ago ( mya ) , lemuriformes - lorisiformes 52 mya , platyrrhini - catarrhini 43 mya and cercopithecoidea - hominoidea 29 mya .
the relative placement of suborder strepsirrhini and infraorder tarsiiformes at an early stage of primate evolution has been difficult to resolve [ 8 ] - [ 11 ] . presently distributed in the islands of borneo , sumatra , sulawesi and the philippines , tarsiiformes had a broad holarctic distribution during the eocene [ 10 ] . phylogenetic placement of tarsiers has alternatively been defined as 1 ) sister taxa to strepsirrhini to form prosimii [ 2 ] , [ 8 ] , [ 12 ] , 2 ) allied with simiiformes ( anthropoidea ) to form haplorrhini [ 1 ] , [ 13 ] , [ 14 ] and 3 ) a separate relict lineage with an independent origin [ 15 ] . here we provide strong evidence that strepsirrhines split with suborder haplorrhini approximately 87 mya ( node 185 ) . the ancient lineage is monophyletic and defined by a long branch and eight shared insertions / deletions ( indels ) ( node 144 ) . rooted by lagomorpha , the phylogeny affirms dermoptera as the closest mammalian order relative to primates , followed by scandentia [ 16 ] , [ 17 ] .
) do not . in the meantime , various primates have been grouped together in modern cladistic arrangements for many reasons , but most obviously according to the presence or lack of a wet nose : the suborder strepsirrhini ( the lowly , wet - nosed prosimians ) versus the suborder haplorhini ( combining dry - nosed anthropoids and tarsiers ) . yet the two taxa combined in haplorhini are incongruent behaviorally , morphologically , and functionally . taxonomic issues become complicated when phylogenetic relationships , fossils , and time frames are added into this taxonomic equation .
concur that 67–69 primate genera originated from a common ancestor during the cretaceous / paleocene boundary roughly 80–90 mya . an eocene expansion formed the major extant lineages of 1 ) strepsirrhini , which is composed of lorisiformes ( galagos , pottos , lorises ) , chiromyiformes ( malagasy aye - aye ) and lemuriformes ( malagasy lemurs ) ; 2 ) tarsiiformes ( tarsiers ) and 3 ) simiiformes composed of platyrrhini ( new world monkeys ) and catarrhini , which includes cercopithecoidea ( old world monkeys ) and hominoidea ( human , great apes , gibbons ) ( see
in strepsirrhini , both members of the relaxin family appear to be encoded by a single gene . southern analysis of genomic dna from g . crassicaudatus with a 32 p - labeled galago - preprorelaxin probe revealed hybridization signals at 9 kb for hin diii - restricted genomic dna and 1 . 4 kb and 4 kb for eco ri - restricted genomic dna , whereas 32 p - labeled galago - rlf cdna probe revealed single hybridization signals at 13 kb ( hin diii digest ) and 2 kb ( eco ri digest ; data not shown ) .
an order of mammalia named by blumenbach in 1791 , including all kinds of apes , monkeys , and lemurs ; the quadrumanous mammals : so called because their hind as well as fore feet can be used as hands . the term is scarcely used now , being superseded by primates ; but primates includes both the bimana ( man alone ) and the quadrumana of the earlier systems . when the name was in vogue the quadrumana were usually divided into catarrhini , old world apes and monkeys ; platyrrhini , new world monkeys ; and strepsirrhini , lemurs .
in contrast to many molecular studies , but in support of others , our analysis has grouped tarsius with the strepsirrhini . whilst the majority of evidence supports a haplorrhine grouping for these taxa , the incongruence of some data demonstrates the uniqueness of these primates and the value of continued efforts to reassess phylogeny as new evidence and novel techniques become available . there are still significant gaps in our understanding of phylogenetic relationships within lorisiformes and platyrrhini ; crucially a range of molecular sequence data is required for various species within these groups before a convincing resolution can be reached .
lemuriformes is an infraorder of primate that falls under the suborder strepsirrhini . it includes the lemurs of madagascar , as well as the galagos and lorisids of africa and asia , although a popular alternative taxonomy places the lorisoids in their own infraorder , lorisiformes ( perelman et al . , 2011 ; roos et al . , 2004 ; stanger - hall , 1997 ) . lemuriform primates are characterized by a toothcomb , a specialized set of teeth in the front , lower part of the mouth that is mostly used for combing fur during grooming . [ 1 ]
) that numerous demonstrable lemuriform apomorphies would have appeared over the course of this interval seems to us to be quite low . more important , however , is the fact that there is no reason to believe that the origin of crown lemuriforms should have been coincident with their colonization of madagascar . it is , in fact , more likely that the dispersal to madagascar occurred earlier than the origin of crown lemuriforms , implying a duration of extra - malagasy stem lemuriform evolution that would be even shorter than the aforementioned 3 ma and 6 . 5 ma molecular estimates separating the origins of crown strepsirrhini and crown lemuriformes . indeed , given the absence of any other satisfying explanation for the lorisiform - lemuriform divergence , there is little reason to suspect that the origin of crown strepsirrhini would not have been essentially coincident with ( and , thus , due to ) the strepsirrhine dispersal to madagascar . such a scenario would imply that stem and crown lemuriforms never existed outside of madagascar , whereas stem strepsirrhines and stem and crown lorisiforms did . thus far , this is the pattern observable in the paleogene fossil record documenting afro - arabian strepsirrhine evolution .
those primates with a philtrum and wet rhinarium have therefore been classified together as strepsirrhini ( geoffroy , 1812 ) because of the structural similarity of these two characters in the nasal area . ( strepho means ' turned inward ' in greek and rhinos is greek for ' nose . ' ) in those primates whose snout is reduced in length and whose incisors are positioned close to each other , the rhinarium disappears . this is the case for tarsiers and anthropoids among primates , and therefore they were labeled together as haplorhini ( haplo means ' simple , single ' in greek ) .
those primates with a philtrum and wet rhinarium have therefore been classified together as strepsirrhini ( geoffroy , 1812 ) because of the structural similarity of these two characters in the nasal area . ( strepho means ' turned inward ' in greek and rhinos is greek for ' nose . ' ) in those primates whose snout is reduced in length and whose incisors are positioned close to each other , the rhinarium disappears . this is the case for tarsiers and anthropoids among primates , and therefore they were labeled together as haplorhini ( haplo means ' simple , single ' in greek ) .
placenta . all indications are that these two suborders separated very early , perhaps 60 million years ago . the strepsirrhini divide further into lemuriformes ( lemurlike ) and lorisiformes ( lorislike ) infraorders , while the haplorrhini divide into tarsiiformes ( tarsiers ) and simiiformes ( “anthropoids”—i . e . , monkeys , apes , and humans ) . the simiiformes divide in turn into platyrrhini ( new world monkeys ) and catarrhini ( old world monkeys and hominoids ) . a group of fossil mammals called the paromomyiformes , known mainly from the paleocene , have usually been classified as primates , but the eminent primate specialist
mitochondrial tree of strepsirrhine species . maximum - clade - credibility subtree of strepsirrhini , inferred from a species - level mitochondrial dna supermatrix using the bayesian phylogenetic software beast . nodes are labelled with a / b , where a represents the bayesian posterior probability expressed as a percentage and b represents the percentage of 1 , 000 maximum - likelihood bootstrap replicates that support the node . asterisks indicate 100 % support ; nodes with 100 % support in both bayesian and maximum - likelihood frameworks are labelled with single asterisks . the tree is drawn to a timescale , with node heights representing mean posterior estimates .
this study offers a novel reconstruction of phylogenetic relationships across the whole of the order primates down to species level and significantly builds upon previous whole - order phylogenies . it provides substantial support for previous studies in a number of key areas including : the primary divergence of lorisiformes within strepsirrhini , the primary divergence of pitheciidae in platyrrhini , a sister grouping of african and non - african colobines within colobinae and of cercopithecini and papionini within cercopthecinae . other advances include a better understanding of species - level relationships within the lemurs , macaques and gibbons , and estimates of divergence dates across the whole tree .
] , with strepsirrhini comprising lorisiformes ( galagos and lorises ) and the sister - pairing of the monophyletic lemuriformes ( lemurs ) and chiromyiformes ( the aye - aye ) , and with haplorrhini consisting of platyrrhini ( new world monkeys ) and catarrhini ( apes and old world monkeys ) . within these groups , however , there are numerous disagreements over interfamilial relationships . molecular evidence has sometimes favored cheirogaleidae as sister group to lemuridae , although current evidence suggests that the four lemuriform families ( lemuridae , cheirogaleidae , lepilemuridae and indriidae ) represent a four - way split , which may be real or may simply reflect a lack of resolution [
employing comparative analysis of the cdna - coding sequences of the unique preprorelaxin of the afro - lorisiform galago crassicaudatus and the malagasy lemur varecia variegata and the relaxin - like factor ( rlf ) of g . crassicaudatus , we demonstrated distinct differences in the dynamics of molecular remodeling of both hormones during primate evolution . the lorisiform and lemuriform preprorelaxin sequences encoded identical hormones , providing the first endocrinological evidence for the monophyletic origin of all strepsirrhini . structural analysis revealed the lemuriform members of the relaxin family to be potentially bioactive single - gene products . in contrast to the “two - prong” relaxin receptor - binding motif ( relvr ) present within the b - domains of other primate relaxins , strepsirrhine relaxin contained a unique “three - prong” motif ( rrlir ) with highest sequence homology to the receptor - binding motif of the evolutionarily much older skate relaxin . in contrast to relaxin , the rlf molecule was highly conserved during primate evolution and contained within its b - domain the putative relaxin receptor - binding motif and a pentameric sequence implicated in binding to specific rlf receptors . mutually exclusive expression of strepsirrhine preprorelaxin and rlf were observed in the fetal villous trophoblast cells of the strepsirrhine placenta and postpubertal testicular leydig cells , respectively , reflecting distinct functional roles for both hormones within the reproductive tract of strepsirrhini .
we estimated the phylogeny and divergence time splits simultaneously using a bayesian approach as implemented in the program beast ver 1 . 5 . 3 [ 76 ] , [ 77 ] . due to computational constraints , analyses were performed with 5 different sets of species : 1 ) genus - level data set including 61 primate genera , two dermoptera genera and one scandentia genus rooted by lagomorpha , 2 ) catarrhini species with outgroups , 3 ) platyrrhini species with outgroups , 4 ) strepsirrhini species with outgroups and 5 ) genus - level analysis with a partitioned data set allowing for rate heterogeneity and different substitution models for autosome , x - chromosome , and y - chromosome sequences .
the late eocene prosimian wadilemur elegans from the jebel qatrani formation , northern egypt , was originally interpreted as an anchomomyin adapiform primate based on limited information from the lower molars and distal premolars . recently recovered fossils attributable to this species , including a proximal femur , the fourth upper premolar and first and second upper molars , and a mandible preserving the lower second premolar and lower canine and incisor alveoli , reveal a number of derived morphological similarities shared with crown strepsirrhines and , in particular , miocene - to - recent stem and crown galagids , to the exclusion of known adapiforms . phylogenetic analysis of 359 morphological features scored across 95 living and extinct crown primate taxa supports a stem galagid placement for wadilemur and older saharagalago , and a close relationship between crown strepsirrhines and the eocene african taxa “ anchomomys ” milleri , djebelemur , and plesiopithecus ( none of which appear to be closely related to european anchomomyins ) . this scheme of relationships supports the hypothesis that crown strepsirrhini is of afro - arabian origin and that lemuriforms likely colonized madagascar by crossing the mozambique channel . wadilemur ' s known dental and postcranial morphology provides additional support for the hypothesis that galagids and lorisids had diverged by the close of the middle eocene , and , by bolstering the ≈37 million - year - old calibration point for crown lorisiform origins provided by saharagalago , indirect support for the hypothesis of an ancient origin of crown strepsirrhini and crown primates .
* ^ 2007 , ankel - simons , f . , primate anatomy , isbn 978 - 0 - 12 - 372576 - 9 : those primates with a philtrum and wet rhinarium have therefore been classified together as strepsirrhini ( geoffroy , 1812 ) because of the structural similarity of these two characters in the nasal area . ( _ strepho _ means ' turned inward ' in greek and _ rhinos _ is greek for ' nose . ' ) in those primates whose snout is reduced in length and whose incisors are positioned close to each other , the rhinarium disappears . this is the case for tarsiers and anthropoids among primates , and therefore they were labeled together as haplorhini ( _ haplo _ means ' simple , single ' in greek ) .
a division of quadrumana , contrasted with catarrhini and strepsirrhini , including all the american or new world members of the order primates and families cebidæ and hapalidæ or mididæ ; the platyrrhine monkeys . there is no bony external auditory meatus , the tympanic bone being annular ; the premolars are three above and below on each side ; the nasal septum is usually broad and fiat , and the nostrils are proportionately far apart , presenting forward or laterally and not downward ; the thumb , when present , is scarcely or not apposable ; there are no cheek - pouches nor ischial callosities ; and the tail is generally long and prehensile or bushy . also written platyrrhina , platyrrhinæ , and in all forms with single or double r . see cuts under cebinæ and hapalidæ .
until recently , naturalists divided primates into prosimians ( lemurs , lorises and tarsiers ) and simians ( monkeys , apes and human beings ) . today , though , the more widely accepted split is between
strepsirrhini
( wet - nosed ) and
haplorhini
( dry - nosed ) primates ; the former includes all the non - tarsier promisimians , and the latter consists of tarsiers and simians . simians themselves are divided into two major groups : old world monkeys and apes (
catarrhines ,
meaning
narrow - nosed
) and new world monkeys (
platyrhines ,
meaning
flat - nosed
) . technically , therefore , all human beings are haplorhine cattarrhines , dry - nosed , narrow - nosed primates . confused yet ?
the fact that the lower dental features of the stem galagid wadilemur are similar enough to afro - arabian stem strepsirrhines such as a . milleri that these taxa were initially placed together in the tribe anchomomyini suggests that the afro - arabian “anchomomyins” are probably of key importance for our understanding of crown strepsirrhine origins . in this regard , it is important that our phylogenetic analysis does not support the hypothesis that a . milleri is a close relative of european anchomomys ( represented in our analysis by a . gaillardi ) , and thus does not necessarily provide evidence for a trans - tethyan migration independent of other eocene afro - arabian non - adapid strepsirrhines ( 52 , 57 ) . given our identification of the eocene african taxa “ a . ” milleri , djebelemur , and plesiopithecus as successive sister taxa of crown strepsirrhini to the exclusion of holarctic adapiforms , it instead appears increasingly likely that these taxa are all derived from an ancient single colonization of afro - arabia that ultimately gave rise to crown strepsirrhini . the extent to which dental resemblances between european anchomomyins , “ a . ” milleri , djebelemur , omanodon , and various crown strepsirrhines are due to convergence , plesiomorphy , or both is unclear because all of these taxa remain so poorly known , but if djebelemur is in fact early eocene ( ypresian ) in age ( 58 ) , then its precocious development of the anchomomyin - like molar features that only appear much later ( mid - lutetian ) in the better sampled european record ( 54 , 59 ) seems , to us , to argue in favor of convergence .
morphological , molecular , and biogeographic data bearing on early primate evolution suggest that the clade containing extant ( or ‘crown’ ) strepsirrhine primates ( lemurs , lorises and galagos ) arose in afro - arabia during the early palaeogene 1 , but over a century of palaeontological exploration on that landmass has failed to uncover any conclusive support for that hypothesis 2 . here we describe the first demonstrable crown strepsirrhines from the afro - arabian palaeogene—a galagid and a possible lorisid from the late middle eocene of egypt , the latter of which provides the earliest fossil evidence for the distinctive strepsirrhine toothcomb . these discoveries approximately double the previous temporal range of undoubted lorisiforms and lend the first strong palaeontological support to the hypothesis of an ancient afro - arabian origin for crown strepsirrhini and an eocene divergence of extant lorisiform families 1 , 3 .
today , primates are split into two major suborders : strepsirrhini ( lemurs , galagos , lorises , and pottos ) and haplorhini ( tarsiers , new world monkeys , old world monkeys , and apes ) . in the past , this division was contentious . tarsiers proved particularly troublesome , as these animals exhibit a blend of anatomical features similar to both major groups . the now obsolete suborder prosimii (
prosimians
) grouped tarsiers with the species now classified as strepsirrhines while pooling the other primate species ( new world monkeys , old world monkeys , and apes ) under the suborder anthropoidea . however , this prosimii - anthropoidea split has since been disproved ; genetic testing has conclusively shown that tarsiers are more closely related to the
anthropoid
species . thus , prosimii is a polyphyletic suborder and is therefore obsolete .
on may 20 of last year , at a remarkable press conference in new york , a group of researchers announced—with much ballyhoo—that they’d found a 47 - million - year - old primate fossil named darwinius masillae ( nicknamed “ida” ) . ida , the finest fossil primate in existence , was touted loudly as the missing link between the two major branches of primates , the haplorhini ( anthropoids [ apes and monkeys ] and tarsiers ) , and the strepsirrhini ( lemurs and lorises ; see figure below ) . concurrent with the press conference was a history channel documentary and a book about ida , colin tudge’s the link , that proclaimed , with much heavy breathing , that ida was , as one of the earliest primate ancestors of our own species , an earthshaking discovery ( see my review of the book here ) .
the description of ida was eventually published in a paper by franzen et al . in the journal plos one , a journal that doesn’t exercise stringent scientific review of submitted papers . the reaction of both bloggers and scientists was very critical , with many pointing out that ida didn’t look like a missing link at all , but may have been only an adapiform primate , a representative of a lineage within the strepsirrhini that went extinct without leaving descendants . the phylogenetic analysis of the plos paper , they claimed , was cursory and incomplete . a later paper in nature by seiffert et al . cast further doubt on ida’s status , suggesting that the features that led ida’s discoverers to lump her with the haplorhines were convergent—that they had evolved independently in haplorhines and adapiforms and thus could not serve as evidence for ida’s anthropoid ancestry .
the succession in time and space of specific germ cell associations , denoted as spermatogenic stages , is a typical feature of mammalian spermatogenesis . the arrangement of these stages is either single stage ( one spermatogenic stage per tubular cross - section ) or multistage ( more than one spermatogenic stage per tubular cross - section ) . it has been proposed that the single - stage versus multistage arrangement is related to spermatogenic efficiency and that the multistage arrangement is typical for hominids . in the present work , the arrangement of spermatogenic stages and the spermatogenic efficiency of 17 primate species , comprising strepsirrhini ( prosimians : lemuriformes , lorisiformes ) , platyrrhini ( new world primates ) , catarrhini ( old world primates ) , and hominoidea ( great apes and humans ) , were analyzed comparatively by quantitative histological and flow cytometric means . we found a predominant single - stage tubular organization in the strepsirrhini , indicating that the single - stage form represents the ancestral state . the highest degree of multistage complexity was found in hominoidea ( except orangutan ) and in platyrrhini , but not in catarrhini . hence , no direct relationship between single - stage / multistage tubular topography and phylogeny could be established across primates . in fact , the tubule arrangement seen in platyrrhini and catarrhini primates is the reverse of what might be expected from phylogeny . interestingly , spermatogenic efficiency was similar in all species . we found no correlation between single - stage / multistage arrangement and spermatogenic efficiency or mating system . we speculate that the presence of a single - stage / multistage organization might simply reflect germ cell clonal size . our findings further indicate that sperm competition in primates is not reflected at the level of testicular function .
simons ' ( 19 ) diagnosis of w . elegans was based on information from two mandibles that together preserve information from p3 - m3 but no alveoli for more anterior teeth . similarities in lower molar morphology shared by wadilemur , the contemporaneous form anchomomys milleri ( also from l - 41 ) , older djebelemur from chambi , tunisia ( 51 ) , slightly younger omanodon and shizarodon from taqah , oman ( 52 ) , and anchomomys species from europe ( 53 ) , led simons to assign wadilemur to the alleged cercamoniine notharctid adapiform clade anchomomyini after the earlier tentative conclusions of gheerbrant et al . ( 52 ) regarding the affinities of omanodon and shizarodon . the alleged anchomomyins from the eocene of afro - arabia have had a convoluted taxonomic history since their initial descriptions : for instance , godinot ( 54 ) considered djebelemur , omanodon , and shizarodon to be either anthropoids or , less likely , crown strepsirrhines ( ”lemuriforms” in his usage ) ; shoshani et al . ( 55 ) placed omanodon and shizarodon within anthropoidea ; the phylogenetic analysis of marivaux et al . ( 56 ) found wadilemur to be an adapiform sister group of a clade containing european adapids and an omanodon - anchomomys - europolemur clade ; and , finally , the phylogenetic analysis of seiffert et al . ( 11 ) recovered an a . milleri - djebelemur clade that was found to be more closely related to plesiopithecus and crown strepsirrhini than either adapidae or notharctidae . the phylogenetic analysis presented herein , which constitutes the most comprehensive available assessment of relationships among early afro - arabian strepsirrhines , clearly supports a stem galagid placement for wadilemur and continues to support a close relationship between an a . milleri - djebelemur clade and crown strepsirrhini .
as recently as 2005 , the wikipedia : aye - aye was tentatively placed in its own infraorder ( chiromyiformes ) . it was uncertain whether this unique primate split off from the ancestral strepsirrhine line before the lemurs and lorises , or after . if the aye - aye represents a group that is ancestral to all the rest of strepsirrhini , then it evolved away from the strepsirrhine line between 63 million years ago ( mya ) ( when the strepsirrhines split from the primitive primate line ) and 50 mya ( the lemur / loris split ) . if chiromyiformes is to be considered as the sister only to the lemurs , then it must have evolved after the lemur / loris split 50 mya . in 2008 , the aye - aye and its family , daubentonia , were placed back in infraorder lemuriformes , making it the fifth lemur family . [ 4 ] molecular research had shown that aye - ayes were basal to the lemuriform clade . [ 5 ]
only two studies have previously involved extensive investigations on primate climatic niche evolution . the first was conducted by kamilar & muldoon [ 18 ] and involved the analysis of 43 malagasy primates . their results differed in many respects to those observed in our study of platyrrhines , particularly : ( 1 ) niche dimensions did not separate the relative effects of precipitation and temperature in niche space ; ( 2 ) no evidence of phylogenetic signal was detected in any pc . there are two main ( non - exclusive ) potential reasons for this discrepancy . the first is methodological : kamilar & muldoon [ 18 ] used individual records instead of species means to map interspecific variation in climatic niches . although that strategy allows for more precise information on the climatic conditions where each species is found , the considerable differences among species in the number of records ( 6 species account for more than half of the records ) might conflate intra - and interspecific variation in climatic niches . the second potential reason is biological : the age of the crown strepsirrhini is nearly twice that of the crown platyrrhini [ 25 ] , yet strepsirrhini evolved within a much smaller geographical area . the combination of more time for competitive interactions among lineages and opportunities for convergence within a smaller region might also have contributed to the observed difference from platyrrhini , but the available analyses cannot discriminate among these potential sources of discrepancy . the second study involved a comprehensive analysis of many ecological and life - history traits for all primates using blomberg et al . ' s k statistic [ 19 ] and suggested consistently weak phylogenetic signal for climatic niche variables . those results were interpreted as possibly advantageous , providing more room to evolve given possible constraints by other life - history traits with higher phylogenetic signal . our results suggest that the low phylogenetic signal in climatic niche variables might actually reflect large - scale differences among primate lineages , such that phylogenetic signal would be stronger within different clades .
adapiform or ' adapoid ' primates first appear in the fossil record in the earliest eocene epoch ( ∼55 million years ( myr ) ago ) , and were common components of palaeogene primate communities in europe , asia and north america 1 . adapiforms are commonly referred to as the ' lemur - like ' primates of the eocene epoch , and recent phylogenetic analyses have placed adapiforms as stem members of strepsirrhini 2 - 4 , a primate suborder whose crown clade includes lemurs , lorises and galagos . an alternative view is that adapiforms are stem anthropoids 5 . this debate has recently been rekindled by the description of a largely complete skeleton of the adapiform darwinius 6 , from the middle eocene of europe , which has been widely publicised as an important ' link ' in the early evolution of anthropoidea 7 . here we describe the complete dentition and jaw of a large - bodied adapiform ( afradapis gen . nov . ) from the earliest late eocene of egypt ( ∼37 myr ago ) that exhibits a striking series of derived dental and gnathic features that also occur in younger anthropoid primates - notably the earliest catarrhine ancestors of old world monkeys and apes . phylogenetic analysis of 360 morphological features scored across 117 living and extinct primates ( including all candidate stem anthropoids ) does not place adapiforms as haplorhines ( that is , members of a tarsiusanthropoidea clade ) or as stem anthropoids , but rather as sister taxa of crown strepsirrhini ; afradapis and darwinius are placed in a geographically widespread clade of caenopithecine adapiforms that left no known descendants . the specialized morphological features that these adapiforms share with anthropoids are therefore most parsimoniously interpreted as evolutionary convergences . as the largest non - anthropoid primate ever documented in afroarabia , afradapis nevertheless provides surprising new evidence for prosimian diversity in the eocene of africa , and raises the possibility that ecological competition between adapiforms and higher primates might have played an important role during the early evolution of stem and crown anthropoidea in afro - arabia .
the time and place of origin of the crown strepsirrhine or “toothcombed” prosimian clade , the group containing the malagasy lemurs , african galagos , and afro - asian lorises , has long been a matter of great interest to primatologists ( 1 - 4 ) . for many decades , the oldest undoubted fossil members of crown strepsirrhini were early miocene [ ≈20 megannum ( ma , megannum = million years old ) ] lorisiforms from kenya and uganda ( 5 - 10 ) , but the recent discovery of a ≈37 ma stem galagid ( saharagalago ) in northern egypt ( 11 ) has almost doubled this minimum paleontological estimate for the time of origin of both crown lorisiforms and crown strepsirrhines . independent estimates for the timing of the loris - galago divergence derived from dna sequence data ( 12 - 14 ) have since come to show remarkable congruence with this fossil evidence in supporting a middle eocene origin for crown lorisiforms , but this critical new calibration point within crown strepsirrhini also lends indirect support to the controversial hypothesis of a ≈80 ma origin for crown primates ( 13 , 15 , 16 ) that is inconsistent with current interpretations of the fossil record documenting early primate evolution ( e . g . , ref . 17 ) . given the pivotal role that saharagalago has come to play in debates surrounding the timing of crown lorisiform , lemuriform , strepsirrhine , and primate origins ( 13 , 14 , 18 ) , it is important that an eocene divergence of lorises and galagos continues to be tested with additional fossil evidence . here , we describe new fossils that indicate that wadilemur elegans , a ≈35 - 35 . 4 ma strepsirrhine primate from late eocene deposits of the jebel qatrani formation exposed in the fayum depression , northern egypt , is not an anchomomyin adapiform as originally thought ( 19 ) but is rather a primitive stem galagid lorisiform . these data strengthen support for the hypothesis of seiffert et al . ( 11 ) , yoder and yang ( 13 ) , and roos et al . ( 14 ) that lorises and galagos had already diverged by the close of the middle eocene .


sevenia benguelae is a butterfly in the nymphalidae family .

it is found in tanzania ( from the western part of the country to the kigoma district ) , the republic of the congo , the central and western part of the democratic republic of the congo , angola , western zambia and namibia .

the habitat consists of savanna woodland .

adults are attracted to fermenting fruit .

the larvae feed on maprounea africana .

sevenia benguelae

children sevenia benguelae , sevenia pechueli subsp . pechueli , sevenia trimeni subsp . trimeni , sevenia morantii , sevenia natalensis , sevenia rosa , sevenia boisduvali subsp . boisduvali
14 african and two madagascan species ( three according to larsen ( 2005 ) ) . ackery et al . ( 1995 ) state that sevenia ( as sallya ) has been viewed as a subgenus of the neotropical genus eunica by authors .
sevenia - species dictionary - southern africa : ispot nature - your place to share nature . ispot is a website aimed at helping anyone identify anything in nature . once you ' ve registered , you can add an observation to the website and suggest an identification yourself or see if anyone else can identify it for you .
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ackery pr , smith cr , and vane - wright ri eds . 1995 . carcasson ' s african butterflies . canberra : csiro .
larsen , t . b . 2005 butterflies of west africa . stenstrup , denmark : apollo books .
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species catalog enables to sort by characteristics such as expansion , flight time , etc . .