The dinosaurs that weren't: osteohistology supports giant ichthyosaur affinity of enigmatic large bone segments from the European Rhaetian.

Very large unidentified elongate and rounded fossil bone segments of uncertain origin recovered from different Rhaetian (Late Triassic) fossil localities across Europe have been puzzling the paleontological community since the second half of the 19th century. Different hypotheses have been proposed regarding the nature of these fossils: (1) giant amphibian bones, (2) dinosaurian or other archosaurian long bone shafts, and (3) giant ichthyosaurian jaw bone segments. We call the latter proposal the 'Giant Ichthyosaur Hypothesis' and test it using bone histology. In presumable ichthyosaur specimens from SW England (Lilstock), France (Autun), and indeterminate cortical fragments from Germany (Bonenburg), we found a combination of shared histological features in the periosteal cortex: an unusual woven-parallel complex of strictly longitudinal primary osteons set in a novel woven-fibered matrix type with intrinsic coarse collagen fibers (IFM), and a distinctive pattern of Haversian substitution in which secondary osteons often form within primary ones. The splenial and surangular of the holotype of the giant ichthyosaur Shastasaurus sikanniensis from Canada were sampled for comparison. The results of the sampling indicate a common osteohistology with the European specimens. A broad histological comparison is provided to reject alternative taxonomic affinities aside from ichthyosaurs of the very large bone segment. Most importantly, we highlight the occurrence of shared peculiar osteogenic processes in Late Triassic giant ichthyosaurs, reflecting special ossification strategies enabling fast growth and achievement of giant size and/or related to biomechanical properties akin to ossified tendons.

Puberty in a Mesozoic reptile.

The histology of bone can be preserved virtually unaltered for hundreds of millions of years in fossils from all environments and all vertebrate taxa, giving rise to the flourishing field of paleohistology.1 The shafts of long bones are formed by the apposition of periosteal bone tissue, similar to the growth of wood, and preserve, an often cyclical, record of the growth of the individual and events in its life history. One such event is sexual maturation or puberty, during which hormonal changes transform the juvenile into a sexually mature adult. Puberty has been well studied in humans and some other living vertebrates. Here, we describe puberty in Keichousaurus, a small sexually dimorphic and live-bearing marine reptile from Middle Triassic rocks of SW China, about 240 million years old. Using a combination of bone histology and morphology, we detected puberty2 as one of the four life stages (the others being fetus, juvenile, and adult). Adult Keichousaurus males have a more robust humerus than females, with pronounced muscle attachment sites and a triangular shaft cross section. Midshaft sections of the humeri of the males show the transition from the rounded juvenile cross section to the triangular adult cross section, as reflected in the contour of the growth marks. This shape change is produced by differential bone apposition of the periosteum, presumably triggered by sex hormones, as in humans,3 and influenced by changes in loading regime during puberty. This is the first report of puberty in a fossil amniote.

Plesiosaurs.

In the early 19th century, long before the discovery of the dinosaurs, scientists and the public alike were faced with the realization that strange beasts, wholly extinct, were once populating Earth's ancient oceans. In no small part, this realization was through the discovery of the first plesiosaurs (and ichthyosaurs) along the Dorset coast of England in the seaside town of Lyme Regis. There was this large marine reptile resembling a large sea turtle, but with four evenly shaped flippers and looking as though a large snake had been pulled through its carapace. It was soon to be named scientifically Plesiosaurus, in reference to its greater similarity to living reptiles than the Ichthyosaurus (Figure 1). While the Ichthyosaurus was relatively easily understood as a fish-shaped reptile descended from land-living ancestors, the Plesiosaurus was beyond comprehension, even though incomplete skeletons had been unearthed already in the early 18th century. Plesiosaurs seemed so alien that the first complete skeleton, discovered by the famed Mary Anning a little more than 200 years ago (Figure 1A), was considered a fake by the leading anatomist of the day, the Baron Georges Cuvier in Paris. Only study of the original specimen convinced him of the authenticity of this animal but reinforced his seminal insight that there is extinction.

Heads or tails first? Evolution of fetal orientation in ichthyosaurs, with a scrutiny of the prevailing hypothesis.

According to a longstanding paradigm, aquatic amniotes, including the Mesozoic marine reptile group Ichthyopterygia, give birth tail-first because head-first birth leads to increased asphyxiation risk of the fetus in the aquatic environment. Here, we draw upon published and original evidence to test two hypotheses: (1) Ichthyosaurs inherited viviparity from a terrestrial ancestor. (2) Asphyxiation risk is the main reason aquatic amniotes give birth tail-first. From the fossil evidence, we conclude that head-first birth is more prevalent in Ichthyopterygia than previously recognized and that a preference for tail-first birth likely arose in derived forms. This weakens the support for the terrestrial ancestry of viviparity in Ichthyopterygia. Our survey of extant viviparous amniotes indicates that fetal orientation at birth reflects a broad diversity of factors unrelated to aquatic vs. terrestrial habitat, further undermining the asphyxiation hypothesis. We propose that birth preference is based on parturitional mechanics or carrying efficiency rather than habitat.

Sauropods.

This article begins as many others on sauropods before it: "Sauropod dinosaurs were the largest animals to ever walk the Earth, by far". The largest sauropods were easily four times heavier than the largest land mammals (and the largest other dinosaur species, for that matter). The iconic body plan of sauropods is dominated by their very long neck, in some species exceeding 14 meters in length, provided with a relatively small head (Figure 1). The neck was mostly held horizontally or at a low angle. The massive but relatively short trunk was supported by four columnar legs, much like in an elephant. The bones in the fore foot of sauropods are oriented vertically, and some late forms even lost their finger bones, walking on their metacarpals (middle bones of the hand). The hind leg, which bore most of the weight, has a half-upright foot. The femur (thigh bone) was the largest bone in the skeleton, like in most other true land vertebrates (amniotes). The long neck was counterbalanced by the long tail, the base of which also functioned as the anchoring region of the giant muscles that pulled back the hind leg during walking.

A globally distributed durophagous marine reptile clade supports the rapid recovery of pelagic ecosystems after the Permo-Triassic mass extinction.

Marine ecosystem recovery after the Permo-Triassic mass extinction (PTME) has been extensively studied in the shallow sea, but little is known about the nature of this process in pelagic ecosystems. Omphalosauridae, an enigmatic clade of open-water durophagous marine reptiles, potentially played an important role in the recovery, but their fragmentary fossils and uncertain phylogenetic position have hindered our understanding of their role in the process. Here we report the large basal ichthyosauriform Sclerocormus from the Early Triassic of China that clearly demonstrates an omphalosaurid affinity, allowing for the synonymy of the recently erected Nasorostra with Omphalosauridae. The skull also reveals the anatomy of the unique feeding apparatus of omphalosaurids, likely an adaptation for feeding on hard-shelled pelagic invertebrates, especially ammonoids. Morphofunctional analysis of jaws shows that omphalosaurids occupy the morphospace of marine turtles. Our discovery adds another piece of evidence for an explosive radiation of marine reptiles into the ocean in the Early Triassic and the rapid recovery of pelagic ecosystems after the PTME.

To be or not to be heavier: The role of dermal bones in the buoyancy of the Late Triassic temnospondyl amphibian Metoposaurus krasiejowensis.

Stereospondyli are a clade of large aquatic temnospondyls known to have evolved a large dermal pectoral girdle. Among the Stereospondyli, metoposaurids in particular possess large interclavicles and clavicles relative to the rest of the postcranial skeleton. Because of the large size of these dermal bones, it was first proposed that they served as ballast during hydrostatic buoyancy control which assisted metoposaurids to live a bottom-dwelling mode of life. However, a large bone need not necessarily be heavy, for which determining the bone compactness becomes crucial for understanding any such adaptation in these dermal bones. Previous studies on the evolution of bone adaptations to aquatic lifestyles such as osteosclerosis, pachyostosis, osteoporotic-like pattern and pachyosteosclerosis have been observed in the long bones of secondarily aquatic amniotes. However, there are no known studies on the analysis of bone compactness in the dermal pectoral girdle of non-amniote taxa including Temnospondyli. This study looks at evidence of changes in bone mass adaptations in the dermal bones of the pectoral girdle of two stereospondyls occurring in the Late Triassic Krasiejόw locality (Southwestern Poland), namely: Metoposaurus krasiejowensis and Cyclotosaurus intermedius. However, because of lack of research on bone compactness of temnospondyls in general, there is no existent frame of reference to infer bone mass increase (BMI) in the M. krasiejowensis samples, and thus the bone compactness results of this taxon are compared with that of the samples of C. intermedius. Results of this study indicate that the interclavicles of M. krasiejowensis partially evolved BMI-like condition rendering these bones to be heavy enough to get selected as ballast during hydrostatic buoyancy control. Additionally, M. krasiejowensis shared its habitat with C. intermedius, however, the dermal pectoral girdle sample of the latter taxon does not display signs of BMI-like condition. Furthermore, the absence of variation in hydrostatic buoyancy control in the ontogenetic series of interclavicles of M. krasiejowensis could imply lack of ontogenetic niche shift along the water column.

Determination of muscle strength and function in plesiosaur limbs: finite element structural analyses of Cryptoclidus eurymerus humerus and femur.

Background: The Plesiosauria (Sauropterygia) are secondary marine diapsids. They are the only tetrapods to have evolved hydrofoil fore- and hindflippers. Once this specialization of locomotion had evolved, it remained essentially unchanged for 135 Ma. It is still controversial whether plesiosaurs flew underwater, rowed, or used a mixture of the two modes of locomotion. The long bones of Tetrapoda are functionally loaded by torsion, bending, compression, and tension during locomotion. Superposition of load cases shows that the bones are loaded mainly by compressive stresses. Therefore, it is possible to use finite element structure analysis (FESA) as a test environment for loading hypotheses. These include muscle reconstructions and muscle lines of action (LOA) when the goal is to obtain a homogeneous compressive stress distribution and to minimize bending in the model. Myological reconstruction revealed a muscle-powered flipper twisting mechanism. The flippers of plesiosaurs were twisted along the flipper length axis by extensors and flexors that originated from the humerus and femur as well as further distal locations. Methods: To investigate locomotion in plesiosaurs, the humerus and femur of a mounted skeleton of Cryptoclidus eurymerus (Middle Jurassic Oxford Clay Formation from Britain) were analyzed using FE methods based on the concept of optimization of loading by compression. After limb muscle reconstructions including the flipper twisting muscles, LOA were derived for all humerus and femur muscles of Cryptoclidus by stretching cords along casts of the fore- and hindflippers of the mounted skeleton. LOA and muscle attachments were added to meshed volumetric models of the humerus and femur derived from micro-CT scans. Muscle forces were approximated by stochastic iteration and the compressive stress distribution for the two load cases, "downstroke" and "upstroke", for each bone were calculated by aiming at a homogeneous compressive stress distribution. Results: Humeral and femoral depressors and retractors, which drive underwater flight rather than rowing, were found to exert higher muscle forces than the elevators and protractors. Furthermore, extensors and flexors exert high muscle forces compared to Cheloniidae. This confirms a convergently evolved myological mechanism of flipper twisting in plesiosaurs and complements hydrodynamic studies that showed flipper twisting is critical for efficient plesiosaur underwater flight.

Early Triassic ichthyopterygian fossils from the Russian Far East.

Ichthyopterygia is a major clade of reptiles that colonized the ocean after the end-Permian mass extinction, with the oldest fossil records found in early Spathian substage (late Olenekian, late Early Triassic) strata in the western USA. Here, we describe reptilian remains found in situ in the early Spathian Neocolumbites insignis ammonoid zone of South Primorye in the Russian Far East. Specimen NSM PV 23854 comprises fragmentary axial elements exhibiting a combination of morphological characteristics typical of Ichthyopterygia. The cylindrical centra suggest that the specimen represents a basal ichthyopterygian, and its size is comparable to that of Utatsusaurus. Specimen NSM PV 24995 is represented by a single limb bone, which is tentatively identified as an ichthyopterygian humerus. With a body length of approximately 5 m estimated from the humeral length, NSM PV 24995 represents one of the largest specimens of early Spathian marine reptiles known to date. Such size variation among the earliest ichthyopterygians might suggest an explosive diversification in size immediately after the end-Permian mass extinction. Both vertebrae and humerus specimens exhibit an extremely cancellous inner structure, suggesting a high degree of aquatic adaptation in ichthyopterygians, despite their short history of evolution in the ocean.

Early giant reveals faster evolution of large body size in ichthyosaurs than in cetaceans.

Body sizes of marine amniotes span six orders of magnitude, yet the factors that governed the evolution of this diversity are largely unknown. High primary production of modern oceans is considered a prerequisite for the emergence of cetacean giants, but that condition cannot explain gigantism in Triassic ichthyosaurs. We describe the new giant ichthyosaur Cymbospondylus youngorum sp. nov. with a 2-meter-long skull from the Middle Triassic Fossil Hill Fauna of Nevada, USA, underscoring rapid size evolution despite the absence of many modern primary producers. Surprisingly, the Fossil Hill Fauna rivaled the composition of modern marine mammal faunas in terms of size range, and energy-flux models suggest that Middle Triassic marine food webs were able to support several large-bodied ichthyosaurs at high trophic levels, shortly after ichthyosaur origins.

Chemistry of porphyrins in fossil plants and animals.

Porphyrins are macrocyclic tetrapyrrole derivatives that are widely distributed in nature. They are often complexed with a metal ion located in the center of the ring system and may be modified by various substituents including additional rings, or by ring opening, which leads to a plethora of different functions. Due to their extended conjugated aromatic ring system, porphyrins absorb light in the visible range and therefore show characteristic colors. Well-known natural porphyrins include the red-colored heme present in hemoglobin, which is responsible for blood oxygen transport, and the chlorophylls in some bacteria and in plants which are utilized for photosynthesis. Porphyrins are mostly lipophilic pigments that display relatively high chemical stability. Therefore, they can even survive hundreds of millions of years. The present review article provides an overview of natural porphyrins, their chemical structures, and properties. A special focus is put on porphyrins discovered in the fossil record. Examples will be highlighted, and information on their chemical analysis will be provided. We anticipate that the development of novel analytical methods with increased sensitivity will prompt new discoveries of porphyrins in fossils.

Bone histology and microanatomy of Edaphosaurus and Dimetrodon (Amniota, Synapsida) vertebrae from the Lower Permian of Texas.

Here we describe the histology and microanatomy of vertebral centra of the iconic pelycosaur-grade synapsids Edaphosaurus boanerges and Dimetrodon spp. Vertebrae from different axial positions and, in the case of Dimetrodon, from different ontogenetic stages were selected. For the histological description, we produced histological petrographic thin sections ground to a thickness of 50-80 µm of the vertebrae in the sagittal and transversal cutting planes. After the preparation process, the thin sections were examined under transmitted and cross-polarized light in a polarized microscope. The analyzed vertebrae reveal similar bone tissues, where both taxa have cortical parallel-fibered bone (PFB). PFB and lamellar bone (LB) forms in the cancellous part. However, in juvenile Dimetrodon, woven-fibered bone (WFB) is also deposited and shows a high degree of vascularity. This suggests that Dimetrodon had slightly faster bone growth than Edaphosaurus, which is mainly made of PFB and LB and shows poorly developed vascular canals. In addition, one specimen of Dimetrodon displays the preservation of an ossified notochord, which can be assumed to be indicative of how the intervertebral tissues were developed. Historically, evidence of how the joint between Dimetrodon vertebral centra was built was lacking until this specimen appeared. If the notochord ran persistently through the vertebrae, it would have possibly increased the stiffness of the vertebral column and would have affected the limbs and locomotion. Furthermore, the organization of trabeculae and relative thickness of the vertebral cortex gives insights into how the animals were adapted to their habitat.

Palaeontological evidence reveals convergent evolution of intervertebral joint types in amniotes.

The intervertebral disc (IVD) has long been considered unique to mammals. Palaeohistological sampling of 17 mostly extinct clades across the amniote tree revealed preservation of different intervertebral soft tissue types (cartilage, probable notochord) seen in extant reptiles. The distribution of the fossilised tissues allowed us to infer the soft part anatomy of the joint. Surprisingly, we also found evidence for an IVD in fossil reptiles, including non-avian dinosaurs, ichthyosaurs, plesiosaurs, and marine crocodiles. Based on the fossil dataset, we traced the evolution of the amniote intervertebral joint through ancestral character state reconstruction. The IVD evolved at least twice, in mammals and in extinct diapsid reptiles. From this reptilian IVD, extant reptile groups and some non-avian dinosaurs independently evolved a synovial ball-and-socket joint. The unique birds dorsal intervertebral joint evolved from this dinosaur joint. The tuatara and some geckos reverted to the ancestral persisting notochord.

Pentadactyl manus of the Metoposaurus krasiejowensis from the Late Triassic of Poland, the first record of pentadactyly among Temnospondyli.

Temnospondyli are commonly believed to have possessed four digits in the manus and five in the pes. However, actual finds of articulated autopodia are extremely rare. Therefore, an articulated, slightly incomplete forelimb skeleton with preserved manus of Metoposaurus krasiejowensis from the Late Triassic of Poland is important in providing new details about the structure and ossification sequence in the temnospondyl limb. The most important observation is the presence of five metacarpals in this specimen. This allows reconstructing the manus as pentadactyl. The number of phalanges and the distribution of distal articulation facets allow reconstruction of the digit formula as (2?)-3-3-(3?)-(2?). The well-developed fifth digit suggests that the Metoposaurus manus shows a unique ossification sequence: the reduction or late ossification of the first digit conforms to the amniote-frog pattern, and the early development of the second and third digit makes Metoposaurus similar to salamanders. Based on the distribution of pentadactyly vs. tetradactyly in the temnospondyl manus, the number of digits was not phylogenetically constrained in temnospondyls, similar to today's amphibians.

Neck mobility in the Jurassic plesiosaur Cryptoclidus eurymerus: finite element analysis as a new approach to understanding the cervical skeleton in fossil vertebrates.

The sauropterygian clade Plesiosauria arose in the Late Triassic and survived to the very end of the Cretaceous. Plesiosauria evolved the greatest species diversity of any marine reptile clade, attaining a global distribution. Plesiosauria consist of two clades, Rhomaleosauridae and Neoplesiosauria. Basal Neoplesiosauria have long necks with at least 30 cervicals, but show qualitative osteological evidence for a stiff neck. Here we quantify neck mobility in lateral, ventral, and dorsal directions based on finite element modeling of neck vertebrae from the Middle Jurassic plesiosaur Cryptoclidus eurymerus. We model the mobility in a single motion segment, consisting of two adjacent cervical vertebrae and the joints connecting them. Based on the model with a maximum intervertebral spacing of 3 mm, we find that in Cryptoclidus, the maximum angle of lateral deflection in the motion segment was 2°. The maximum angle of ventral deflection was 5° and of dorsal deflection was 5°. When these values are multiplied by the number of cervical vertebrae, it becomes apparent that neck mobility was limited in all directions. The maximum angle of total lateral deflection in the neck was 67°. The maximum angle of total ventral deflection was 148° and of total dorsal deflection was 157°. This raises the question of the function of such a long, multi-segment but immobile neck. We posit that the long neck served in hydrodynamic and visual camouflage, hiding the bulk of the body from the small but abundant prey, such as schooling fish and squid. Neck immobility may have been advantageous in withstanding strong hydrodynamic forces acting on the neck during predatory strikes.

Hematological convergence between Mesozoic marine reptiles (Sauropterygia) and extant aquatic amniotes elucidates diving adaptations in plesiosaurs.

Plesiosaurs are a prominent group of Mesozoic marine reptiles, belonging to the more inclusive clades Pistosauroidea and Sauropterygia. In the Middle Triassic, the early pistosauroid ancestors of plesiosaurs left their ancestral coastal habitats and increasingly adapted to a life in the open ocean. This ecological shift was accompanied by profound changes in locomotion, sensory ecology and metabolism. However, investigations of physiological adaptations on the cellular level related to the pelagic lifestyle are lacking so far. Using vascular canal diameter, derived from osteohistological thin-sections, we show that inferred red blood cell size significantly increases in pistosauroids compared to more basal sauropterygians. This change appears to have occurred in conjunction with the dispersal to open marine environments, with cell size remaining consistently large in plesiosaurs. Enlarged red blood cells likely represent an adaptation of plesiosaurs repeated deep dives in the pelagic habitat and mirror conditions found in extant marine mammals and birds. Our results emphasize physiological aspects of adaptive convergence among fossil and extant marine amniotes and add to our current understanding of plesiosaur evolution.

Structure and evolutionary implications of the earliest (Sinemurian, Early Jurassic) dinosaur eggs and eggshells.

One of the fossil record's most puzzling features is the absence of preserved eggs or eggshell for the first third of the known 315 million year history of amniote evolution. Our meagre understanding of the origin and evolution of calcareous eggshell and amniotic eggs in general, is largely based on Middle Jurassic to Late Cretaceous fossils. For dinosaurs, the most parsimonious inference yields a thick, hard shelled egg, so richly represented in the Late Cretaceous fossil record. Here, we show that a thin calcareous layer (≤100 µm) with interlocking units of radiating crystals (mammillae) and a thick shell membrane already characterize the oldest known amniote eggs, belonging to three coeval, but widely distributed Early Jurassic basal sauropodomorph dinosaurs. This thin shell layer strongly contrasts with the considerably thicker calcareous shells of Late Jurassic dinosaurs. Phylogenetic analyses and their Sinemurian age indicate that the thin eggshell of basal sauropodomorphs represents a major evolutionary innovation at the base of Dinosauria and that the much thicker eggshell of sauropods, theropods, and ornithischian dinosaurs evolved independently. Advanced mineralization of amniote eggshell (≥150 µm in thickness) in general occurred not earlier than Middle Jurassic and may correspond with a global trend of increase in atmospheric oxygen.

Fossilization transforms vertebrate hard tissue proteins into N-heterocyclic polymers.

Vertebrate hard tissues consist of mineral crystallites within a proteinaceous scaffold that normally degrades post-mortem. Here we show, however, that decalcification of Mesozoic hard tissues preserved in oxidative settings releases brownish stained extracellular matrix, cells, blood vessels, and nerve projections. Raman Microspectroscopy shows that these fossil soft tissues are a product of diagenetic transformation to Advanced Glycoxidation and Lipoxidation End Products, a class of N-heterocyclic polymers generated via oxidative crosslinking of proteinaceous scaffolds. Hard tissues in reducing environments, in contrast, lack soft tissue preservation. Comparison of fossil soft tissues with modern and experimentally matured samples reveals how proteinaceous tissues undergo diagenesis and explains biases in their preservation in the rock record. This provides a target, focused on oxidative depositional environments, for finding cellular-to-subcellular soft tissue morphology in fossils and validates its use in phylogenetic and other evolutionary studies.

Dinosaurs: Four Legs Good, Two Legs Bad.

The quadrupedal Sauropods - the biggest dinosaurs to walk the Earth - evolved from bipedal ancestors. Two new early sauropodomorphs from South Africa and Argentina indicate that very large, flexed-limbed sauropodomorphs coexisted with early columnar-limbed sauropods for 20 million years.

Fossil eggshell cuticle elucidates dinosaur nesting ecology.

The cuticle layer consisting mainly of lipids and hydroxyapatite (HAp) atop the mineralized avian eggshell is a protective structure that prevents the egg from dehydration and microbial invasions. Previous ornithological studies have revealed that the cuticle layer is also involved in modulating the reflectance of eggshells in addition to pigments (protoporphyrin and biliverdin). Thus, the cuticle layer represents a crucial trait that delivers ecological signals. While present in most modern birds, direct evidence for cuticle preservation in stem birds and non-avian dinosaurs is yet missing. Here we present the first direct and chemical evidence for the preservation of the cuticle layer on dinosaur eggshells. We analyze several theropod eggshells from various localities, including oviraptorid Macroolithus yaotunensis eggshells from the Late Cretaceous deposits of Henan, Jiangxi, and Guangdong in China and alvarezsaurid Triprismatoolithus eggshell from the Two Medicine Formation of Montana, United States, with the scanning electron microscope (SEM), electron probe micro-analysis (EPMA), and Raman spectroscopy (RS). The elemental analysis with EPMA shows high concentration of phosphorus at the boundary between the eggshell and sediment, representing the hydroxyapatitic cuticle layer (HAp). Depletion of phosphorus in sediment excludes the allochthonous origin of the phosphorus in these eggshells. The chemometric analysis of Raman spectra collected from fossil and extant eggs provides further supportive evidence for the cuticle preservation in oviraptorid and probable alvarezsaurid eggshells. In accordance with our previous discovery of pigments preserved in Cretaceous oviraptorid dinosaur eggshells, we validate the cuticle preservation on dinosaur eggshells through deep time and offer a yet unexplored resource for chemical studies targeting the evolution of dinosaur nesting ecology. Our study also suggests that the cuticle structure can be traced far back to maniraptoran dinosaurs and enhance their reproductive success in a warm and mesic habitat such as Montana and southern China during the Late Cretaceous.