Stephen G. B. Chester

Evolution of the Earliest Horses Driven by Climate Change in the Paleocene-Eocene Thermal Maximum

Warming and Shrinking In most mammals, individual body sizes tend to be smaller in warmer regions and larger in cooler regions. Secord et al. (p. 959; see the Perspective by Smith) examined a high-resolution 175,000-year record of equid fossils deposited over a past climate shift—the Paleocene-Eocene Thermal Maximum—for changes in body size. Using oxygen isotopes collected from the teeth of co-occurring mammal species to track prevailing environmental temperature, a clear decrease in equid body size was seen during 130,000 years of warming, followed by a distinct increase as the climate cooled at the end of the period. These results indicate that temperature directly influenced body size in the past and may continue to have an influence as our current climate changes. Oxygen isotope measurements of fossil teeth show that the body size of the horse Sifrhippus decreased as temperature increased. Body size plays a critical role in mammalian ecology and physiology. Previous research has shown that many mammals became smaller during the Paleocene-Eocene Thermal Maximum (PETM), but the timing and magnitude of that change relative to climate change have been unclear. A high-resolution record of continental climate and equid body size change shows a directional size decrease of ~30% over the first ~130,000 years of the PETM, followed by a ~76% increase in the recovery phase of the PETM. These size changes are negatively correlated with temperature inferred from oxygen isotopes in mammal teeth and were probably driven by shifts in temperature and possibly high atmospheric CO2 concentrations. These findings could be important for understanding mammalian evolutionary responses to future global warming.

New fossils of the oldest North American euprimate Teilhardina brandti (Omomyidae) from the paleocene–eocene thermal maximum

More than 25 new specimens of Teilhardina brandti, one of the oldest known euprimates, are reported from earliest Eocene strata of the southern Bighorn Basin, Wyoming. The new fossils include the first upper dentitions, a dentary showing the lower dental formula for the first time, and the first postcrania ascribed to T. brandti (tarsals and terminal phalanges). The elongated navicular and long talar neck suggest that T. brandti was an active arboreal quadruped, and the terminal phalanges constitute the oldest evidence for nails in Euprimates. Phylogenetic analysis incorporating the new data indicates that T. brandti is more derived than T. belgica but less so than T. americana. The hypothesis that Teilhardina originated in Asia (T. asiatica) and dispersed westward to Europe (T. belgica) and then to North America (T. brandti and T. magnoliana) during the earliest Eocene Paleocene-Eocene Thermal Maximum is most consistent with available evidence, including the relative age of fossil samples and their stage of evolution.

Hands of early primates

Questions surrounding the origin and early evolution of primates continue to be the subject of debate. Though anatomy of the skull and inferred dietary shifts are often the focus, detailed studies of postcrania and inferred locomotor capabilities can also provide crucial data that advance understanding of transitions in early primate evolution. In particular, the hand skeleton includes characteristics thought to reflect foraging, locomotion, and posture. Here we review what is known about the early evolution of primate hands from a comparative perspective that incorporates data from the fossil record. Additionally, we provide new comparative data and documentation of skeletal morphology for Paleogene plesiadapiforms, notharctines, cercamoniines, adapines, and omomyiforms. Finally, we discuss implications of these data for understanding locomotor transitions during the origin and early evolutionary history of primates. Known plesiadapiform species cannot be differentiated from extant primates based on either intrinsic hand proportions or hand-to-body size proportions. Nonetheless, the presence of claws and a different metacarpophalangeal [corrected] joint form in plesiadapiforms indicate different grasping mechanics. Notharctines and cercamoniines have intrinsic hand proportions with extremely elongated proximal phalanges and digit rays relative to metacarpals, resembling tarsiers and galagos. But their hand-to-body size proportions are typical of many extant primates (unlike those of tarsiers, and possibly Teilhardina, which have extremely large hands). Non-adapine adapiforms and omomyids exhibit additional carpal features suggesting more limited dorsiflexion, greater ulnar deviation, and a more habitually divergent pollex than observed plesiadapiforms. Together, features differentiating adapiforms and omomyiforms from plesiadapiforms indicate increased reliance on vertical prehensile-clinging and grasp-leaping, possibly in combination with predatory behaviors in ancestral euprimates.

Oldest known euarchontan tarsals and affinities of Paleocene Purgatorius to Primates.

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. 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.

RUNX2 tandem repeats and the evolution of facial length in placental mammals

BackgroundWhen simple sequence repeats are integrated into functional genes, they can potentially act as evolutionary ‘tuning knobs’, supplying abundant genetic variation with minimal risk of pleiotropic deleterious effects. The genetic basis of variation in facial shape and length represents a possible example of this phenomenon. Runt-related transcription factor 2 (RUNX2), which is involved in osteoblast differentiation, contains a functionally-important tandem repeat of glutamine and alanine amino acids. The ratio of glutamines to alanines (the QA ratio) in this protein seemingly influences the regulation of bone development. Notably, in domestic breeds of dog, and in carnivorans in general, the ratio of glutamines to alanines is strongly correlated with facial length.ResultsIn this study we examine whether this correlation holds true across placental mammals, particularly those mammals for which facial length is highly variable and related to adaptive behavior and lifestyle (e.g., primates, afrotherians, xenarthrans). We obtained relative facial length measurements and RUNX2 sequences for 41 mammalian species representing 12 orders. Using both a phylogenetic generalized least squares model and a recently-developed Bayesian comparative method, we tested for a correlation between genetic and morphometric data while controlling for phylogeny, evolutionary rates, and divergence times. Non-carnivoran taxa generally had substantially lower glutamine-alanine ratios than carnivorans (primates and xenarthrans with means of 1.34 and 1.25, respectively, compared to a mean of 3.1 for carnivorans), and we found no correlation between RUNX2 sequence and face length across placental mammals.ConclusionsResults of our diverse comparative phylogenetic analyses indicate that QA ratio does not consistently correlate with face length across the 41 mammalian taxa considered. Thus, although RUNX2 might function as a ‘tuning knob’ modifying face length in carnivorans, this relationship is not conserved across mammals in general.

A New Small-Bodied Species of Palaeonictis (Creodonta, Oxyaenidae) from the Paleocene-Eocene Thermal Maximum

Oxyaenid creodonts are extinct carnivorous mammals known from the Paleogene of North America, Europe, and Asia. The genus Palaeonictis is represented by three species that together span the late Paleocene to early Eocene of North America, and at least one species from the early Eocene of Europe. Previously, only a single trigonid of Palaeonictis was known from the interval encompassing the Paleocene-Eocene Thermal Maximum (PETM) in North America. We describe Palaeonictis wingi sp. nov. from the PETM in the Cabin Fork drainage, southeastern Bighorn Basin, Wyoming, based on associated right and left dentaries with P2-M2. Palaeonictis wingi sp. nov. is substantially smaller than the other North American congeners, making it similar in size to P. gigantea from the earliest Eocene of Europe and the previously described PETM specimen. We suggest that a form similar to the large-bodied late Paleocene P. peloria from North America gave rise to two smaller species in the earliest Eocene of North America (P. wingi) and Europe (P. gigantea). Palaeonictis wingi may have given rise to P. occidentalis following the PETM in North America. Dispersal of Palaeonictis to Europe coincided with rapid global warming of 5–10°C and related geographic range shifts in plants and other animals during the PETM. It has been suggested that certain mammalian lineages decreased in body size during the PETM, possibly in response to elevated temperature and/or higher CO2 levels. Results from a dietary analysis of Palaeonictis indicate that it was an omnivore that primarily consumed meat. This suggests that the decreased nutritious quality of vegetation caused by increased CO2 levels was not the direct contributing factor that caused body size reduction of this lineage during the PETM. Other selective pressures such as temperature, aridity, and prey size may have also contributed to the smaller body size of carnivorous mammals during this interval, although the presence of smaller species could also be explained by latitudinal range shifts of mammals during the PETM.

Mammalian distal humeri from the Late Cretaceous of Uzbekistan

We describe mammalian distal humeri recovered from the Bissekty Formation, Dzharakuduk, Kyzylkum Desert, Uzbekistan (90 Ma). Isolated elements were sorted into groups that likely correspond to species (or genera). These groups were allocated to taxa known mostly from the dentition, petrosals, and/or tarsals at this site. We identified one humerus of a multituberculate and one of a zalambdalestid. Several eutherian humeri have been tentatively assigned to Zhelestidae based on their dissimilarity to zalambdalestids and the abundance of zhelestids in the dental record. The zalambdalestids and zhelestids were probably terrestrial. At least two metatherian taxa have also been identified, and both were likely arboreal. Although the dental record suggests twelve eutherian species and only one metatherian, crurotarsal evidence supports the presence of at least four metatherian species at Dzharakuduk. The humeri analyzed here also provide support for the presence of multiple metatherians in the fauna, further demonstrating that postcrania are critical to understanding the taxonomic diversity present at these Late Cretaceous localities.

Dinosaur extinction: closing the '3 m gap'

Modern debate regarding the extinction of non-avian dinosaurs was ignited by the publication of the Cretaceous–Tertiary (K–T) asteroid impact theory and has seen 30 years of dispute over the position of the stratigraphically youngest in situ dinosaur. A zone devoid of dinosaur fossils reported from the last 3 m of the Upper Cretaceous, coined the ‘3 m gap’, has helped drive controversy. Here, we report the discovery of the stratigraphically youngest in situ dinosaur specimen: a ceratopsian brow horn found in a poorly rooted, silty, mudstone floodplain deposit located no more than 13 cm below the palynologically defined boundary. The K–T boundary is identified using three criteria: (i) decrease in Cretaceous palynomorphs without subsequent recovery, (ii) the existence of a ‘fern spike’, and (iii) correlation to a nearby stratigraphic section where primary extraterrestrial impact markers are present (e.g. iridium anomaly, spherules, shocked quartz). The in situ specimen demonstrates that a gap devoid of non-avian dinosaur fossils does not exist and is inconsistent with the hypothesis that non-avian dinosaurs were extinct prior to the K–T boundary impact event.

The evolutionary radiation of plesiadapiforms

Very shortly after the disappearance of the non‐avian dinosaurs, the first mammals that had features similar to those of primates started appearing. These first primitive forms went on to spawn a rich diversity of plesiadapiforms, often referred to as archaic primates. Like many living primates, plesiadapiforms were small arboreal animals that generally ate fruit, insects, and, occasionally, leaves. However, this group lacked several diagnostic features of euprimates. They also had extraordinarily diverse specializations, represented in eleven families and more than 140 species, which, in some cases, were like nothing seen since in the primate order. Plesiadapiforms are known from all three Northern continents, with representatives that persisted until at least 37 million years ago. In this article we provide a summary of the incredible diversity of plesiadapiform morphology and adaptations, reviewing our knowledge of all eleven families. We also discuss the challenges that remain in our understanding of their ecology and evolution.

Therian Femora from the Late Cretaceous of Uzbekistan

Femora referable to metatherians and eutherians recovered from the Bissekty Formation, Dzharakuduk, Kyzylkum Desert, Uzbekistan (90 Mya), are described. Fourteen isolated specimens were sorted based on size and morphology into groups that likely correspond to the species level or higher. Groups were then tentatively assigned to taxa known from teeth, petrosals, and/or other postcrania at these localities. One distal femur of a small arboreal metatherian, and several eutherian distal femora that probably represent zhelestids and/or zalambdalestids were identified. With the exception of one proximal femur that is similar in some aspects to the zalambdalestid Barunlestes, and a previously described multituberculate specimen, all other proximal femora from the Bissekty Formation exhibit a metatherian-like morphology. The dental record currently suggests the presence of twelve eutherian species and only one metatherian at Dzharakuduk, whereas the humeral and crurotarsal evidence supports the presence of at least two or four metatherian species, respectively. Given the sample size of the proximal femora, the morphological diversity present, and the overwhelming presence of eutherians at these localities, it is highly unlikely that the overwhelming majority of proximal femora actually represent metatherians. Therefore, this sample may suggest that the metatherian proximal femoral condition is primitive for Theria and that some eutherian taxa (probably including Zhelestidae, which are dentally most abundant at these localities) retain this condition.