Mostrando postagens com marcador Purgatorius. Mostrar todas as postagens
Mostrando postagens com marcador Purgatorius. Mostrar todas as postagens

sexta-feira, 1 de janeiro de 2021

 

70 milhões de anos de evolução primata

A Evolução dos Primatas, do Purgatório ao Homo Sapiens

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Muitas pessoas têm uma visão compreensivelmente centrada no humano da evolução dos primatas , focalizando os hominídeos bípedes e de cérebro grande que povoaram as selvas da África alguns milhões de anos atrás. Mas o fato é que os primatas como um todo - uma categoria de mamíferos da megafauna que inclui não apenas humanos e hominídeos, mas macacos, macacos, lêmures, babuínos e társios - têm uma história evolutiva profunda que remonta à era dos dinossauros .

O primeiro mamífero que os paleontólogos identificaram como possuidores de características semelhantes às dos primatas foi o Purgatorius , uma minúscula criatura do tamanho de um camundongo do final do período Cretáceo (pouco antes do Evento de Impacto K / T que extinguiu os dinossauros). Embora parecesse mais com um musaranho-das-árvores do que com um macaco ou símio, o Purgatorius tinha uma dentição muito parecida com a de primata e (ou um parente próximo) pode ter gerado os primatas mais familiares da Era Cenozóica . (Estudos de sequenciamento genético sugerem que o ancestral primata mais antigo pode ter vivido espantosos 20 milhões de anos antes de Purgatório, mas ainda não há evidências fósseis para esta besta misteriosa.)

Os cientistas têm elogiado o Archicebus igualmente semelhante a um rato, que viveu 10 milhões de anos após o Purgatório, como o primeiro primata verdadeiro, e as evidências anatômicas em apoio a essa hipótese são ainda mais fortes. O que é confuso sobre isso é que o Arquicebu asiático parece ter vivido na mesma época que o Plesiadapis norte-americano e euro-asiático , um primata semelhante a um lêmure muito maior, com 60 centímetros de comprimento e com cabeça de roedor. Os dentes de Plesiadapis exibiram as primeiras adaptações necessárias para uma dieta onívora - uma característica chave que permitiu que seus descendentes, dezenas de milhões de anos depois, se diversificassem para longe das árvores e em direção aos campos abertos.

Evolução dos primatas durante a época do Eoceno

Durante a época do Eoceno - de cerca de 55 milhões a 35 milhões de anos atrás - pequenos primatas parecidos com lêmures assombraram florestas em todo o mundo, embora a evidência fóssil seja frustrantemente escassa. A mais importante dessas criaturas era Notharctus, que tinha uma mistura reveladora de traços símios: um rosto achatado com olhos voltados para a frente, mãos flexíveis que podiam agarrar galhos, uma espinha dorsal sinuosa e (talvez o mais importante) um cérebro maior, proporcional seu tamanho que pode ser visto em qualquer vertebrado anterior. Curiosamente, Notharctus foi o último primata a ser nativo da América do Norte; provavelmente descendeu de ancestrais que cruzaram a ponte de terra da Ásia no final do Paleoceno . Semelhante a Notharctus era o Darwinius da Europa Ocidental, o assunto de uma grande blitz de relações públicas alguns anos atrás, apregoando-o como o mais antigo ancestral humano; poucos especialistas estão convencidos.

Outro importante primata do Eoceno foi o asiático Eosimias ("macaco da madrugada"), que era consideravelmente menor que Notharctus e Darwinius, apenas alguns centímetros da cabeça à cauda e pesando um ou dois onças, no máximo. O Eosimias noturno, que habita as árvores - que era aproximadamente do tamanho de um mamífero mesozóico médio - foi postulado por alguns especialistas como prova de que os macacos se originaram na Ásia e não na África, embora isso esteja longe de ser uma conclusão amplamente aceita. O Eoceno também testemunhou os Smilodectes norte-americanos e o divertido Necrolemur da Europa Ocidental, ancestrais dos macacos pequeninos que eram remotamente aparentados com os lêmures e társios modernos.

Uma breve digressão: Os lêmures de Madagascar

Por falar em lêmures, nenhum relato da evolução dos primatas estaria completo sem uma descrição da rica variedade de lêmures pré-históricos que uma vez habitaram a ilha de Madagascar, no Oceano Índico, na costa leste da África. A quarta maior ilha do mundo, depois da Groenlândia, Nova Guiné e Bornéu, Madagascar se separou do continente africano há cerca de 160 milhões de anos, durante o final do período Jurássico , e depois do subcontinente indiano em qualquer lugar de 100 a 80 milhões de anos atrás, durante o período médio ao final do Cretáceo. O que isso significa, é claro, é que é virtualmente impossível para qualquer primata mesozóico ter evoluído em Madagascar antes dessas grandes divisões - então, de onde vieram todos esses lêmures?

A resposta, pelo que os paleontólogos podem dizer, é que alguns primatas do Paleoceno ou Eoceno sortudos conseguiram flutuar da costa africana para Madagascar em colmo emaranhado de madeira flutuante, uma jornada de 320 quilômetros que poderia ter sido realizada em questão de dias. Crucialmente, os únicos primatas que fizeram essa viagem com sucesso foram os lêmures, e não outros tipos de macacos - e uma vez instalados em sua enorme ilha, esses minúsculos progenitores ficaram livres para evoluir em uma ampla variedade de nichos ecológicos durante as dezenas de milhões de anos (ainda hoje, o único lugar na Terra onde você pode encontrar lêmures é Madagascar; esses primatas morreram há milhões de anos na América do Norte, na Eurásia e até na África).

Dado seu relativo isolamento e a falta de predadores eficazes, os lêmures pré-históricos de Madagascar estavam livres para evoluir em algumas direções estranhas. A época do Pleistoceno testemunhou lêmures gigantes como Archaeoindris, que era do tamanho de um gorila moderno, e o menor Megaladapis, que "apenas" pesava 45 quilos ou mais. Totalmente diferentes (mas, claro, intimamente relacionados) eram os chamados lêmures "preguiças", primatas como Babakotia e Palaeopropithecus que pareciam e se comportavam como preguiças, subindo preguiçosamente em árvores e dormindo de cabeça para baixo nos galhos. Infelizmente, a maioria desses lêmures lentos, confiantes e estúpidos foram condenados à extinção quando os primeiros colonos humanos chegaram a Madagascar, cerca de 2.000 anos atrás.

Macacos do Velho Mundo, Macacos do Novo Mundo e os primeiros macacos

Freqüentemente usada de forma intercambiável com "primata" e "macaco", a palavra "símio" deriva de Simiiformes, a infraordem de mamíferos que inclui macacos e macacos do velho mundo (ou seja, africano e eurasiano) e do novo mundo (ou seja, central e sul-americana ) macacos; os pequenos primatas e lêmures descritos na página 1 deste artigo são geralmente chamados de "prosímios". Se tudo isso parece confuso, o importante a lembrar é que os macacos do novo mundo se separaram do ramo principal da evolução símia há cerca de 40 milhões de anos, durante a época do Eoceno, enquanto a divisão entre macacos do velho mundo e macacos ocorreu há cerca de 25 milhões de anos mais tarde.

A evidência fóssil de macacos do novo mundo é surpreendentemente pequena; até o momento, o gênero mais antigo já identificado é Branisella, que viveu na América do Sul entre 30 e 25 milhões de anos atrás. Normalmente para um macaco do novo mundo, Branisella era relativamente pequena, com um nariz achatado e uma cauda preênsil (estranhamente, os macacos do velho mundo nunca conseguiram desenvolver esses apêndices ágeis e flexíveis). Como Branisella e seus companheiros macacos do novo mundo fizeram todo o caminho da África à América do Sul? Bem, o trecho do Oceano Atlântico que separa esses dois continentes era cerca de um terço mais curto 40 milhões de anos atrás do que é hoje, então é concebível que alguns pequenos macacos do velho mundo tenham feito a viagem acidentalmente, em colmo flutuante de troncos.

Justa ou injustamente, os macacos do velho mundo costumam ser considerados significativos apenas na medida em que eventualmente geraram macacos, e depois hominídeos, e depois humanos. Um bom candidato para uma forma intermediária entre macacos do velho mundo e macacos do velho mundo era Mesopithecus, um primata parecido com o macaco que, como os macacos, buscava folhas e frutos durante o dia. Outra forma de transição possível era Oreopithecus (chamado de "monstro do biscoito" pelos paleontólogos), um primata europeu residente na ilha que possuía uma estranha mistura de características de macaco e macaco, mas (de acordo com a maioria dos esquemas de classificação) quase não era um verdadeiro hominídeo.

A evolução de macacos e hominídeos durante a época do Mioceno

É aqui que a história fica um pouco confusa. Durante a época do Mioceno , de 23 a 5 milhões de anos atrás, uma variedade desconcertante de macacos e hominídeos habitou as selvas da África e da Eurásia (os macacos se distinguem dos macacos principalmente por sua falta de cauda e braços e ombros mais fortes, e os hominídeos são diferentes dos macacos principalmente por suas posturas eretas e cérebros maiores). O macaco africano não hominídeo mais importante foi o Pliopithecus , que pode ter sido ancestral dos gibões modernos; um primata ainda mais antigo, o Propliopithecus , parece ter sido ancestral do Pliopithecus. Como seu status não hominídeo indica, Pliopithecus e macacos relacionados (como o Proconsul) não eram diretamente ancestrais dos humanos; por exemplo, nenhum desses primatas andava sobre dois pés.

A evolução dos macacos (mas não dos hominídeos) realmente atingiu seu ritmo durante o Mioceno posterior, com o Dryopithecus , que vive em árvores , o enorme Gigantopithecus (que tinha cerca de duas vezes o tamanho de um gorila moderno) e o ágil Sivapithecus , que agora é considerado o mesmo gênero de Ramapithecus (descobriu-se que fósseis menores de Ramapithecus eram provavelmente fêmeas de Sivapithecus!) Sivapithecus é especialmente importante porque este foi um dos primeiros macacos a se aventurar a descer das árvores para as pastagens africanas, uma transição evolutiva crucial que pode foram estimulados pelas mudanças climáticas .

Os paleontólogos discordam dos detalhes, mas o primeiro hominídeo verdadeiro parece ter sido o Ardipithecus, que caminhava (embora desajeitadamente e ocasionalmente) sobre dois pés, mas tinha apenas um cérebro do tamanho de um chimpanzé; Ainda mais assustadoramente, não parece ter havido muita diferenciação sexual entre os machos e as fêmeas de Ardipithecus, o que torna este gênero assustadoramente semelhante aos humanos. Alguns milhões de anos depois do Ardipithecus vieram os primeiros hominídeos indiscutíveis: Australopithecus (representado pelo famoso fóssil "Lucy"), que tinha apenas cerca de quatro ou cinco pés de altura, mas andava sobre duas pernas e tinha um cérebro invulgarmente grande, e Paranthropus, que já foi considerado uma espécie de Australopithecus, mas desde então ganhou seu próprio gênero graças ao seu invulgarmente grande e musculoso cabeça e cérebro correspondentemente maior.

Tanto o Australopithecus quanto o Paranthropus viveram na África até o início da época do Pleistoceno; os paleontólogos acreditam que uma população de Australopithecus foi o progenitor imediato do gênero Homo, a linha que eventualmente evoluiu (no final do Pleistoceno) em nossa própria espécie, Homo sapiens .

 

 

quarta-feira, 9 de setembro de 2020

B. Chester, Jonathan I. Bloch, Doug M. Boyer, and William A. Clemens
  1. Edited by Neil H. Shubin, The University of Chicago, Chicago, IL, and approved December 24, 2014 (received for review November 12, 2014)

Significance

Purgatorius has been considered a plausible ancestor for primates since it was discovered, but this fossil mammal has been known only from teeth and jaw fragments. We attribute to Purgatorius the first (to our knowledge) nondental remains (ankle bones) which were discovered in the same ∼65-million-year-old deposits as dentitions of this putative primate. This attribution is based mainly on size and unique anatomical specializations known among living euarchontan mammals (primates, treeshrews, colugos) and fossil plesiadapiforms. Results of phylogenetic analyses that incorporate new data from these fossils support Purgatorius as the geologically oldest known primate. These recently discovered tarsals have specialized features for mobility and provide the oldest fossil evidence that suggests arboreality played a key role in earliest primate evolution.

Abstract

Earliest Paleocene Purgatorius often is regarded as the geologically oldest primate, but it has been known only from fossilized dentitions since it was first described half a century ago. The dentition of Purgatorius is more primitive than those of all known living and fossil primates, leading some researchers to suggest that it lies near the ancestry of all other primates; however, others have questioned its affinities to primates or even to placental mammals. Here we report the first (to our knowledge) nondental remains (tarsal bones) attributed to Purgatorius from the same earliest Paleocene deposits that have yielded numerous fossil dentitions of this poorly known mammal. Three independent phylogenetic analyses that incorporate new data from these fossils support primate affinities of Purgatorius among euarchontan mammals (primates, treeshrews, and colugos). Astragali and calcanei attributed to Purgatorius indicate a mobile ankle typical of arboreal euarchontan mammals generally and of Paleocene and Eocene plesiadapiforms specifically and provide the earliest fossil evidence of arboreality in primates and other euarchontan mammals. Postcranial specializations for arboreality in the earliest primates likely played a key role in the evolutionary success of this mammalian radiation in the Paleocene.
Evidence from the fossil record suggests that placental mammals diversified following the Cretaceous–Paleogene (K–Pg) boundary ∼66 Mya (1, 2). Among the oldest known placental mammals, the putative primate Purgatorius has been documented in the western interior of North America during the first million years after the K–Pg boundary (25) to within the first few hundred thousand years of the Paleocene (6). Although the fossil record of Purgatorius has been restricted to dentitions long recognized as uniquely similar to those of primates (3, 4), these anatomical data are limited. Some researchers who have preferred to restrict the order Primates to the crown-clade (i.e., Euprimates) have also questioned the primate affinities of Purgatorius and other Paleogene plesiadapiforms (reputed stem primates) (e.g., refs 7 and 8). Furthermore, several recent phylogenetic analyses have not supported Purgatorius in Primates (9) or even in Placentalia (crown-clade eutherians) (1012). New evidence supporting Purgatorius as the oldest plesiadapiform primate is derived from tarsal bones collected at the late Puercan (Pu3; ∼65 Mya) Garbani Channel fauna localities in Garfield County, northeastern Montana. Four decades of fieldwork have resulted in the recovery of hundreds of Purgatorius teeth and fragmentary jaws (13). Here we attribute tarsals (astragali and calcanei) to Purgatorius based on size and diagnostic euarchontan and plesiadapiform features (Fig. 1 and SI Appendix). This taxonomic attribution is supported further by the absence of other euarchontan taxa from the Garbani Channel fauna.

Phylogenetic Analysis

Results from recent broad cladistic analyses that focused on relationships among eutherian mammals do not support primate affinities of Purgatorius and instead place Purgatorius in a clade directly outside Placentalia with the contemporary condylarths (archaic ungulates) Protungulatum and Oxyprimus (1012). However, the addition of new tarsal data for Purgatorius and increased taxon sampling, including a colugo and four plesiadapiforms, using this same matrix, results in a strict consensus tree that supports a monophyletic Euarchonta with Sundatheria (treeshrews and colugos) as the sister group to a fairly unresolved Primates clade that includes Purgatorius (Fig. 2A). This result is driven mainly by the addition of euarchontan taxa rather than by new character data for Purgatorius, which strongly suggests that the previous support for Purgatorius outside Placentalia and Euarchonta is primarily an artifact of taxon sampling (10, 14). To address this issue further, we included the new Purgatorius tarsal data in two additional analyses that were designed to evaluate relationships within Euarchonta (15) or more broadly within Euarchontoglires (16). Results from both analyses support Purgatorius as the most basal primate (Fig. 2 B and C).
Fig. 2.
Hypotheses of evolutionary relationships of Purgatorius and other eutherian mammals. (A) Simplified resulting strict consensus cladogram based on data modified from ref. 12, Purgatorius tarsals, and five additional euarchontan taxa (colugo Cynocephalus, the micromomyid plesiadapiforms Foxomomys, Dryomomys, and Tinimomys, and the carpolestid plesiadapiform Carpolestes). Asterisks indicate results from a previously published analysis with Laurasiatheria excluding Eulipotyphla and Afrotheria excluding Afrosoricida (12). (B) Simplified resulting single-most-parsimonious cladogram based on data from ref. 15 and Purgatorius tarsals. (C) Simplified resulting strict consensus cladogram based on data from ref. 16 and Purgatorius tarsals. In all cladograms, Sundatheria is supported and indicated in green, Primates is supported and indicated in violet, and Purgatorius is supported as a primate and indicated in orange. See SI Appendix for methods, full tree topologies, and support values.

Description and Comparison of Tarsal Bones

Although previously published cladistic analyses support a close relationship between Purgatorius and the condylarth Protungulatum (1012), the tarsals attributed to Purgatorius differ considerably from those of Protungulatum by having many characteristics of euarchontan mammals that relate to arboreality (Fig. 3). As in other euarchontans, the upper ankle joint of Purgatorius is more mobile than that of Protungulatum, which has a contact between the fibula and calcaneus that restricts medial–lateral movements at this joint (17). The astragalar trochlea (lateral tibial facet) of Purgatorius is relatively longer than that of Protungulatum, allowing a greater range of dorsi- and plantarflexion (Fig. 3). The trochlea of Purgatorius is medially sloping, is aligned oblique to the long axis of the astragalus, and extends slightly onto the dorsal surface of the astragalar neck, as is consistent with mammals whose feet abduct during dorsiflexion for climbing on vertical supports (18). The lower ankle joint of Purgatorius also is considerably more mobile than that of Protungulatum, especially in having increased capacity for movements between the sustentacular facets of the astragalus and calcaneus. Purgatorius has a saddle-shaped astragalar ectal facet that articulates with and rotates along a longer, moderately proximodistally aligned calcaneal ectal facet (Fig. 1). This morphology suggests a pronounced capacity for inversion and eversion of the foot, which is supported further by the presence of a well-developed distal calcaneal sustentacular facet and a distally extensive astragalar sustentacular facet that contacts the navicular facet (Fig. 3). These distal articular regions would have come into close contact only during strong inversion of the foot. Such movements are facilitated further at the transverse tarsal joint of Purgatorius by the rounded, concave, gliding articulation of the calcaneocuboid facet and its fairly transverse orientation and by the pronounced, rounded navicular facet on the medial side of the astragalar head (Fig. 1). In contrast, Protungulatum has a more ovoid, asymmetrical calcaneocuboid facet that is oriented more obliquely to the long axis of the calcaneus and a less pronounced medial side of the astragalar head, suggesting that Protungulatum had less capacity for pedal inversion and used level-oriented foot positions for locomotion on a flat substrate, as do terrestrial quadrupeds (Fig. 3) (17).
Fig. 3.
Comparison of micro-CT scan images of tarsal bones. Columns illustrate tarsals of condylarth Protungulatum (A), colugo Cynocephalus (B), treeshrew Ptilocercus (C), Purgatorius (D), the micromomyid plesiadapiform Dryomomys (E), and the adapoid euprimate Notharctus (F) right astragali (rows 1–3) and calcanei (rows 4–6) in dorsal (rows 1 and 4), plantar (rows 2 and 5), and distal (rows 3 and 6) views, respectively. Some elements are reversed for clarity. (Scale bars: 1 mm.) aef, astragalar ectal facet; aff, astragalar groove for tendon of musculus flexor fibularis (violet); ah, astragalar head (blue); asf, astragalar sustentacular facet (red); caf, calcaneal fibular facet; cef, calcaneal ectal facet; cf, calcaneocuboid facet (orange); cff, calcaneal groove for tendon of musculus flexor fibularis; csf, calcaneal sustentacular facet (red); ltf, lateral tibial facet (yellow); nav, astragalonavicular facet (red); pt, peroneal tubercle. See SI Appendix for specimen numbers.
Among euarchontans, the tarsals attributed to Purgatorius are uniquely similar to those of other plesiadapiforms in having an astragalus with a medially sloping trochlea and a relatively broad head and a calcaneus with a large peroneal tubercle (Figs. 3 and 4 and SI Appendix) (18). However, it should be noted that currently euarchontan tarsal comparisons outside Primates are limited to the presumably more derived morphologies of extant colugos and treeshrews, given the paucity of postcranial fossils representing these clades. Unlike the level astragalar trochlea of colugos and the most basally divergent treeshrew Ptilocercus, the medially sloping trochlea of Purgatorius may have reduced the potential for lateral sheer of the tibia on the astragalus when inverted foot postures were used during locomotion on large-diameter supports (Fig. 3). The astragalar head of Purgatorius and other plesiadapiforms is broad and ovoid, suggesting frequent use of inverted and everted postures. The large medial aspect of the astragalonavicular facet of Purgatorius likely reflects forces frequently transmitted on the medial side of the head during habitual pedal inversion (17), but the more spherical head of colugos, Ptilocercus, and many euprimates indicates even greater emphasis on inverted postures in these taxa (Fig. 3). Purgatorius also differs from colugos, Ptilocercus, and euprimates in having a calcaneus with a much larger and more laterally projecting peroneal tubercle (Fig. 3), which provides more leverage for tendons of peroneal muscles that contribute to eversion (musculus peroneus longus) and abduction (musculus peroneus brevis) and counterbalance forces that invert the foot (19). The smaller peroneal tubercle in other euarchontans suggests less emphasis on the peroneal muscles for eversion movements and rotational stability, possibly as a mechanical consequence of the greater degree of distal calcaneal elongation present in these taxa (18, 20).
Fig. 4.
Results of principal component analysis of 23 astragalar measurements for 34 species (A) and 25 calcaneal measurements for 33 species (B) (SI Appendix). Lines connecting data points reflect a minimum-spanning tree computed from a Euclidean distance matrix. Polygons encompass taxa including living strepsirrhine and haplorhine euprimates (red), treeshrews (blue), and colugos (yellow). Gray polygons encompass fossil groups including adapoid and omomyoid euprimates, plesiadapiforms, and earliest Paleocene mammals Protungulatum donnae (Pd) and Procerberus formicarum (Pf). Results support tarsals attributed to Purgatorius (starred) as a plesiadapiform, which, like Protungulatum and Cretaceous Deccanolestes (D. hislopi, Dh; D. robustus, Dr), has tarsal features that may be plesiomorphic, such as a large calcaneal peroneal tubercle. See SI Appendix for eigenvalue, percentage variance, and variable component loadings for each principal component.
Micromomyids are the most primitive plesiadapiforms known from skeletons (Fig. 2 B and C) and have been reconstructed as being most similar to Ptilocercus among extant mammals (15). Thus it is significant that Purgatorius, which has teeth very similar to those of primitive micromomyids (21, 22), also shares with that group unique tarsal features including a slightly grooved astragalar trochlea with a relatively high medial ridge and a fairly consistent mediolateral width (Fig. 3), whereas other stem primates have a flat and more medially sloping trochlea that is widest distally. Purgatorius and micromomyids also are most similar in tarsal features related to the tendon of musculus flexor (digitorum) fibularis, which contributes to digital flexion and plantarflexion of the foot and is important for pedal grasping. These taxa have a very large and mediolaterally wide flexor fibularis groove on the astragalus (Fig. 3), as is consistent with the large origination areas indicating sizeable flexor muscles on the tibia and fibula of micromomyids (23). However, the corresponding groove for the tendon of flexor fibularis on the plantar aspect of the calcaneal sustentaculum is shallow in Purgatorius and micromomyids, as it is in treeshrews and colugos (Fig. 3). The presence of a deep flexor fibularis groove on the calcaneal sustentaculum has been considered a synapomorphy for primates related to stronger hallucal grasping, whereas this muscle has been considered to play a less active role in treeshrews and colugos (18). In fact, a deep flexor fibularis groove is present on the calcaneus of euprimates and more derived plesiadapiforms (including paromomyids and plesiadapoids). The combination of a large groove for the tendon of the flexor fibularis on the astragalus and absence of a deep groove on the calcaneus in Purgatorius and micromomyids may be a primitive retention in these taxa. Similar characteristics are present in Protungulatum and the Cretaceous eutherian Deccanolestes, whose affinities lie well outside the Euarchonta (Fig. 2).

Discussion

The evolution of diagnostic euprimate traits associated with grasping, leaping, and an enhanced visual system has long been thought to relate in part to arboreality (24), although substrate preferences of our earliest primate ancestors have been less clear. Certain features of euprimates, such as grasping hands and feet, already had evolved to various degrees among plesiadapiforms (15, 20, 23, 2528). Nearly all plesiadapiform species are known only from fossil dentitions, and the several known partial skeletons belong to fairly derived and relatively late-occurring members of their respective clades (23). Based on ecological inferences from the shape of the skull and teeth, it has been suggested that arboreality and herbivory evolved independently in plesiadapiforms and euprimates following their divergence from a ground-dwelling, insectivorous ancestor (29). However, tarsals of Purgatorius reported here indicate instead that arboreality was characteristic of the oldest and most primitive known stem primate. Purgatorius is more primitive than other plesiadapiforms and euprimates in retaining three lower incisors, four lower premolars, and molars with taller trigonids and more acute cusps that likely reflect an omnivorous diet that included a large proportion of insects (3, 13). Like the dentition (3, 4), the tarsals of Purgatorius reflect a plesiomorphic state that is sufficiently primitive to have given rise to the more derived morphologies present in all later primates.
The major radiation of angiosperms in the Late Cretaceous continued throughout the earliest Paleocene and dominated megafloras in the North American western interior (30, 31). Within this context, the immigration of Purgatorius represents the infusion of a unique arboreal mammal into North America during the first million years following the K–Pg boundary (32). Increased size of seeds and fruits is correlated with increases in the proportions of animal-dispersed taxa during this time (33) and would have provided an arboreal and omnivorous primate such as Purgatorius with angiosperm products including fruits, flowers, and associated insect pollinators (15, 3436). Therefore, the postcranial specializations for arboreality documented in Purgatorius would have allowed this animal to access resources that were not directly available to many contemporary terrestrial mammals, such as Protungulatum. The fossil record provides a direct test to evaluate adaptive scenarios, however incremental (8), and future recovery and analysis of early euarchontan fossils will continue to improve our understanding of primate origins. The previously unidentified fossils of Purgatorius described here suggest that the divergence of primates from other mammals was not a dramatic event. Instead, the beginning of primate evolutionary history likely involved subtle changes in the postcranial skeleton that allowed easier navigation and improved access to food resources in an arboreal setting.

Materials and Methods

Regression Analysis.

To assess whether tarsals described here (SI Appendix, Table S1) are of a size consistent with their attribution to the Purgatorius dental sample from the Garbani Channel fauna (SI Appendix, Fig. S1), least squares linear regression analyses were run in Microsoft Excel to evaluate the scaling relationship between the natural log area of the second lower molar and astragalar tibial trochlea width, as well as between the natural log second lower molar area and calcaneal cuboid facet area for euarchontan mammals. Skeletal elements from a sample of 60 dentally associated skeletons of euarchontans including extant taxa and fossil plesiadapiforms were microCT scanned, and digital reconstructions were measured using Avizo 6 software (SI Appendix, Table S2). The 95% confidence limits on the prediction interval of tooth size from postcranial element dimensions were generated using equation 17.29 of ref. 37. Dimensions from isolated tarsals and teeth of Purgatorius (SI Appendix, Table S3) then were plotted on the resulting regression equations (SI Appendix, Fig. S2).

Principal Component Analysis.

To evaluate our qualitative observations that the tarsals attributed to Purgatorius are generally similar to those of euarchontan mammals and are specifically similar to those of plesiadapiforms, we ran principal component analysis on the correlation matrix derived from 18 linear and 5 angular astragalar measurements (SI Appendix, Fig. S3A) following ref. 38 for 48 individuals representing 34 species (SI Appendix, Table S4) and 19 linear and 6 angular calcaneal measurements (SI Appendix, Fig. S3B) following ref. 39 for 54 individuals representing 33 species (SI Appendix, Table S5). Additionally, we ran a cluster analysis using the correlation matrix as our similarity metric and using the paired group method for linking cases. All analyses were run using PAST v. 2.16 (40). All linear measurements were size-standardized using the geometric mean of a subset of the measures. Angular measurements are reported in degrees but were analyzed in radians. The expanded taxonomic sample includes Puercan mammals, fossil plesiadapiforms and euprimates, and extant euarchontans (SI Appendix, Tables S4 and S5). All tarsals were microCT scanned, and digital reconstructions were created and measured using Avizo 6 software. Eigenvalue, percentage variance, and variable component loadings were recorded for each principal component (SI Appendix, Tables S6 and S7).

Phylogenetic Analysis.

Cladistic analysis using maximum parsimony was performed in TNT (41) on three revised character matrices (12, 15, 16). Four plesiadapiforms, one colugo, and new Purgatorius tarsal data were added to the character matrix of ref. 12, and new Purgatorius tarsal data were added to the character matrices of refs. 15 and 16. In all analyses, New Technology Search was used to obtain the stabilized consensus five times, and resulting most parsimonious trees (MPTs) were used as starting trees in a Traditional Heuristic Search that was carried out using tree bisection reconnection (TBR). All resulting MPTs were used to obtain a strict consensus, and, following the methods of ref. 12, the Pruned Trees function was used to identify the least stable taxa, which were removed using the Prune Taxa function if large polytomies were present. The Tree Filter function was used to delete longer trees and duplicate MPTs. Bremer branch supports were calculated using the Traditional Search option (10 replicates per run with TBR enforced) from 50,000 suboptimal trees up to 10 steps longer than the most parsimonious tree. See SI Appendix for more detailed methodology, list of specimens analyzed, and specific modifications to character matrices (SI Appendix, Tables S8–S10).

Acknowledgments

We thank P. Holroyd, E. Sargis, C. Manz, G. Wilson, L. Debey, K. Pugh, C. Sprain, P. Renne, W. Mitchell III, M. Silcox, and A. Hill for helpful discussions; eight anonymous reviewers and the editor for helpful comments; the Engdahl family and Bureau of Land Management for help with fieldwork; J. VanHouten, S. Judex, and C. Ruben for CT scanning assistance; and G. Yapuncich, A. Garberg, J. Butler, and J. Lovoi for segmenting CT scans. S.G.B.C. was supported by National Science Foundation (NSF) Grant SBE-1028505 (to E. J. Sargis and S.G.B.C.), the Leakey Foundation, and a Brooklyn College Tow Faculty Travel Fellowship. J.I.B. was supported by NSF Grant SBR-9616194 (to G. F. Gunnell, P. D. Gingerich, and J.I.B.) and Yale Institute for Biospheric Studies. D.M.B. was supported by NSF Grant BCS 1317525 (to E. Seiffert and D.M.B.). W.A.C. was supported by NSF Grant EAR 9505847. This work also was supported by the Doris O. and Samuel P. Welles Research Fund, University of California Museum of Paleontology.

Footnotes