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the (134), and (93), placental (30), mammals (30), doi (27), pmid (23), from (22), order (22), #placentalia (17), with (17), bibcode (16), for (13), placentals (13), that (12), are (12), edit (11), evolution (11), cretaceous (11), which (11), mammal (10), boreoeutheria (10), afrotheria (10), eutheria (10), s2cid (10), primates (9), relatives (8), shrews (8), xenarthra (8), mammalia (8), other (8), wikipedia (7), articles (7), 2025 (7), different (7), extant (7), moles (7), orders (7), see (7), pmc (7), molecular (7), biology (7), journal (7), eutherian (7), these (7), page (6), all (6), paleocene (6), radiation (6), mammalian (6), basal (6), relationships (6), science (6), waddell (6), three (6), late (6), rodents (6), modern (6), mya (6), mirorder (6), may (5), statements (5), list (5), atlantogenata (5), 2022 (5), evolutionary (5), proceedings (5), society (5), phylogeny (5), wible (5), nature (5), phylogenetic (5), boundary (5), data (5), history (5), marsupials (5), more (5), large 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eduardo, brien, 1061, 12552136, 298725, 0334222100, 2003pnas, 1056s, 1056, tertiary, ota, 154, 11791233, 141, informatics, series, nishihara, maruyama, 5240, 19286970, 2655268, 0809297106, 2009pnas, 5235n, 5235, geological, suggest, superorders, averianov, ekdale, november, 6859, 11689942, 35102048, 2001natur, 62a, kriegs, jan, ole, churakov, gennady, kiefmann, jordan, ursula, brosius, schmitz, 2006, e91, 16515367, 1395351, pbio, 0040091, retroposed, elements, archives, towards, interordinal, 12078634, 1093, sysbio, haibing, jernvall, jukka, yilun, bian, congyu, bai, bin, qian, zhu, zifan, hai, lun, yuanqing, 2061, 1471, 2954, 1934, 292, unravelling, metatherian, velut, destrieux, kakou, 1998, morphologic, suppl, 9757322, neuro, chirurgie, marvalee, wake, september, 1992, university, chicago, press, 583, 978, 226, 87013, isbn, hyman, yuan, georgi, 822, 4330626, 11976675, 416816a, 816j, 816, mckenna, dashzeveg, horovitz, 1997, mongolia, 6650, 486, 205026882, 9333234, 39020, 1997natur, 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peared with waddell hasegawa and okada in 1999 18 jumping genes type retroposon presence absence patterns have provided corroboration of phylogenetic relationships inferred from molecular sequences 19 it is now widely accepted that there are three major subdivisions or lineages of placental mammals boreoeutheria xenarthra and afrotheria all of these diverged from common ancestors citation needed 2022 studies of bertrand o c and sarah l shelley have identified leptictids palaeoryctids and taeniodonts as basal placental mammal clades 2 4 leptictids were also proposed to be close relatives of boreoeutheria 3 the 19 living orders of placentalia in the three groups are 20 citation needed magnorder atlantogenata superorder xenarthra order cingulata armadillos order pilosa sloths and anteaters superorder afrotheria grandorder afroinsectiphilia order tubulidentata aardvarks mirorder afroinsectivora order afrosoricida tenrecs otter shrews and golden moles order macroscelidea elephant shrews grandorder paenungulata order hyracoidea hyraxes mirorder tethytheria order proboscidea elephants order sirenia dugongs and manatees magnorder boreoeutheria superorder euarchontoglires grandorder gliriformes mirorder glires order lagomorpha rabbits hares and pikas order rodentia rodents mice rats hamsters guinea pigs chinchillas capybaras porcupines voles squirrels beavers etc grandorder euarchonta order scandentia treeshrews mirorder primatomorpha order dermoptera colugos order primates primates monkeys apes including humans lemurs lorisids galagos and tarsiers superorder laurasiatheria order eulipotyphla hedgehogs gymnures shrews moles and solenodons order chiroptera bats grandorder ferungulata mirorder euungulata order artiodactyla even toed ungulates cattle antelopes sheep deer camelids pigs giraffes cetaceans hippopotamuses goats buffalo etc order perissodactyla odd toed ungulates horses asses zebras rhinoceroses and tapirs mirorder ferae order pholidota pangolins order carnivora carnivorans dogs cats bears pinnipeds mongooses raccoons skunks hyenas weasels etc the exact relationships among these three lineages is currently a subject of debate and four different hypotheses have been proposed with respect to which group is basal or diverged first from other placentals these hypotheses are atlantogenata basal boreoeutheria epitheria basal xenarthra exafroplacentalia basal afrotheria and a hypothesis supporting a near simultaneous divergence 21 estimates for the divergence times among these three placental groups mostly range from 105 to 120 million years ago mya depending on the type of dna whether it is translated and the phylogenetic method e g nuclear or mitochondrial 22 23 and varying interpretations of paleogeographic data 21 in addition a strict molecular clock does not hold so it is necessary to assume models of how evolutionary rates change along lineages these assumptions alone can make substantial differences to the relative ages of different mammal groups estimated with genomic data 24 placentalia atlantogenata xenarthra afrotheria boreoeutheria euarchontoglires glires euarchonta laurasiatheria eulipotyphla scrotifera chiroptera ferungulata ferae pholidota carnivora euungulata perissodactyla artiodactyla cladogram and classification based on amrine madsen h et al 2003 25 and asher r j et al 2009 26 compare with waddell hasegawa and okada 1999 18 and waddell et al 2001 22 genomics edit as of 2020 update the genome has been sequenced for at least one species in each extant placental order and in 83 of families 105 of 127 extant placental families 27 see list of sequenced animal genomes evolutionary history edit true placental mammals the crown group including all modern placentals arose from stem group members of the clade eutheria which had existed since at least the middle jurassic epoch about 170 mya these early eutherians were small nocturnal insect eaters with adaptations for life in trees 14 true placentals may have originated in the late cretaceous around 90 mya but the earliest undisputed fossils are dated to the cretaceous paleogene boundary k pg boundary the genus protungulatum is sometimes placed as a stem ungulate 28 with probably the earliest known species p coombsi from the strata within the hell creek formation specifically dated to at least 300 000 years before the k pg boundary 1 the genus purgatorius sometimes considered a stem primate appears no more than 300 000 years after the k pg boundary 29 one study has recovered both genera to be closely related and as stem eutherians outside modern placental mammals 30 but others have recovered protungulatum as a pan euungulate based on phylogenetic analysis and inner ear anatomy different from non placentals 31 32 the rapid appearance of placentals after the mass extinction at the end of the cretaceous suggests that the group had already originated and undergone an initial diversification in the late cretaceous as suggested by molecular clocks 33 the lineages leading to xenarthra and afrotheria probably originated around 90 mya and boreoeutheria underwent an initial diversification around 70 80 mya 33 producing the lineages that eventually would lead to modern primates rodents insectivores artiodactyls and carnivorans however modern members of the placental orders originated in the paleogene around 66 to 23 mya following the cretaceous paleogene extinction event the evolution of crown orders such modern primates rodents and carnivores appears to be part of an adaptive radiation 34 that took place as mammals quickly evolved to take advantage of ecological niches that were left open when most dinosaurs and other animals disappeared following the chicxulub asteroid impact as they occupied new niches mammals rapidly increased in body size and began to take over the large herbivore and large carnivore niches that had been left open by the decimation of the dinosaurs and perhaps more relevantly competing synapsids 35 mammals also exploited niches that the non avian dinosaurs had never touched for example bats evolved flight and echolocation allowing them to be highly effective nocturnal aerial insectivores and whales first occupied freshwater lakes and rivers and then moved into the oceans primates meanwhile acquired specialized grasping hands and feet which allowed them to grasp branches and large eyes with keener vision which allowed them to forage in the dark the evolution of land placentals followed different pathways on different continents since they cannot easily cross large bodies of water an exception is smaller placentals such as rodents and primates who left laurasia and colonized africa and then south america via rafting in africa the afrotheria underwent a major adaptive radiation which led to elephants elephant shrews tenrecs golden moles aardvarks and manatees 36 37 in south america a similar event occurred with radiation of the xenarthra which led to modern sloths anteaters and armadillos as well as the extinct ground sloths and glyptodonts expansion in laurasia was dominated by boreoeutheria which includes primates and rodents insectivores carnivores perissodactyls and artiodactyls these groups expanded beyond a single continent when land bridges formed linking africa to eurasia and south america to north america a study on eutherian diversity suggests that placental diversity was constrained during the paleocene while multituberculate mammals diversified afterwards multituberculates decline and placentals explode in diversity 35 notes edit genera alveugena ambilestes and procerberus are the immediate outgroups to order taeniodonta with genera alveugena and procerberus classified as a close sister taxa to this order exceptional adult placentals that retain a cloaca are golden moles tenrecs beavers and some shrews references edit 1 2 archibald j david zhang yue harper tony cifelli richard l may 6 2011 protungulatum confirmed cretaceous occurrence of an otherwise paleocene eutherian placental mammal pdf journal of mammalian evolution 18 3 153 161 doi 10 1007 s10914 011 9162 1 s2cid 16724836 archived from the original pdf on march 3 2016 retrieved april 28 2013 1 2 3 4 bertrand o c shelley s l williamson t e wible j r chester s g b flynn j j holbrook l t lyson t r meng j miller i m püschel h p smith t spaulding m tseng z j brusatte s l 2022 brawn before brains in placental mammals after the end cretaceous extinction science 376 6588 80 85 bibcode 2022sci 376 80b doi 10 1126 science abl5584 hdl 20 500 11820 d7fb8c6e 886e 4c1d 9977 0cd6406fda20 pmid 35357913 1 2 stephen brusatte robin m beck ornella c c bertrand fernando a perini hans püschel eleonora rossi sarah l shelley john r wible thomas williamson shawn p zack 2025 resolving early placental mamma relationships with morphological and molecular data in the society of vertebrate paleontology 85th annual meeting 1 2 sarah l shelley 2022 the phylogeny of paleocene mammals and the 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798 799 bibcode 2002natur 416 798w doi 10 1038 416798a pmid 11976661 s2cid 4332049 1 2 reilly s m white t d january 2003 hypaxial motor patterns and the function of epipubic bones in primitive mammals science 299 5605 400 402 bibcode 2003sci 299 400r doi 10 1126 science 1074905 pmid 12532019 s2cid 41470665 novacek m j rougier g w wible j r mckenna m c dashzeveg d and horovitz i october 1997 epipubic bones in eutherian mammals from the late cretaceous of mongolia nature 389 6650 483 486 bibcode 1997natur 389 483n doi 10 1038 39020 pmid 9333234 s2cid 205026882 cite journal cs1 maint multiple names authors list link 1 2 3 ji q luo z x yuan c x wible j r zhang j p and georgi j a april 2002 the earliest known eutherian mammal nature 416 6883 816 822 bibcode 2002natur 416 816j doi 10 1038 416816a pmid 11976675 s2cid 4330626 cite journal cs1 maint multiple names authors list link marvalee h wake 15 september 1992 hyman s comparative vertebrate anatomy university of chicago press p 583 isbn 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