Search

Temperatures in the Arctic have increased by an astounding 1[degrees]C in response to anthropogenic forcing over the past 20 years and are expected to rise further in the coming decades. The Pliocene (2.6-5.3 Ma) is of particular interest as an analog for future warming because global temperatures were significantly warmer than today for a sustained period of time, with continental configurations similar to present. Here, we estimate mean annual temperature (MAT) based upon three independent proxies from an early Pliocene peat deposit in the Canadian High Arctic. Our proxies, including oxygen isotopes and annual ring widths (MAT = -0.5 [+ or -] 1.9[degrees]C), coexistence of paleovegetation (MAT = -0.4 [+ or -] 4.1[degrees]C), and bacterial tetraether composition in paleosols (MAT = -0.6 [+ or -] 5.0[degrees]C), yield estimates that are statistically indistinguishable. The consensus among these proxies suggests that Arctic temperatures were ~19[degrees]C warmer during the Pliocene than at present, while atmospheric C[O.sub.2] concentrations were ~390 ppmv. These elevated Arctic Pliocene temperatures result in a greatly reduced and asymmetrical latitudinal temperature gradient that is probably the result of increased poleward heat transport and decreased albedo. These results indicate that Arctic temperatures may be exceedingly sensitive to anthropogenic C[O.sub.2] emissions. doi: 10.1130/G30815.1

The beaver (Castor canadensis) is more specialized for swimming than any other living rodent and represents a good model for investigating questions pertaining to the evolutionary transition from terrestrial to aquatic habits. Aquatic specializations in the beaver include a streamlined body, enlarged webbed hind-feet, and a specialized tail. The tail is comprised of a muscular base and a distal paddle, which is dorsoventrally flattened and possesses a scaly integument. Most of the paddle is composed of fibrous connective-tissues embedded in a 'matrix' of adipose tissue. Lateral to the caudal vertebrae crossed structural fibers are attached to the dorsal and ventral dermis, and are oriented between 90 and 45[degrees] to the horizontal. Previous workers have suggested that crossed-array patterns of connective fibers are associated with structures that exhibit stereotyped movements. Stereotyped movements by the beaver tail during swimming can occur, but have not been described as few studies have examined swimming by this aquatic rodent. Observations of two captive beavers found that the rail is not held rigidly during swimming, but undulates. Posteriorly traveling waves in the tail were observed as the animals were simultaneously stroking with the hind feet. The waves moved on average 72% faster than the forward velocity of the animal, indicating generation of thrust by the broad tail (N = 25, Standard deviation = 55). These findings are consistent with the hypothesis that the beaver rail is specialized to produce undulatory propulsion. The addition of undulatory propulsion to hind foot paddling is consistent with proposed evolutionary stages associated with transition to fully aquatic specializations in mammals.

Castorids present an unusual opportunity to study behavioral evolution because their robust fossil record includes numerous behavioral artifacts. For some castorids the behavioral evidence indicates a close correspondence between behavioral and morphological specialization, but in other cases the behavioral trait appear unassociated with any morphological traits. Thus castorids may represent an excellent case study for examining the evolutionary relationships between morphology and behavior. Toward this end I have begun by examining patterns of behavioral evolution in a phylogenetic context. Within castorids two types of behavioral specialists were recognized, tooth-diggers and wood-cutters. These behavioral specialists were identified using morphological (quantitative comparative) and/or behavioral evidence. I conducted a series of cladistic analyses (36 taxa, 89 morphological characters) that varied in the exclusion and inclusion of various taxa and character weighting regimes. Mapping characters onto all the most parsimonious cladograms yielded similar patterns of behavioral evolution. The results suggest that fossoriality arose primitively within castorids, whereas tooth-digging, a fossorial specialization, arose at least twice. Assuming parsimony, wood-cutting appears to have evolved only once from a morphologically unspecialized, common ancestor that lived at least 20 million years ago. At least six genera descended from this ancestor, suggesting that the acquisition of wood-cutting behavior may have been associated with an adaptive radiation. Supported by NSF DIG grant IBN-0073119.

Fossilized cut wood found associated with the skeletal remains of the Pliocene beaver, Dipoides, provides the first evidence of wood-cutting behavior in an extinct beaver. Unlike the modern beaver, Castor, the lower incisor tips of Dipoides are not squared-off in anterior view, but are rounded. In addition, Dipoides is approximately 1/3 smaller than Castor, and is likely to have a lower maximum bite force. The aim of this study was to compare the woodcutting performance of these two incisor shapes. Models of the lower incisors were made in steel and honed to reflect incisor sharpness measurements. The models were of similar widths to one another, and were intermediate in size between the lower incisors of adult Dipoides and Castor. I investigated cutting performance of the models using a motorized test apparatus, fitted with a dynamometer for measuring cutting forces. Fixed parameters, defining the relative orientation of the wood sample and model, were derived from behavioral observations. Parameters that varied were wood-type (sweetgum, white oak, redcedar or pine), and width of cut. Analysis of 257 cutting events found that for cuts of similar size average peak cutting-forces were significantly higher for the Castor-tooth model than for the Dipoides tooth-model. However, unlike the Castor-tooth model, the Dipoides-tooth model was often unable to complete wide cuts in most wood samples (redcedar excepted). Therefore, although cutting with the Dipoides model is generally more efficient than the Castor-tooth model, in terms of force requirements, its cuts are usually smaller. The possibility that rounded incisors might promote specialization on certain trees is also discussed. I thank North Carolina University, and NSF (DIG-0073119).

The repeated returns of vertebrates to the marine ecosystems since the Triassic serve as an evolutionary model to understand macroevolutionary change. Here we investigate the effects of the land-to-sea transition on disparity and constraint of the vertebral column in aquatic carnivorans (Carnivora; Pinnipedia) to assess how their functional diversity and evolutionary innovations influenced major radiations of crown pinnipeds. We use three-dimensional geometric morphometrics and multivariate analysis for high-dimensional data under a phylogenetic framework to quantify vertebral size and shape in living and extinct pinnipeds. Our analysis demonstrates an important shift in vertebral column evolution by 10-12 million years ago, from an unconstrained to a constrained evolutionary scenario, a point of time that coincides with the major radiation of crown pinnipeds. Moreover, we also demonstrate that the axial skeleton of phocids and otariids followed a different path of morphological evolution that was probably driven by their specialized locomotor strategies. Despite this, we found a significant effect of habitat preference (coastal versus pelagic) on vertebral morphology of crown taxa regardless of the family they belong. In summary, our analysis provides insights into how the land-to-sea transition influenced the complex evolutionary history of pinniped vertebral morphology.

Secondary aquatic vertebrates exhibit a diversity of swimming modes that use paired limbs and/or the tail. Various secondarily aquatic tetrapod clades, including amphibians, reptiles, and mammals use transverse undulations or oscillations of the tail for swimming. These movements have often been classified according to a kinematic gradient that was established for fishes but may not be appropriate to describe the swimming motions of tetrapods. To understand the evolution of movements and design of the tail in aquatic tetrapods, we categorize the types of tails used for swimming and examine swimming kinematics and hydrodynamics. From a foundation of a narrow, elongate ancestral tail, the tails used for swimming by aquatic tetrapods are classified as tapered, keeled, paddle, and lunate. Tail undulations are associated with tapered, keeled, and paddle tails for a diversity of taxa. Propulsive undulatory waves move down the tail with increasing amplitude toward the tail tip, while moving posteriorly at a velocity faster than the anterior motion of the body indicating that the tail is used for thrust generation. Aquatic propulsion is associated with the transfer of momentum to the water from the swimming movements of the tail, particularly at the trailing edge. The addition of transverse extensions and flattening of the tail increases the mass of water accelerated posteriorly and affects vorticity shed into the wake for more aquatically adapted animals. Digital Particle Image Velocimetry reveals that the differences were exhibited in the vortex wake between the morphological and kinematic extremes of the alligator with a tapering undulating tail and the dolphin with oscillating wing-like flukes that generate thrust. In addition to exploring the relationship between the shape of undulating tails and the swimming performance across aquatic tetrapods, the role of tail reduction or loss of a tail in aquatic-tetrapod swimming was also explored. For aquatic tetrapods, the reduction would have been due to factors including locomotor and defensive specializations and phylogenetic and physiological constraints. Possession of a thrust-generating tail for swimming, or lack thereof, guided various lineages of secondarily aquatic vertebrates into different evolutionary trajectories for effective aquatic propulsion (i.e., speed, efficiency, and acceleration)., (© The Author(s) 2021. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology.)

Modern beavers (Castor) are prolific ecosystem engineers and dramatically alter the landscape through tree harvesting and dam building. Little is known, however, about the evolutionary drivers of their woodcutting behaviour. Here we investigate if early woodcutting behaviour in Castoridae was driven by nutritional needs. We measured stable carbon and nitrogen isotopes (δ 13 C and δ 15 N) of coeval subfossil plants and beaver collagen (Dipoides sp.) from the Early Pliocene, High Arctic Beaver Pond fossil locality (Ellesmere Island), in order to reconstruct Dipoides sp. diet. Isotopic evidence indicates a diet of woody plants and freshwater macrophytes, supporting the hypothesis that this extinct semiaquatic beaver engaged in woodcutting behaviour for feeding purposes. In a phylogenetic context, the isotopic evidence implies that woodcutting and consumption of woody plants can be traced back to a small-bodied, semiaquatic Miocene castorid, suggesting that beavers have been consuming woody plants for over 20 million years. We propose that the behavioural complex (swimming, woodcutting, and consuming woody plants) preceded and facilitated the evolution of dam building. Dam building and food caching behaviours appear to be specializations for cold winter survival and may have evolved in response to late Neogene northern cooling.

Proteomic analyses of ancient remains are increasing in number and offer great potential to recover phylogenetic information on extinct animals beyond the reach of ancient DNA, but limitations in proteomic techniques remain unclear. Here we carry out LC-MS/MS sequence analysis of a ~3.5 million year old giant camel specimen from Nunavut along with the younger Pleistocene remains of the Yukon giant camel (c.f. Paracamelus) and the western camel (Camelops hesternus) for comparison with complete sequences to both extant camels (Bactrian and Dromedary) and the alpaca. Although not complete (~75-80% sequence coverage), no amino acid sequence differences were confidently observed between the giant camels and the extant Dromedary, indicative of a closer relationship than that of the extant Bactrian lineage. However, multiple amino acid changes were observed for the western camel (Camelops) collagen sequence, placing it as a sister group to these members of the Camelini tribe consistent recent ancient DNA analyses. Although this supports a role for the sequencing of ancient collagen in the understanding of vertebrate evolution, these analyses highlight the limitations in phylogenetic reconstructions based on partial sequence data retrieved from proteomic analyses, particularly, the impact of omitting even only a single peptide on the resulting tree topology. The presence of other non-collagenous proteins, such as biglycan and PEDF, indicates a further resource for phylogenetic information, but none more promising than the degraded camel albumin seemingly observed in the Pliocene specimen. SIGNIFICANCE: As proteomics is becoming more frequently used in the study of ancient proteins, an emerging field known as 'palaeoproteomics' (or 'paleoproteomics'), understanding the limitations of the technique is essential. Here, through the study of the oldest undisputed collagen sequences obtained from proteomics, we confirm that some peptides following diagenetic modifications of tryptic sites are no longer matched with standard searches, but can be matched with Error Tolerant searches. We also demonstrate the ability to retrieve phylogenetic information consistent with that of ancient DNA methods, but that with the omission of only one or more key peptides, the inferred evolutionary relationships change. This is a significant finding for the field of palaeoproteomics implying a need for better understanding the particular composition of the partial sequences retrieved from proteomic analyses., (Copyright © 2018. Published by Elsevier B.V.)

The skeletal remains of a small bear (Protarctos abstrusus) were collected at the Beaver Pond fossil site in the High Arctic (Ellesmere I., Nunavut). This mid-Pliocene deposit has also yielded 12 other mammals and the remains of a boreal-forest community. Phylogenetic analysis reveals this bear to be basal to modern bears. It appears to represent an immigration event from Asia, leaving no living North American descendants. The dentition shows only modest specialization for herbivory, consistent with its basal position within Ursinae. However, the appearance of dental caries suggest a diet high in fermentable-carbohydrates. Fossil plants remains, including diverse berries, suggests that, like modern northern black bears, P. abstrusus may have exploited a high-sugar diet in the fall to promote fat accumulation and facilitate hibernation. A tendency toward a sugar-rich diet appears to have arisen early in Ursinae, and may have played a role in allowing ursine lineages to occupy cold habitats.

The evolution of robust jaws, hypsodont teeth, and large chewing muscles among grazing ruminants is a quintessential example of putative morphological adaptation. However, the degree of correlated evolution (i.e., to what extent the grazer feeding apparatus represents an evolutionary module), especially of soft and hard tissues, remains poorly understood. Recent generation of large datasets and phylogenetic information has made testing hypotheses of correlated evolution possible. We, therefore, test for correlated evolution among various traits of the ruminant masticatory apparatus including tooth crown height, jaw robustness, chewing muscle size, and characters of the molar occlusal surfaces, using phylogenetic and nonphylogenetic comparative methods as well as phylogenetic evolutionary model selection. We find that the large masseter muscles of grazing ruminants evolved with the inclusion of grass in the diet, an increase in the proportion of occlusal enamel bands oriented parallel to the chewing stroke, and possibly hypsodonty. We suggest that the masseter evolved under two evolutionary regimes: i) selection for higher masticatory forces during chewing and ii) flattening of the tooth profile, which resulted in reduced tooth guidance and, thus, a requirement for more chewing muscle activity during each chewing stroke, in agreement with previous research. The linear jaw metrics (depth of the mandibular angle, mandibular angle width, and length of the superficial masseteric scar) all show correlated evolution with hypsodonty and the proportion of enamel bands oriented parallel to the chewing stroke. We suggest that changes in the shape of the mandible represent the combined effects of selection for a reorientation of the chewing stroke, so as to emphasize horizontal translation of the teeth, and accommodation of high-crowned teeth. Our analyses show that the ruminant feeding apparatus is an evolutionary mosaic with its various components showing both correlated and independent evolution., (Copyright © 2014 Wiley Periodicals, Inc.)

Spatial diversity patterns are thought to be driven by climate-mediated processes. However, temporal patterns of community composition remain poorly studied. We provide two complementary analyses of North American mammal diversity, using (i) a paleontological dataset (2077 localities with 2493 taxon occurrences) spanning 21 discrete subdivisions of the Cenozoic based on North American Land Mammal Ages (36 Ma--present), and (ii) climate space model predictions for 744 extant mammals under eight scenarios of future climate change. Spatial variation in fossil mammal community structure (β diversity) is highest at intermediate values of continental mean annual precipitation (MAP) estimated from paleosols (∼ 450 mm/year) and declines under both wetter and drier conditions, reflecting diversity patterns of modern mammals. Latitudinal gradients in community change (latitudinal turnover gradients, aka LTGs) increase in strength through the Cenozoic, but also show a cyclical pattern that is significantly explained by MAP. In general, LTGs are weakest when continental MAP is highest, similar to modern tropical ecosystems in which latitudinal diversity gradients are weak or undetectable. Projections under modeled climate change show no substantial change in β diversity or LTG strength for North American mammals. Our results suggest that similar climate-mediated mechanisms might drive spatial and temporal patterns of community composition in both fossil and extant mammals. We also provide empirical evidence that the ecological processes on which climate space models are based are insufficient for accurately forecasting long-term mammalian response to anthropogenic climate change and inclusion of historical parameters may be essential.

Sexual selection is one of the earliest areas of interest in evolutionary biology. And yet, the evolutionary history of sexually dimorphic traits remains poorly characterized for most vertebrate lineages. Here, we report on evidence for the early evolution of dimorphism within a model mammal group, the pinnipeds. Pinnipeds show a range of sexual dimorphism and mating systems that span the extremes of modern mammals, from monomorphic taxa with isolated and dispersed mating to extreme size dimorphism with highly ordered polygynous harem systems. In addition, the degree of dimorphism in pinnipeds is closely tied to mating system, with strongly dimorphic taxa always exhibiting a polygynous system, and more monomorphic taxa possessing weakly polygynous systems. We perform a comparative morphological description, and provide evidence of extreme sexual dimorphism (similar to sea lions), in the Miocene-aged basal pinniped taxon Enaliarctos emlongi. Using a geometric morphometric approach and combining both modern and fossil taxa we show a close correlation between mating system and sex-related cranial dimorphism, and also reconstruct the ancestral mating system of extant pinnipeds as highly polygynous. The results suggest that sexual dimorphism and extreme polygyny in pinnipeds arose by 27 Ma, in association with changing climatic conditions., (© 2014 The Author(s). Evolution © 2014 The Society for the Study of Evolution.)

The mid-Pliocene was a global warm period, preceding the onset of Quaternary glaciations. Here we use cosmogenic nuclide dating to show that a fossiliferous terrestrial deposit that includes subfossil trees and the northern-most evidence of Pliocene ice wedge casts in Canada's High Arctic (Ellesmere Island, Nunavut) was deposited during the mid-Pliocene warm period. The age estimates correspond to a general maximum in high latitude mean winter season insolation, consistent with the presence of a rich, boreal-type forest. Moreover, we report that these deposits have yielded the first evidence of a High Arctic camel, identified using collagen fingerprinting of a fragmentary fossil limb bone. Camels originated in North America and dispersed to Eurasia via the Bering Isthmus, an ephemeral land bridge linking Alaska and Russia. The results suggest that the evolutionary history of modern camels can be traced back to a lineage of giant camels that was well established in a forested Arctic.

The highly specialized tooth morphology and arrangement of the dental battery of hadrosaurids has led to much speculation surrounding the chewing mechanics of this successful group of herbivorous dinosaurs. Pleurokinesis, a long established hypothesis explaining the ornithopod chewing mechanism, proposes a transverse power stroke in hadrosaurids that was accommodated by vertical adduction of the mandible, lateral rotation of the maxilla at the maxilla-premaxilla joint, lateral rotation of the jugal-maxilla complex at its contact with the lacrimal, and posterolateral rotation of the quadrate at its contact with the squamosal. A secondary series of movements were also thought to have occurred as a consequence of these primary movements. In this article, the intracranial joint morphology is described for both Brachylophosaurus canadensis and Edmontosaurus regalis and their permissive kinematics are established. Based on this evidence, the movements associated with pleurokinesis are not accommodated in these hadrosaurine dinosaurs. Rather, the movements that seem most likely to have produced the observed dental wear patterns are those associated with the mandible about the jaw joint. The structure of this joint appears well-suited to have accommodated some translation as well as rotation of the mandible about the quadrate condyle. Three-dimensional modeling of the alternate mandibular movements reveals that not all the combined labiolingual width of the lingual and buccal facets of the tooth row was involved in the power stroke. Rather, limits on the degree of mandibular long axis rotation suggest that only the lingual facet and the more medial portion of the buccal facet were utilized., (Copyright © 2012 Wiley Periodicals, Inc.)

The extant beaver, Castor, has played an important role shaping landscapes and ecosystems in Eurasia and North America, yet the origins and early evolution of this lineage remain poorly understood. Here we use a geometric morphometric approach to help re-evaluate the phylogenetic affinities of a fossil skull from the Late Miocene of China. This specimen was originally considered Sinocastor, and later transferred to Castor. The aim of this study was to determine whether this form is an early member of Castor, or if it represents a lineage outside of Castor. The specimen was compared to 38 specimens of modern Castor (both C. canadensis and C. fiber) as well as fossil specimens of C. fiber (Pleistocene), C. californicus (Pliocene) and the early castorids Steneofiber eseri (early Miocene). The results show that the specimen falls outside the Castor morphospace and that compared to Castor, Sinocastor possesses a: 1) narrower post-orbital constriction, 2) anteroposteriorly shortened basioccipital depression, 3) shortened incisive foramen, 4) more posteriorly located palatine foramen, 5) longer rostrum, and 6) longer braincase. Also the specimen shows a much shallower basiocciptal depression than what is seen in living Castor, as well as prominently rooted molars. We conclude that Sinocastor is a valid genus. Given the prevalence of apparently primitive traits, Sinocastor might be a near relative of the lineage that gave rise to Castor, implying a possible Asiatic origin for Castor.

Modern pinnipeds (seals, sea lions and the walrus) are semi-aquatic, generally marine carnivores the limbs of which have been modified into flippers. Recent phylogenetic studies using morphological and molecular evidence support pinniped monophyly, and suggest a sister relationship with ursoids (for example bears) or musteloids (the clade that includes skunks, badgers, weasels and otters). Although the position of pinnipeds within modern carnivores appears moderately well resolved, fossil evidence of the morphological steps leading from a terrestrial ancestor to the modern marine forms has been weak or contentious. The earliest well-represented fossil pinniped is Enaliarctos, a marine form with flippers, which had appeared on the northwestern shores of North America by the early Miocene epoch. Here we report the discovery of a nearly complete skeleton of a new semi-aquatic carnivore from an early Miocene lake deposit in Nunavut, Canada, that represents a morphological link in early pinniped evolution. The new taxon retains a long tail and the proportions of its fore- and hindlimbs are more similar to those of modern terrestrial carnivores than to modern pinnipeds. Morphological traits indicative of semi-aquatic adaptation include a forelimb with a prominent deltopectoral ridge on the humerus, a posterodorsally expanded scapula, a pelvis with relatively short ilium, a shortened femur and flattened phalanges, suggestive of webbing. The new fossil shows evidence of pinniped affinities and similarities to the early Oligocene Amphicticeps from Asia and the late Oligocene and Miocene Potamotherium from Europe. The discovery suggests that the evolution of pinnipeds included a freshwater transitional phase, and may support the hypothesis that the Arctic was an early centre of pinniped evolution.

Herbivores can increase their digestion rate by mechanically reducing particle size through oral trituration. Groups of terrestrial vertebrates with the greatest capacity to reduce tough plant foods orally are also the most abundant and diverse, as exemplified by ornithopod dinosaurs during the Mesozoic and extant artiodactyl and perissodactyl mammals. Thus, the effective oral processing of high-fibre plant material seems to represent an evolutionary innovation of both functional and macroevolutionary significance. However, evidence for oral processing is poorly documented in the fossil record, especially during the initial stages of terrestrial vertebrate diversification. Here we report on the basal anomodont Suminia getmanovi, the only known Palaeozoic vertebrate in which unequivocal specializations in its cranium and teeth for high-fibre herbivory are well preserved. We propose that the capacity to comminute tough plant foods was critical to the diversification of anomodonts, the most diverse, widely dispersed and abundant group of Palaeozoic terrestrial vertebrates, and to the onset of modern terrestrial ecosystems.