Additional postcranial remains of omomyid primates from the Uinta Formation, Utah and implications for the locomotor behavior of large-bodied omomyids

Most omomyids are relatively small bodied (e.g. <500 g), but beginning in the middle Eocene, some omomyids began to grow larger. The largest omomyids occur in the late middle Eocene during the Uintan NALMA, reaching an estimated body mass over 1 kg. The hind limb skeleton of small omomyids is relatively well known, and is generally thought to show active arboreal quadrupedal and leaping adaptations. New postcranial specimens of previously unknown elements from the larger Uintan omomyids, Ourayia (two species), Chipetaia lamporea, and Mytonius hopsoni have recently been recovered from the Uinta Formation, Utah, and from the Mission Valley Formation, California, and they provide additional information concerning their locomotor behavior.The new specimens include several distal tibiae, partial calcanei, a complete talus and a proximal first metatarsal of Chipetaia; distal femora, distal tibiae, cuboids, and partial calcanei of Ourayia uintensis; a complete calcaneus of Ourayia sp.; and a partial calcaneus and talus of Mytonius. Metric analysis of these elements, together with qualitative observations of non-metric traits, indicate that Ourayia and Chipetaia show equal or greater development of traits associated with leaping behavior (including elongation of the calcaneus, navicular and cuboid) than do smaller omomyids from North America. The elements of Mytonius, although fragmentary, lack some leaping features that are well-developed in Ourayia and Chipetaia, suggesting that Mytonius may have relied more on arboreal quadrupedal locomotion than on leaping.     

Hindlimb adaptations in Ourayia and Chipetaia, relatively large-bodied omomyine primates from the Middle Eocene of Utah

North American omomyids represent a tremendous Eocene radiation of primates exhibiting a wide range of body sizes and dietary patterns. Despite this adaptive diversity, relatively little is known of the postcranial specializations of the group. Here we describe hindlimb and foot bones of Ourayia uintensis and Chipetaia lamporea that were recovered from the Uinta B member (early Uintan Land Mammal Age), Uinta Formation, Utah. These specimens provide insights into the evolution of postcranial adaptations across different body sizes and dietary guilds within the Eocene primate radiation. Body mass estimates based on talar measurements indicate that Ourayia uintensis and Chipetaia lamporea weighed about 1,500-2,000 g and 500-700 g, respectively. Skeletal elements recovered for Ourayia include the talus, navicular, entocuneiform, first metatarsal, and proximal tibia; bones of Chipetaia include the talus, navicular, entocuneiform, and proximal femur. Both genera had opposable grasping big toes, as indicated by the saddle-shaped joint between the entocuneiform and first metatarsal. Both taxa were arboreal leapers, as indicated by a consistent assemblage of characters in all represented bones, most notably the somewhat elongated naviculars, the high and distinct trochlear crests of the talus, the posteriorly oriented tibial plateau (Ourayia), and the cylindrical head of the femur (Chipetaia). The closest resemblances to Ourayia and Chipetaia are found among the Bridger omomyines, Omomys and Hemiacodon. The results of our comparisons suggest that the later, larger, more herbivorous omomyines from Utah retained a skeletal structure characteristic of earlier, smaller North American omomyids.     

Significance of primate petrosal from Middle Eocene fissure-fillings at Shanghuang, Jiangsu Province, People's Republic of China

An isolated petrosal bone belonging to a diminutive primate is reported from Middle Eocene fissure-fills near Shanghuang (southern Jiangsu Province, People's Republic of China), the type locality of several newly described primates (Eosimias sinensis, a basal anthropoid; Adapoides troglodytes, a basal adapinan; Tarsius eocaenus, a congener of extant tarsiers; and Macrotarsius macrorhysis, the first Asian representative of an otherwise exclusively North American genus). Because of its fragmentary condition and unique combination of characters, the Shanghuang petrosal cannot be assigned unambiguously to any of the Shanghuang primate taxa known from dental remains. However, the possibility that the petrosal represents either an adapid or a tarsiid can be dismissed because it lacks defining basicranial apomorphines of these groups. By contrast, the element does present arterial features consistent with its being haplorhine. Deciding between the likeliest candidates for its allocation—Omomyidae and Eosimiidae—is difficult, in part because it is not known what (or even whether) basicranial characters can be used to distinguish these clades. If the Shanghuang petrosal is that of an cosimiid, as both direct and indirect evidence appears to indicate, the following implications emerge: (1) as long suspected on other grounds, anthropoids share a closer evolutionary history with Omomyidae (and Tarsiiformes) than they do with Adapidae (and Strepsirhini); (2) the specialised basicranial anatomy of extant anthropoids and their immediate cladistic relatives is derived from a primitive precursor whose otic morphology was like that of omomyids in most known respects; (3) the evolution of the defining dental and basicranial apomorphies of extant Anthropoidea has been distinctly mosaic in pattern.     

Body size and premolar evolution in the early‐middle eocene euprimates of Wyoming

The earliest euprimates to arrive in North America were larger‐bodied notharctids and smaller‐bodied omomyids. Through the Eocene, notharctids generally continued to increase in body size, whereas omomyids generally radiated within small‐ and increasingly mid‐sized niches in the middle Eocene. This study examines the influence of changing body size and diet on the evolution of the lower fourth premolar in Eocene euprimates. The P₄ displays considerable morphological variability in these taxa. Despite the fact that most studies of primate dental morphology have focused on the molars, P₄ can also provide important paleoecological insights. We analyzed the P₄ from 177 euprimate specimens, representing 35 species (11 notharctids and 24 omomyids), in three time bins of approximately equal duration: early Wasatchian, late Wasatchian, and Bridgerian. Two‐dimensional surface landmarks were collected from lingual photographs, capturing important variation in cusp position and tooth shape. Disparity metrics were calculated and compared for the three time bins. In the early Eocene, notharctids have a more molarized P₄ than omomyids. During the Bridgerian, expanding body size range of omomyids was accompanied by a significant increase in P₄ disparity and convergent evolution of the semimolariform condition in the largest omomyines. P₄ morphology relates to diet in early euprimates, although patterns vary between families. Am J Phys Anthropol 153:15–28, 2014. © 2013 Wiley Periodicals, Inc.     

Using extant patterns of dental variation to identify species in the primate fossil record: a case study of middle Eocene <Emphasis Type="Italic">Omomys</Emphasis> from the Bridger Basin,southwestern Wyoming

Patterns of extant primate dental variation provide important data for interpreting taxonomic boundaries in fossil forms. Here I use dental data from several well-known living primates (as well as data from selected Eocene forms) to evaluate dental variation in Middle Eocene Omomys, the first North American fossil primate identified by paleontologists. Measurements were collected from a sample of 148 omomyid dental specimens recovered from Bridger B localities in the Bridger Basin, Wyoming. Most of these specimens have not previously been described. Nonmetric traits were also scored for this sample. Lower molar coefficients of variation range from 4.01 for M₂ length (n = 80) to 6.73 for M₃ talonid width (n = 57). All of the nonmetric traits scored exhibit less than 100% presence in the overall sample, including traits previously described as representative of Omomys (e.g., P₄ metaconids present in 91%, n = 55; M² pericones present in 80%, n = 15). Dental traits also vary in a set of spatially restricted localities from the same fossil horizon and in a separate, single fossil locality (DMNH 868, P₄ metaconids present in 67%, n = 6). An increasing frequency in several premolar traits across time in these more restricted samples suggests an anagenetic change in Bridger B Omomys. However, this degree of morphological variability is consistent with that seen in extant primate species from single locations. Metric variation in this sample is comparable to that seen in other Eocene primates, such as new data presented here for the omomyid Arapahovius gazini from the Washakie Basin, southern Wyoming. Omomys metric variation is also comparable to that found in several samples of well-known extant primates from single localities (e.g., ring-tailed lemurs and gray–brown mouse lemurs). These metric data also correspond to the patterns of variability described in previously published studies of Omomys carteri. In sum, a single species interpretation (O. carteri) for this new Bridger B Omomys sample from southern Wyoming is affirmed, and this study illustrates the usefulness of dental data from extant primates for evaluating primate fossil samples.     

New early Eocene anaptomorphine primate (Omomyidae) from the Washakie Basin,Wyoming, with comments on the phylogeny and paleobiology of anaptomorphines

Recent paleontological collecting in the Washakie Basin, southcentral Wyoming, has resulted in the recovery of over 100 specimens of omomyid primates from the lower Eocene Wasatch Formation. Much of what is known about anaptomorphine omomyids is based upon work in the Bighorn and Wind River Basins of Wyoming. This new sample documents greater taxonomic diversity of omomyids during the early Eocene and contributes to our understanding of the phylogeny and adaptations of some of these earliest North American primates. A new middle Wasatchian (Lysitean) anaptomorphine, Anemorhysis savagei, n. sp., is structurally intermediate between Teilhardina americana and other species of Anemorhysis and may be a sister group of other Anemorhysis and Trogolemur. Body size estimates for Anemorhysis, Tetonoides, Trogolemur, and Teilhardina americana indicate that these animals were extremely small, probably less than 50 grams. Analysis of relative shearing potential of lower molars of these taxa indicates that some were primarily insectivorous, some primarily frugivorous, and some may have been more mixed feeders. Anaptomorphines did not develop the extremes of molar specialization for frugivory or insectivory seen in extant prosimians. Incisor enlargement does not appear to be associated with specialization in either fruits or insects but may have been an adaptation for specialized grooming or food manipulation. © 1994 Wiley-Liss, Inc.     

Mandibular form and function in North American and European Adapidae and Omomyidae

Previous experimental and comparative studies among a wide variety of primate and nonprimate mammals provide a unique source of information for investigating the functional and phylogenetic significance of variation in the masticatory apparatus of Eocene primates. To provide a quantitative study of mandibular form and function in Eocene primates, the scaling of jaw dimensions and the development of symphyseal fusion was considered in a broad sample of North American and European Adapidae and Omomyidae. Statistical analyses indicate a significant size-related pattern of symphyseal fusion across Eocene primates, with larger taxa often having a greater degree of fusion than smaller species; this trend is also evident at the family level. As adapids are mostly larger than omomyids and these taxa show allometry of symphyseal fusion, this may explain why no omomyids evince complete fusion. Controlling for jaw size, species with greater symphyseal fusion tend to have more robust jaws than those with a lesser amount of fusion. Upon further examination, a primary reason why adapids have more robust mandibles than omomyids is associated with the presence of taxa with fused symphyses, and thus more robust jaws, in the adapid sample, whereas no omomyids have fused symphyses. In addition, there is little indication of a dietary effect, as measured by molar shear-crest development, on symphyseal fusion. Moreover, as there is no correlation between molar shear-crest development and skull size, this also points to the absence of a size-related pattern of dietary preference underlying the allometry of symphyseal fusion. Based on the interspecific and ontogenetic allometry of symphyseal ossification in Eocene primates, jaw-scaling patterns are used to further examine the functional determinants of fusion in this group. This study indicates that greater dorsoventral shear during mastication is a more likely factor than lateral transverse bending (“wishboning”) in the evolution of symphyseal fusion among “late-fusing” mammals like adapids and omomyids. Given that wishboning is an important functional determinant of symphyseal form in recent anthropoids, apparently the evolutionary development of marked wishboning occurs only in taxa that shift the timing of fusion to a growth stage preceding the onset of weaning (before adult masticatory patterns are fully developed) and perhaps first ossified the symphysis to counter elevated dorsoventral shear stress. As early anthropoids probably consisted of members varying interspecifically and ontogenetically in the degree of ossification, it is especially informative to analyze the adaptive setting in which anthropoid symphyseal fusion evolved from a similar primitive “prosimian” perspective. © 1996 Wiley-Liss, Inc.     

Brain,body and encephalization in early primates

Encephalization—the evolution of relatively enlarged brains—was probably a characteristic adaptation in the order Primates from the earliest times. Evidence for this generalization is reviewed by reanalyzing data on brain size and body size in Paleocene, Eocene and Oligocene genera: Plesiadapis, Tetonius, Necrolemur, Smilodectes, Adapis, Rooneyia and Aegyptopithecus. Uncertainties about the generalization are based primarly on problems in the estimation of body size. Mathematical (dimensional) and statistical issues in those estimations are reviewed and the errors-of-estimate are presented quantitatively. These are small enough to suggest that the generalization is correct. Although the early primates were progressive with respect to encephalization, only the omomyids appear to have reached present (tarsiid) grades during the earliest times. Plesiadapis and the adapids appeared to be somewhat below the present strepsorhine grade, and Aegyptopithecus was at a strepsorhine rather than haplorhine grade of encephalization, according to presently available evidence.     

A new basilosaurid (Cetacea, Pelagiceti) from the Late Eocene to Early Oligocene Otuma Formation of Peru

Cynthiacetus peruvianus nov. sp. is a new basilosaurid species, from Late Eocene to Early Oligocene Otuma Formation of Peru. It is the first described archaeocete in South-America and is represented by a sub-complete skeleton. C. peruvianus differs from C. maxwelli (middle to Late Eocene of Egypt and United States) principally in having one cuspid less on both mesial and distal sides of p3 and p4. Cynthiacetus is among the largest basilosaurids. Its more characteristic features are located on its postcranial skeleton: large vertebrarterial foramina on cervical vertebrae and absence of ventral expansion of the transverse process on C3-C5. Besides, C. peruvianus presents the greatest number of thoracic vertebrae (20) and ribs observed in Cetacea and the first thoracics have an almost vertical neural spine. A preliminary parsimony analysis establishes the monophyly of the Basilosauridae on the basis of three unambiguous cranial synapomorphies. However, within the Basilosauridae, the most diagnostic characters are observed on the postcranial skeleton.     

A new tarkadectine primate from the Eocene of Inner Mongolia,China: phylogenetic and biogeographic implications

Tarka and Tarkadectes are Middle Eocene mammals known only from the Rocky Mountains region of North America. Previous work has suggested that they are members of the Plagiomenidae, an extinct family often included in the order Dermoptera. Here we describe a new primate, Tarkops mckennai gen. et sp. nov., from the early Middle Eocene Irdinmanha Formation of Inner Mongolia, China. The new taxon is particularly similar to Tarka and Tarkadectes, but it also displays many features observed in omomyids. A phylogenetic analysis based on a data matrix including 59 taxa and 444 dental characters suggests that Tarkops, Tarka and Tarkadectes form a monophyletic group—the Tarkadectinae—that is nested within the omomyid clade. Within Omomyidae, tarkadectines appear to be closely related to Macrotarsius. Dermoptera, including extant and extinct flying lemurs and plagiomenids, is recognized as a clade nesting within the polyphyletic group of plesiadapiforms, therefore supporting the previous suggestion that the relationship between dermopterans and primates is as close as that between plesiadapiforms and primates. The distribution of tarkadectine primates on both sides of the Pacific Ocean basin suggests that palaeoenvironmental conditions appropriate to sustain primates occurred across a vast expanse of Asia and North America during the Middle Eocene.     

Phylogeny of Schisandraceae based on morphological data: evidence from modern plants and the fossil record

Schisandraceae are traditionally subdivided in two genera, Schisandra and Kadsura, based on differences in the organisation of the floral receptacle, the carpels, and the presence or absence of a ``pseudostigma'. Recently, phylogenetic analyses utilizing ITS sequence data and morphological data resulted in incongruent tree topologies, with the morphological trees suggesting monophyly of the two genera, whereas ITS trees did not resolve Schisandra and Kadsura as monophyletic clades. In the present paper we study seed morphology and leaf epidermal features of 22 species of Schisandraceae in order to provide additional data for a morphological data matrix. Seed morphological characters are highly homoplastic and do not yield further evidence for monophyly of the two genera. Instead, a number of characters appear to support sister group relationships between taxa within the genera, such as, for instance, for K. coccinea and K. scandens, both of which have large seeds along with a multi-layered mesotesta. Considering leaf epidermal characteristics, species of Kadsura were found to be consistently amphistomatic, whereas species of Schisandra are always hypostomatic. Phylogenetic analysis using the extended data matrix resulted in weakly supported Kadsura and Schisandra clades with five and four synapomorphies indicating monophyly of Kadsura and Schisandra, respectively. Fossils ascribed to Schisandraceae date back to the Late Cretaceous. These are tri-and hexacolpate pollen types displaying a combination of features found in modern Schisandraceae and partly also in Illiciaceae. Leaf remains from this period are poorly preserved and difficult to ascribe to Schisandraceae because of the lack of synapomorphies for the family. In the Early Cainozoic, leaf and seed remains from North America and Europe unambiguously belong to the family. Seeds from the Eocene of North America show some similarities to the modern Schisandra glabra from North America, while fossils from Europe show more similarities to modern Asian species.     

In vivo baseline measurements of hip joint range of motion in suspensory and nonsuspensory anthropoids

Hominoids and atelines are known to use suspensory behaviors and are assumed to possess greater hip joint mobility than nonsuspensory monkeys, particularly for range of abduction. This assumption has greatly influenced how extant and fossil primate hip joint morphology has been interpreted, despite the fact that there are no data available on hip mobility in hominoids or Ateles. This study uses in vivo measurements to test the hypothesis that suspensory anthropoids have significantly greater ranges of hip joint mobility than nonsuspensory anthropoids. Passive hip joint mobility was measured on a large sample of anesthetized captive anthropoids (nonhuman hominids = 43, hylobatids = 6, cercopithecids = 43, Ateles = 6, and Cebus = 6). Angular and linear data were collected using goniometers and tape measures. Range of motion (ROM) data were analyzed for significant differences by locomotor group using ANOVA and phylogenetic regression. The data demonstrate that suspensory anthropoids are capable of significantly greater hip abduction and external rotation. Degree of flexion and internal rotation were not larger in the suspensory primates, indicating that suspension is not associated with a global increase in hip mobility. Future work should consider the role of external rotation in abduction ability, how the physical position of the distal limb segments are influenced by differences in ROM proximally, as well as focus on bony and soft tissue differences that enable or restrict abduction and external rotation at the anthropoid hip joint. Am J Phys Anthropol 153:417–434, 2014. © 2013 Wiley Periodicals, Inc.     

Isolation and characterization of 12 microsatellite loci for population studies of Sulawesi tarsiers (Tarsius spp.)

This study reports the development and characterization of the first 12 microsatellite markers for tarsiers. Nine loci were isolated from Dian's tarsier, Tarsius dianae and three from the Philippine tarsier, Tarsius syrichta. The 12 markers were used to screen 40 individuals of Dian's tarsier and 40 individuals of the Lariang tarsier, Tarsius lariang for allelic diversity. This suite of highly polymorphic microsatellites provides the first chance to genetically study parentage patterns in tarsiers.     

A South American snake lineage from the Eocene Greenhouse of North America and a reappraisal of the fossil record of “anilioid” snakes

“Anilioidea” is a likely paraphyletic assemblage of pipe snakes that includes extant Aniliidae from equatorial South America, Uropeltoidea from South and Southeast Asia, and a fossil record that consists primarily of isolated precloacal vertebrae ranging from the earliest Late Cretaceous and includes geographic distributions in North America, South America, Europe, and Africa. Articulated precloacal vertebrae from the middle Eocene Bridger Formation of Wyoming, attributed to Borealilysia nov. gen., represent an unambiguous North American aniliid record and prompts a reconsideration of described pipe snakes and their resultant biogeographic histories. On the basis of vertebral apomorphies, the vast majority of reported fossils cannot be assigned to “Anilioidea”. Instead, most records represent stem taxa and macrostomatans erroneously assigned to anilioids on the basis of generalized features associated with fossoriality. A revised fossil record demonstrates that the only extralimital distributions of fossil “anilioids” consist of the North American aniliid record, and there is no unambiguous fossil record of Old World taxa. The occurrence of aniliids in the mid-high latitudes of the late early Eocene of North America is consistent with histories of northward shifts in equatorial ecosystems during the early Paleogene Greenhouse.     

Skeletal adaptations and phylogeny of the oldest mole Eotalpa (Talpidae,Lipotyphla, Mammalia) from the UK Eocene: the beginning of fossoriality in moles

The oldest talpid, Eotalpa, was previously known only from isolated cheek teeth from the European late Middle Eocene to earliest Oligocene. Screenwashing of Late Eocene sediments of the Hampshire Basin, UK, has yielded cranial and postcranial elements: maxilla, dentary, ulna, metacarpals, distal tibia, astragalus, calcaneum, metatarsals and phalanges. In addition to M_1–2 myotodonty, typical talpid features are as follows: ulna with long medially curved olecranon and deep abductor fossa and astragalar body with lateral process. However, Eotalpa retains certain soricid‐like primitive states (M¹ preparacrista, P⁴ with prominent mesiolingual protocone lobe, strongly angled astragalar neck and calcaneum with no space for a cuboid medial process) not found in modern talpids. Eotalpa is more derived than the most primitive living talpid Uropsilus in having lost the M^1–2 talon shelf, developed a convex radial facet on the ulna, an incipient proximal olecranon crest, relatively shorter metapodials and depressed manual unguals. Its astragalus with medial trochlear ridge taller than the lateral one and massive medial plantar process is typical of the Lipotyphla. Eotalpa lacks synostosis of tibia and fibula, found in other Talpidae, Soricidae and Erinaceidae, suggesting that synostosis in these groups has been independently acquired. Cladistic analysis places Eotalpa as stem member of the Talpidae and shows that much homoplasy arose during the early evolution of the family. Ground dwelling in Eotalpa is indicated by the following: astragalus with a medially dipping head, curved in a single plane; calcaneum with distal peroneal process and strongly overlapping ectal and sustentacular facets; and matching sized ectal and sustentacular facets on calcaneum and astragalus. These features would have restricted ankle mobility. Ungual and metatarsal shape and ulnar structure suggest a primitive stage in fossorial evolution and argue against a semiaquatic precursor stage in talpid fossoriality. Shrew‐moles may represent a reversal to surface foraging rather than an intermediate stage in fossoriality.     

Comparative study of leaf architecture and cuticles of Nelumbo changchangensis from the Eocene of Hainan Island,China, and the two extant species of Nelumbo (Nelumbonaceae)

Fossil leaves of Nelumbo changchangensis, collected from the Eocene of Hainan Island, China, were studied and compared with those of the extant species of Nelumbo, N. nucifera Gaertn. and N. lutea Willd. The fossil leaves have all the specialized features of extant Nelumbo in leaf architecture, except that the organization of the areolae looks much more irregular than that of extant Nelumbo. Comparisons of the cuticle and epicuticular ultrastructure indicate that: (1) N. changchangensis resembles N. nucifera in that anticlinal cell walls of the lower epidermis are straight along the major veins and near leaf bases and are shallowly undulate with U‐ to V‐shaped undulations inside the areolae; (2) N. changchangensis differs from N. lutea in that anticlinal cell walls of the lower epidermis of the latter are deeply undulate with U‐, V‐ to reversed Ω‐shaped undulations inside the areolae; and (3) epicuticular wax crystals are more densely distributed on the leaves of N. changchangensis and N. nucifera than they are in N. lutea. These findings shed significant light on the cuticle differentiation of fossil and extant Nelumbo species. The morphometric comparisons indicate that almost all the synapomorphies of extant Nelumbo were already present by the Eocene, © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 180 , 123–137.     

A new kinosternoid from the Late Cretaceous Hell Creek Formation of North Dakota and Montana and the origin of the <Emphasis Type="Italic">Dermatemys mawii</Emphasis> lineage

A nearly complete turtle shell from the Late Cretaceous (Maastrichtian) Hell Creek Formation of Slope County, North Dakota, represents the most complete remains to date of a Mesozoic kinosternoid turtle and a new species, Hoplochelys clark nov. sp. The new taxon is diagnosable from other representatives of Hoplochelys by the plesiomorphic placement of the humeral/femoral sulcus behind the hyo/hypoplastral suture and the autapomorphic development of an interrupted median (neural) keel. All six previously named Paleocene (Puercan and Torrejonian) representatives of Hoplochelys lack diagnostic characters and are synonymized as Hoplochelys crassa. A phylogenetic analysis reveals that Hoplochelys spp. and Agomphus pectoralis are most parsimoniously placed within Kinosternoidea along the phylogenetic stem of the extant Mesoamerican River Turtle Dermatemys mawii, extending that taxon’s stem lineage from the early Eocene to the late Maastrichtian. The two primary crown lineages of Kinosternoidea are thus known from the Mesozoic and split prior to the late Campanian. The presence of a thickened cruciform plastron, true costiform processes, only three inframarginals, and the reduction of the medial contact of the abdominals are synapomorphies of Chelydroidea, the clade formed by Chelydridae and Kinosternoidae.