Neural canal ridges: A novel osteological correlate of postcranial neuroanatomy in dinosaurs.

In this article, we document the widespread presence of bony ridges in the neural canals of non-avian dinosaurs, including a wide diversity of sauropods, two theropods, a thyreophoran, and a hadrosaur. These structures are present only in the caudal vertebrae. They are anteroposteriorly elongate, found on the lateral walls of the canal, and vary in size and position both taxonomically and serially. Similar bony projections into the neural canal have been identified in extant teleosts, dipnoans, and urodelans, in which they are recognized as bony spinal cord supports. In most non-mammals, the dura mater that surrounds the spinal cord is fused to the periosteum of the neural canal, and the denticulate ligaments that support the spinal cord can pass through the dura and periosteum to anchor directly to bone. The function of these structures in dinosaurs remains uncertain, but in sauropods they might have stabilized the spinal cord during bilateral movement of the tail and use of the tail as a weapon. Of broader significance, this study emphasizes that important new discoveries at the gross anatomical level can continue to be made in part by closely examining previously overlooked features of known specimens.

A giant tyrannosaur from the Campanian-Maastrichtian of southern North America and the evolution of tyrannosaurid gigantism.

Tyrannosaurid dinosaurs dominated as predators in the Late Cretaceous of Laurasia, culminating in the evolution of the giant Tyrannosaurus rex, both the last and largest tyrannosaurid. Where and when Tyrannosaurini (T. rex and kin) originated remains unclear. Competing hypotheses place tyrannosaurin origins in Asia, or western North America (Laramidia). We report a new tyrannosaurin, Tyrannosaurus mcraeensis, from the Campanian-Maastrichtian Hall Lake Formation of New Mexico, based on a fossil previously referred to T. rex. T. mcraeensis predates T. rex by ~ 6-7 million years, yet rivaled it in size. Phylogenetic analysis recovers T. mcraeensis as sister to T. rex and suggests Tyrannosaurini originated in southern Laramidia. Evolution of giant tyrannosaurs in southern North America, alongside giant ceratopsians, hadrosaurs, and titanosaurs suggests large-bodied dinosaurs evolved at low latitudes in North America.

Introduction to Dinosaurs: New Ideas from Old Bones papers honoring Peter Dodson.

This special volume of The Anatomical Record honors the life-long commitment to anatomy and paleontology by Professor Peter Dodson (Emeritus, University of Pennsylvania). Peter's legacy is not only rooted in his own research interests, but also in the wealth of former students he mentored over his career, many of whom have made their own new contributions to the fields of anatomy and paleontology through original scientific investigation. Within these 18 scientific papers, which cover multiple taxa, multiple continents, and multiple methodologies, each of the contributors brought to this volume their own unique work that can be traced back to some form of inspiration by the honoree.

Evolutionary insights from an anatomical network analysis of the hyolaryngeal apparatus in extant archosaurs (birds and crocodilians).

Adaptive radiation of archosaurs, represented by crocodilians, non-avian dinosaurs, and birds, since the Mesozoic has been studied mainly based on their major skeletal elements (skull, vertebrae, and limbs). However, little is known about the evolution of their hyolaryngeal apparatus, which is involved with feeding, respiration, and vocalization, because of poor fossil preservation and the difficulty in determining the musculoskeletal homology of the apparatus. Network analysis is a framework to quantitatively characterize the topological organization of anatomical structures for comparing structural integration and modularity regardless of ambiguous homology. Herein, we modeled the musculoskeletal system of hyolarynx in six species of extant archosaurs and its sister-taxon turtle, and conducted a network analysis using network parameters, modular partition, and bone centrality in a phylogenetic framework. The network parameters reveal that ancestral archosaurs have reduced the numbers of elements and links and acquired complex networks as a whole domain with strong modularity in the hyolarynx. Furthermore, the modular partition and centrality reveal that the hyoids are highly evolvable, while the larynx is constrained and less evolvable. The archosaur hyolarynx exhibits different evolutionary trends: crocodilians with the system integration, basihyal simplification, and ceratobranchial centralization; and birds with the simplicity, weak integration, and modularity of the hyolarynx, laryngeal integration with cricoid centrality, and tongue-module expansion with the acquisition of paraglossal. Four hyolaryngeal bones (ceratobranchial, basihyal, paraglossal, and cricoid) have played important roles in archosaur evolution, and their fossil records are keys to understanding the two major archosaur lineages toward crocodilians and birds.

Description of the first definitive Corythosaurus (Dinosauria, Hadrosauridae) specimens from the Judith River Formation in Montana, USA and their paleobiogeographical significance.

Despite the long history of research in the late Campanian Judith River Formation in northern Montana, most of the vertebrate fossils are represented by fragmentary remains, making precise taxonomic identifications difficult. Contrary to this, the partially contemporaneous Dinosaur Park Formation, Alberta, Canada is known for its tremendous fossil preservation, permitting rigorous studies of dinosaur diversity, evolution, and biostratigraphy. Hadrosaurids comprise one of the most abundant dinosaur clades in the Dinosaur Park Formation, but taxonomic affinities of hadrosaurid specimens remain poorly understood in the Judith River Formation. Corythosaurus is the most common hadrosaurid in the Dinosaur Park Formation and, to date, has been restricted to this formation. This study reports the first definitive Corythosaurus specimens from the Judith River Formation, which were discovered on two private ranches in northern Montana. The attribution of the most complete skeleton to Corythosaurus is indicated by: wide crest-snout angle, presence of premaxilla-nasal fontanelle, dorsoventrally expanded nasal, laterally exposed ophthalmic canal of the laterosphenoid, and tall neural spines. A second specimen preserves a large ilium that can be positively identified as Corythosaurus based on its associated skull, which is now in private hands. The specimens were recovered from the Coal Ridge Member of the Judith River Formation, which is approximately time equivalent to the Dinosaur Park Formation. Thus, the discovery of Corythosaurus in the Judith River Formation extends the biogeographic range of this genus and establishes a framework for future interformational biostratigraphic studies of Late Cretaceous dinosaur faunas in North America.

Paleobiological inferences from paleopathological occurrences in the Arctic ceratopsian Pachyrhinosaurus perotorum.

As a key tool for understanding how animals lived in the past, paleopathology informs us about the lives and deaths of fossil animals. We identify paleopathologies within an assemblage of bones of the pachyrostran centrosaurine Pachyrhinosaurus perotorum, an Arctic ceratopsian. More than 1,000 bones of this dinosaur were collected from the Prince Creek Formation of North Slope, Alaska from fossil sites along the Colville River. Our survey shows the occurrence of paleopathology to be very low and comparable to other populations of horned dinosaurs from the lower latitudes, suggesting that the ancient Arctic environment did not impose intense hardships on these dinosaurs greater than in other environments, as expressed by paleopathological modification of the skeleton. This result may be due to the more equable mean annual temperatures in the Arctic region during the Cretaceous. Also of interest, the frequency of occurrence of paleopathology in the Arctic Pachyrhinosaurus population is very low compared to populations of fossil and historic quadrupedal artiodactyls that are recognized as long distance wanderers.

New therizinosaurid dinosaur from the marine Osoushinai Formation (Upper Cretaceous, Japan) provides insight for function and evolution of therizinosaur claws.

The record of therizinosaurs is rich in Asian countries such as Mongolia and China. Fragmentary therizinosaur specimens have been reported from the Lower and Upper Cretaceous deposits in Japan. One of these specimens, from the lower Campanian Osoushinai Formation in Nakagawa Town of Hokkaido Prefecture, was previously identified as a maniraptoran theropod dinosaur, possibly therizinosaur, but its taxonomic status remained unresolved. This study re-examines the specimen and provides a more detailed description and attempts to resolve its taxonomic status. Our study demonstrates that it is a new taxon, Paralitherizinosaurus japonicus gen. et sp. nov., because it shows a unique combination of characters in the metacarpal I and unguals. Our phylogenetic analysis places this new taxon within an unresolved clade of Therizinosauridae in the strict consensus tree. The 50% majority-rule consensus tree shows better resolution within Therizinosauridae, showing an unresolved monophyletic clade of Paralitherizinosaurus, Therizinosaurus, Suzhousaurus, and the Bissekty form. Geometric morphometric analysis suggests that Paralitherizinosaurus unguals most closely resemble Therizinosaurus unguals in being slender and has weak flexor tubercles. This study also shows an evolutionary trend in ungual shape, which associates a decrease in mechanical advantage, development of flexor tubercle, and hypothesized output (product of mechanical advantage and development of flexor tubercle) in derived therizinosaurs, supporting the hook-and-pull function of claws to bring vegetation to its mouth. Paralitherizinosaurus is the youngest therizinosaur from Japan and the first recovered from the marine deposits in Asia. This suggests a long temporal existence of therizinosaurs at the eastern edge of the Asian continent and adaptation of therizinosaurs to coastal environments.

A new basal hadrosaurid (Dinosauria: Ornithischia) from the latest Cretaceous Kita-ama Formation in Japan implies the origin of hadrosaurids.

Here we describe a partial hadrosaurid skeleton from the marine Maastrichtian Kita-ama Formation in Japan as a new taxon, Yamatosaurus izanagii gen. et sp. nov., based on unique characters in the dentition. Our phylogenetic analysis demonstrates Yamatosaurus izanagii belongs to Hadrosauridae, composed of Hadrosaurus foulkii + (Yamatosaurus izanagii + (Saurolophinae + Lambeosaurinae)). The coracoid lacks a biceps tubercle as in non-hadrosaurid hadrosauroids, suggesting its presence is a key feature for the clade of Saurolophinae and Lambeosaurinae. The evolutionary rates analysis further supports that shoulder and forelimb features, which are likely to have been involved in locomotion, were important for the early evolution of Hadrosauridae. Our biogeographic analyses show that basal hadrosaurids were widely distributed in Asia and Appalachia, that the clade of Saurolophinae and Lambeosaurinae originated in Asia, and that eastern Asia may have served as a refugium of relict hadrosauroid taxa such as Plesiohadros djadokhtaensis, Tanius sinensis, and Yamatosaurus izanagii during the Late Cretaceous. The contemporaneous occurrence of basal (Yamatosaurus izanagii) and derived (Kamuysaurus japonicus) hadrosaurids during the Maastrichtian in Japan is the first record in Asia. Because of the long geographical distance between these localities, they likely did not co-exist, but instead demonstrate some level of provinciality.

Differing trabecular bone architecture in dinosaurs and mammals contribute to stiffness and limits on bone strain.

The largest dinosaurs were enormous animals whose body mass placed massive gravitational loads on their skeleton. Previous studies investigated dinosaurian bone strength and biomechanics, but the relationships between dinosaurian trabecular bone architecture and mechanical behavior has not been studied. In this study, trabecular bone samples from the distal femur and proximal tibia of dinosaurs ranging in body mass from 23-8,000 kg were investigated. The trabecular architecture was quantified from micro-computed tomography scans and allometric scaling relationships were used to determine how the trabecular bone architectural indices changed with body mass. Trabecular bone mechanical behavior was investigated by finite element modeling. It was found that dinosaurian trabecular bone volume fraction is positively correlated with body mass similar to what is observed for extant mammalian species, while trabecular spacing, number, and connectivity density in dinosaurs is negatively correlated with body mass, exhibiting opposite behavior from extant mammals. Furthermore, it was found that trabecular bone apparent modulus is positively correlated with body mass in dinosaurian species, while no correlation was observed for mammalian species. Additionally, trabecular bone tensile and compressive principal strains were not correlated with body mass in mammalian or dinosaurian species. Trabecular bone apparent modulus was positively correlated with trabecular spacing in mammals and positively correlated with connectivity density in dinosaurs, but these differential architectural effects on trabecular bone apparent modulus limit average trabecular bone tissue strains to below 3,000 microstrain for estimated high levels of physiological loading in both mammals and dinosaurs.

The first juvenile dromaeosaurid (Dinosauria: Theropoda) from Arctic Alaska.

Compared to the osteological record of herbivorous dinosaurs from the Late Cretaceous Prince Creek Formation of northern Alaska, there are relatively fewer remains of theropods. The theropod record from this unit is mostly comprised of isolated teeth, and the only non-dental remains known can be attributed to the troodontid cf. Troodon and the tyrannosaurid Nanuqsaurus. Thus far, the presence of members of Dromaeosauridae has been limited to isolated teeth. Here we describe a symphyseal portion of a small dentary with two ziphodont teeth. Based on tooth shape, denticle morphology, and the position of the Meckelian groove, we attribute this partial dentary to a saurornitholestine dromaeosaurid. The fibrous bone surface, small size, and higher number of mesial denticles compared to distal ones point to a juvenile growth stage for this individual. Multivariate comparison of theropod teeth morphospace by means of principal component analysis reveals an overlap between this dentary and Saurornitholestinae dromaeosaurid morphospace, a result supported by phylogenetic analyses. This is the first confirmed non-dental fossil specimen from a member of Dromaeosauridae in the Arctic, expanding on the role of Beringia as a dispersal route for this clade between Asia and North America. Furthermore, the juvenile nature of this individual adds to a growing body of data that suggests Cretaceous Arctic dinosaurs of Alaska did not undergo long-distance migration, but rather they were year-round residents of these paleopolar latitudes.

Re-examination of the cranial osteology of the Arctic Alaskan hadrosaurine with implications for its taxonomic status.

Hadrosaurid fossils from the Liscomb Bonebed (Prince Creek Formation, North Slope, Alaska) were the first dinosaur bones discovered from the Arctic. While the Prince Creek Formation hadrosaurids were long identified as Edmontosaurus, a member of the sub-clade Hadrosaurinae, they were recently assigned to a newly-erected taxon, Ugrunaaluk kuukpikensis. However, taxonomic status of the new taxon is ambiguous largely due to the immature nature of the specimens upon which it was based. Here we reexamine cranial elements of the Prince Creek Formation hadrosaurine in order to solve its taxonomic uncertainties. The Prince Creek Formation hadrosaurine possesses a short dorsolateral process of the laterosphenoid, one of the diagnostic characters of Edmontosaurus. The Prince Creek Formation hadrosaurine also shows affinity to Edmontosaurus regalis in the presence of a horizontal shelf of the jugal. Our morphological comparisons with other North American Edmontosaurus specimens and our phylogenetic analyses demonstrate that the Prince Creek Formation hadrosaurine should be re-assigned to Edmontosaurus. Because the Prince Creek Formation Edmontosaurus shows differences with lower latitude Edmontosaurus in a dorsoventrally short maxilla, presence of a secondary ridge on the dentary teeth, and the absence of the transverse ridge between basipterygoid processes of the basisphenoid, we consider that the Prince Creek Formation Edmontosaurus should be regarded as Edmontosaurus sp. until further discoveries of mature hadrosaurines from the Prince Creek Formation Bonebed and/or equivalently juvenile Edmontosaurus specimens from the lower latitudes allow direct comparisons. The retention of the Prince Creek Formation hadrosaurine as Edmontosaurus re-establishes a significant latitudinal distribution for this taxon. Despite the large latitudinal distribution of the taxon, the morphological disparity of Edmontosaurus is small within Hadrosaurinae. The small morphological disparity may be related to the relatively low latitudinal temperature gradient during the latest Cretaceous compared to present day, a gradient which might not have imposed significant pressure for much morphological adaptations across a broad latitudinal range.

Filling in Gaps in the Ceratopsid Histologic Database: Histology of Two Basal Centrosaurines and an Assessment of the Utility of Rib Histology in the Ceratopsidae.

Bone histology grants substantial insight into the growth and biology of fossil vertebrates. Many of the major non-avian dinosaurian clades have been extensively sampled for bone histologic data allowing reconstruction of their growth as well as the assessment of the evolution of growth changes along phylogenies. However, horned ceratopsians are poorly represented in paleohistologic studies. Further, the ceratopsian taxa that have been examined are unevenly sampled phylogenetically with very basal forms and highly derived forms making up the majority of studied taxa. In order to rectify this, we have histologically sampled Avaceratops from Montana and Yehuecauhceratops from northern Mexico to assess how mid-sized basal centrosaurines grew relative to more basal and derived forms. Based on results from these taxa, basal centrosaurines present a mosaic of growth characters intermediate between those seen in basal ceratopsians and more derived centrosaurines. Further, Yehuecauhceratops has many lines of arrested growth preserved, suggesting that the large number of lines of arrested growth found in a high-latitude Pachyrhinosaurus specimen may be a result of phylogeny rather than geography. Since lines of arrested growth are not preserved in long bones of many ceratopsians, especially chasmosaurines, we also histologically sampled ribs of Avaceratops and Pachyrhinosaurus. However, the largest ribs were highly remodeled obscuring lines of arrested growth, making it unlikely that rib histology will clarify growth trends in ceratopsians. These centrosaurines add to the growing ceratopsian histological database and demonstrate that basal centrosaurines grew in a manner intermediate between non-ceratopsid taxa and derived centrosaurines. Anat Rec, 303:935-948, 2020. © 2019 Wiley Periodicals, Inc.

Author Correction: A New Hadrosaurine (Dinosauria: Hadrosauridae) from the Marine Deposits of the Late Cretaceous Hakobuchi Formation, Yezo Group, Japan.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Dinosaur ichnology and sedimentology of the Chignik Formation (Upper Cretaceous), Aniakchak National Monument, southwestern Alaska; Further insights on habitat preferences of high-latitude hadrosaurs.

While there are now numerous records of dinosaurs from Cretaceous rocks around the state of Alaska, very few fossil records of terrestrial vertebrates are known from the Mesozoic rocks of the southwestern part of the state. Here we report the new discovery of extensive occurrences of dinosaur tracks from Aniakchak National Monument of the Alaska Peninsula. These tracks are in the Late Cretaceous (Maastrichtian) Chignik Formation, a cyclic sequence of rocks, approximately 500-600 m thick, representing shallow marine to nearshore marine environments in the lower part and continental alluvial coastal plain environments in the upper part of the section. These rocks are part of the Peninsular Terrane and paleomagnetic reconstructions based on the volcanic rocks of this terrane suggest that the Chignik Formation was deposited at approximately its current latitude which is almost 57° N. Recent field work in Aniakchak National Monument has revealed over 75 new track sites, dramatically increasing the dinosaur record from the Alaska Peninsula. Most of the combined record of tracks can be attributed to hadrosaurs, the plant-eating duck-billed dinosaurs. Tracks range in size from those made by full-grown adults to juveniles. Other tracks can be attributed to armored dinosaurs, meat-eating dinosaurs, and two kinds of fossil birds. The track size of the predatory dinosaur suggests a body approximately 6-7 m long, about the estimated size of the North Slope tyrannosaurid Nanuqsaurus. The larger bird tracks resemble Magnoavipes denaliensis previously described from Denali National Park, while the smaller bird tracks were made by a bird about the size of a modern Willet. Previous interdisciplinary sedimentologic and paleontologic work in the correlative and well-known dinosaur bonebeds of the Prince Creek Formation 1400km-1500km further north in Alaska suggested that high-latitude hadrosaurs preferred distal coastal plain or lower delta plain habitats. The ichnological record being uncovered in the Chignik Formation of southwestern Alaska is showing that the hadrosaur tracks here were also made in distal coastal and delta plain conditions. This similarity may corroborate the habitat preference model for Cretaceous high-latitude dinosaurs proposed for the data gathered from the Prince Creek Formation, and may indicate that at least Beringian hadrosaurids had similar habitat preferences regardless of latitude.

A New Hadrosaurine (Dinosauria: Hadrosauridae) from the Marine Deposits of the Late Cretaceous Hakobuchi Formation, Yezo Group, Japan.

A nearly complete skeleton of a new hadrosaurid, Kamuysaurus japonicus gen. et sp. nov., was discovered from the outer shelf deposits of the Upper Cretaceous Hakobuchi Formation of the Yezo Group in Hobetsu area of Mukawa town in Hokkaido, Japan. Kamuysaurus belongs to the sub-clade of Hadrosaurinae, Edmontosaurini, and forms a monophyly with Laiyangosaurus and Kerberosaurus from the northern Far East. Kamuysaurus has a long anterior platform for the nasofrontal sutural surface, which may indicate the presence of a small supracranial crest, similar to a sub-adult form of Brachylophosaurus based on the extension of the nasofrontal sutural surface. The Dispersal Extinction Cladogenesis analysis with the 50% Majority Rule consensus tree suggests that the clade of Kamuysaurus, Laiyangosaurus, and Kerberosaurus may have dispersed into Asia prior to the late Campanian and the potential endemism of this clade during the late Campanian and early Maastrichtian in the northern Far East. The results of both Dispersal Extinction Cladogenesis and Ancestral State Reconstruction analyses imply that the marine-influenced environment in North America during the Campanian may have played an important role for the hadrosaurid diversification in its early evolutionary history.

The First Definite Lambeosaurine Bone From the Liscomb Bonebed of the Upper Cretaceous Prince Creek Formation, Alaska, United States.

The Prince Creek Formation of Alaska, a rock unit that represents lower coastal plain and delta deposits, is one of the most important formations in the world for understanding vertebrate ecology in the Arctic during the Cretaceous. Here we report on an isolated cranial material, supraoccipital, of a lambeosaurine hadrosaurid from the Liscomb Bonebed of the Prince Creek Formation. The lambeosaurine supraoccipital has well-developed squamosal bosses and a short sutural surface with the exoccipital-opisthotic complex, and is similar to lambeosaurine supraoccipitals from the Dinosaur Park Formation in having anteriorly positioned squamosal bosses. Affinities with Canadian lambeosaurines elucidate more extensive faunal exchange between the Arctic and lower paleolatitudes which was previously suggested by the presence of Edmontosaurus, Pachyrhinosaurus, tyrannosaurids, and troodontids in both regions. The presence of one lambeosaurine and nine hadrosaurine supraoccipitals in the Liscomb Bonebed suggests hadrosaurine dominated faunal structure as in the Careless Creek Quarry of the USA that was also deposited under a near-shore environment. It differs from the lambeosaurine dominant structures of localities in Russia and China interpreted as inland environments. This may suggest that lambeosaurines had less preference for near-shore environments than hadrosaurines in both Arctic and lower paleolatitudes.

An unusual association of hadrosaur and therizinosaur tracks within Late Cretaceous rocks of Denali National Park, Alaska.

We report details of a unique association of hadrosaur and therizinosaur tracks found in the Late Cretaceous lower Cantwell Formation, Denali National Park, central Alaska Range, Alaska. This rock unit is now well-documented as a source of thousands of fossil footprints of vertebrates such as fishes, pterosaurs, and avialan and non-avialan dinosaurs. The lower Cantwell Formation in this area consists of numerous fining-upward successions of conglomerates and pebbly sandstones, cross-stratified and massive sandstones, interbedded sandstones and siltstones, organic-rich siltstones and shales, and rare, thin, bentonites, typically bounded by thin coal seams, and it contains a diverse fossil flora. We report the first North American co-occurrence of tracks attributable to hadrosaurs and therizinosaurs in the lower Cantwell Formation. Although previously un-reported in North America, this association of hadrosaur and therizinosaur tracks is more characteristic of the correlative Nemegt Formation in central Asia, perhaps suggesting that parameters defining the continental ecosystem of central Asia were also present in this part of Alaska during the Latest Cretaceous.

Lizards ran bipedally 110 million years ago.

Four heteropod lizard trackways discovered in the Hasandong Formation (Aptian-early Albian), South Korea assigned to Sauripes hadongensis, n. ichnogen., n. ichnosp., which represents the oldest lizard tracks in the world. Most tracks are pes tracks (N = 25) that are very small, average 22.29 mm long and 12.46 mm wide. The pes tracks show "typical" lizard morphology as having curved digit imprints that progressively increase in length from digits I to IV, a smaller digit V that is separated from the other digits by a large interdigital angle. The manus track is 19.18 mm long and 19.23 mm wide, and shows a different morphology from the pes. The predominant pes tracks, the long stride length of pes, narrow trackway width, digitigrade manus and pes prints, and anteriorly oriented long axis of the fourth pedal digit indicate that these trackways were made by lizards running bipedally, suggesting that bipedality was possible early in lizard evolution.

A diminutive new tyrannosaur from the top of the world.

Tyrannosaurid theropods were dominant terrestrial predators in Asia and western North America during the last of the Cretaceous. The known diversity of the group has dramatically increased in recent years with new finds, but overall understanding of tyrannosaurid ecology and evolution is based almost entirely on fossils from latitudes at or below southern Canada and central Asia. Remains of a new, relatively small tyrannosaurine were recovered from the earliest Late Maastrichtian (70-69Ma) of the Prince Creek Formation on Alaska's North Slope. Cladistic analyses show the material represents a new tyrannosaurine species closely related to the highly derived Tarbosaurus+Tyrannosaurus clade. The new taxon inhabited a seasonally extreme high-latitude continental environment on the northernmost edge of Cretaceous North America. The discovery of the new form provides new insights into tyrannosaurid adaptability, and evolution in an ancient greenhouse Arctic.