Quantifying the effects of exceptional fossil preservation on the global availability of phylogenetic data in deep time.

Fossil deposits with exceptional preservation ("lagerstätten") provide important details not typically preserved in the fossil record, such that they hold an outsized influence on our understanding of biodiversity and evolution. In particular, the potential bias imparted by this so-called "lagerstätten effect" remains a critical, but underexplored aspect of reconstructing evolutionary relationships. Here, we quantify the amount of phylogenetic information available in the global fossil records of 1,327 species of non-avian theropod dinosaurs, Mesozoic birds, and fossil squamates (e.g., lizards, snakes, mosasaurs), and then compare the influence of lagerstätten deposits on phylogenetic information content and taxon selection in phylogenetic analyses to other fossil-bearing deposits. We find that groups that preserve a high amount of phylogenetic information in their global fossil record (e.g., non-avian theropods) are less vulnerable to a "lagerstätten effect" that leads to disproportionate representation of fossil taxa from one geologic unit in an evolutionary tree. Additionally, for each taxonomic group, we find comparable amounts of phylogenetic information in lagerstätten deposits, even though corresponding morphological character datasets vary greatly. Finally, we unexpectedly find that ancient sand dune deposits of the Late Cretaceous Gobi Desert of Mongolia and China exert an anomalously large influence on the phylogenetic information available in the squamate fossil record, suggesting a "lagerstätten effect" can be present in units not traditionally considered lagerstätten. These results offer a phylogenetics-based lens through which to examine the effects of exceptional fossil preservation on biological patterns through time and space, and invites further quantification of evolutionary information in the rock record.

An Italian dinosaur Lagerstätte reveals the tempo and mode of hadrosauriform body size evolution.

During the latest Cretaceous, the European Archipelago was characterized by highly fragmented landmasses hosting putative dwarfed, insular dinosaurs, claimed as fossil evidence of the "island rule". The Villaggio del Pescatore quarry (north-eastern Italy) stands as the most informative locality within the palaeo-Mediterranean region and represents the first, multi-individual Konservat-Lagerstätte type dinosaur-bearing locality in Italy. The site is here critically re-evaluated as early Campanian in age, thus preceding the final fragmentation stages of the European Archipelago, including all other European localities preserving hypothesized dwarfed taxa. New skeletal remains allowed osteohistological analyses on the hadrosauroid Tethyshadros insularis indicating subadult features in the type specimen whereas a second, herein newly described, larger individual is likely somatically mature. A phylogenetic comparative framework places the body-size of T. insularis in range with other non-hadrosaurid Eurasian hadrosauroids, rejecting any significant evolutionary trend towards miniaturisation in this clade, confuting its 'pygmy' status, and providing unmatched data to infer environmentally-driven body-size trends in Mesozoic dinosaurs.

Novel Approach to Tooth Chemistry. Quantification of the Dental-Enamel Junction.

The dentin-enamel junction (DEJ) is known for its special role in teeth. Several techniques were applied for the investigation of the DEJ in human sound molar teeth. The electron (EPMA) and proton (PIXE) microprobes gave consistent indications about the variability of elemental concentrations on this boundary. The locally increased and oscillating concentrations of Mg and Na were observed in the junction, in the layer adhering to the enamel and covering roughly half of the DEJ width. The chemical results were compared with the optical profiles of the junction. Our chemical and optical results were next compared with the micromechanical results (hardness, elastic modulus, friction coefficient) available in the world literature. A strong correlation of both result sets was proven, which testifies to the self-affinity of the junction structures for different locations and even for different kinds of teeth and techniques applied for studies. Energetic changes in tooth strictly connected with crystallographic transformations were calculated, and the minimum energetic status was discovered for DEJ zone. Modeling of both walls of the DEJ from optical data was demonstrated. Comparing the DEJ in human teeth with the same structure found in dinosaur, shark, and alligator teeth evidences the universality of dentin enamel junction in animal world. The paper makes a contribution to better understanding the joining of the different hard tissues.

A new styracosternan hadrosauroid (Dinosauria: Ornithischia) from the Early Cretaceous of Portell, Spain.

A new styracosternan ornithopod genus and species is described based on the right dentary of a single specimen from the Mirambell Formation (Early Cretaceous, early Barremian) at the locality of Portell, (Castellón, Spain). Portellsaurus sosbaynati gen. et sp. nov. is diagnosed by two autapomorphic features as well as a unique combination of characters. The autapomorphies include: the absence of a bulge along the ventral margin directly ventral to the base of the coronoid process and the presence of a deep oval cavity on the medial surface of the mandibular adductor fossa below the eleventh-twelfth tooth position. Phylogenetic analyses reveal that the new Iberian form is more closely related to the African taxon Ouranosaurus nigeriensis than to its synchronic Iberian taxa Magnamanus soriaensis and Iguanodon galvensis. In addition, Portellsaurus sosbaynati is less related to other Iberian taxa such as Iguanodon bernissartensis and Proa valdearinnoensis than to the other Early Cretaceous Iberian styracosternans Mantellisaurus atherfieldensis and Morelladon beltrani. A new phylogenetic hypothesis is proposed that resolves Iguanodon (I. bernissartensis, I. galvensis) with the Valanginian Barilium dawsoni into a monophyletic clade (Iguanodontoidea). The recognition of Portellsaurus sosbaynati gen. et sp. nov. as the first styracosternan dinosaur species identified from the Margas de Mirambell Formation (early Barremian-early late Barremian) in the Morella sub-basin (Maestrat Basin, eastern Spain) indicates that the Iberian Peninsula was home to a highly diverse assemblage of medium-to-large bodied styracosternan hadrosauriforms during the Early Cretaceous.

Empirical and Methodological Challenges to the Model-Based Inference of Diversification Rates in Extinct Clades.

Changes in speciation and extinction rates are key to the dynamics of clade diversification, but attempts to infer them from phylogenies of extant species face challenges. Methods capable of synthesizing information from extant and fossil species have yielded novel insights into diversification rate variation through time, but little is known about their behavior when analyzing entirely extinct clades. Here, we use empirical and simulated data to assess how two popular methods, PyRate and Fossil BAMM, perform in this setting. We inferred the first tip-dated trees for ornithischian dinosaurs and combined them with fossil occurrence data to test whether the clade underwent an end-Cretaceous decline. We then simulated phylogenies and fossil records under empirical constraints to determine whether macroevolutionary and preservation rates can be teased apart under paleobiologically realistic conditions. We obtained discordant inferences about ornithischian macroevolution including a long-term speciation rate decline (BAMM), mostly flat rates with a steep diversification drop (PyRate) or without one (BAMM), and episodes of implausibly accelerated speciation and extinction (PyRate). Simulations revealed little to no conflation between speciation and preservation, but yielded spuriously correlated speciation and extinction estimates while time-smearing tree-wide shifts (BAMM) or overestimating their number (PyRate). Our results indicate that the small phylogenetic data sets available to vertebrate paleontologists and the assumptions made by current model-based methods combine to yield potentially unreliable inferences about the diversification of extinct clades. We provide guidelines for interpreting the results of the existing approaches in light of their limitations and suggest how the latter may be mitigated. [BAMM; diversification; fossils; macroevolutionary rates; Ornithischia; PyRate.].

A neoceratopsian dinosaur from the early Cretaceous of Mongolia and the early evolution of ceratopsia.

Ceratopsia is a diverse dinosaur clade from the Middle Jurassic to Late Cretaceous with early diversification in East Asia. However, the phylogeny of basal ceratopsians remains unclear. Here we report a new basal neoceratopsian dinosaur Beg tse based on a partial skull from Baruunbayan, Ömnögovi aimag, Mongolia. Beg is diagnosed by a unique combination of primitive and derived characters including a primitively deep premaxilla with four premaxillary teeth, a trapezoidal antorbital fossa with a poorly delineated anterior margin, very short dentary with an expanded and shallow groove on lateral surface, the derived presence of a robust jugal having a foramen on its anteromedial surface, and five equally spaced tubercles on the lateral ridge of the surangular. This is to our knowledge the earliest known occurrence of basal neoceratopsian in Mongolia, where this group was previously only known from Late Cretaceous strata. Phylogenetic analysis indicates that it is sister to all other neoceratopsian 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.

More than one way to be a giant: Convergence and disparity in the hip joints of saurischian dinosaurs.

Saurischian dinosaurs evolved seven orders of magnitude in body mass, as well as a wide diversity of hip joint morphology and locomotor postures. The very largest saurischians possess incongruent bony hip joints, suggesting that large volumes of soft tissues mediated hip articulation. To understand the evolutionary trends and functional relationships between body size and hip anatomy of saurischians, we tested the relationships among discrete and continuous morphological characters using phylogenetically corrected regression. Giant theropods and sauropods convergently evolved highly cartilaginous hip joints by reducing supraacetabular ossifications, a condition unlike that in early dinosauromorphs. However, transitions in femoral and acetabular soft tissues indicate that large sauropods and theropods built their hip joints in fundamentally different ways. In sauropods, the femoral head possesses irregularly rugose subchondral surfaces for thick hyaline cartilage. Hip articulation was achieved primarily using the highly cartilaginous femoral head and the supraacetabular labrum on the acetabular ceiling. In contrast, theropods covered their femoral head and neck with thinner hyaline cartilage and maintained extensive articulation between the fibrocartilaginous femoral neck and the antitrochanter. These findings suggest that the hip joints of giant sauropods were built to sustain large compressive loads, whereas those of giant theropods experienced compression and shear forces.

Modeling Dragons: Using linked mechanistic physiological and microclimate models to explore environmental, physiological, and morphological constraints on the early evolution of dinosaurs.

We employed the widely-tested biophysiological modeling software, Niche Mapper™ to investigate the metabolic function of the Late Triassic dinosaurs Plateosaurus and Coelophysis during global greenhouse conditions. We tested a variety of assumptions about resting metabolic rate, each evaluated within six microclimate models that bound paleoenvironmental conditions at 12° N paleolatitude, as determined by sedimentological and isotopic proxies for climate within the Chinle Formation of the southwestern United States. Sensitivity testing of metabolic variables and simulated "metabolic chamber" analyses support elevated "ratite-like" metabolic rates and intermediate "monotreme-like" core temperature ranges in these species of early saurischian dinosaur. Our results suggest small theropods may have needed partial to full epidermal insulation in temperate environments, while fully grown prosauropods would have likely been heat stressed in open, hot environments and should have been restricted to cooler microclimates such as dense forests or higher latitudes and elevations. This is in agreement with the Late Triassic fossil record and may have contributed to the latitudinal gap in the Triassic prosauropod record.

Forelimb shortening of Carcharodontosauria (Dinosauria: Theropoda): an update on evolutionary anterior micromelias in non-avian theropods.

Evolutionary teratology recognises certain anatomical modifications as developmental anomalies. Within non avian-theropod dinosaurs, the strong forelimb shortening of Tyrannosauridae, Carnotaurinae and Limusaurus - associated with a reduction or loss of autonomy - have been previously diagnosed as evolutionary anterior micromelias. The feature is here examined with Acrocanthosaurus atokensis (Carcharodontosauridae) and Gualicho shinyae (Neovenatoridae). The micromelic diagnosis is confirmed for Acrocanthosaurus, without supplementary malformations. Gualicho is considered as a borderline case, outside of the micromelic spectrum, but shows a total phalangeal loss on digit III. The reduction in the biomechanical range of Acrocanthosaurus' forelimbs was compensated by the skull and jaws as main predatory organs. The same is assumed for Gualicho, but its robust first digit and raptorial claw are to be underlined. Other gigantic-sized and derived representatives of Carcharodontosauridae probably shared the anterior micromelia condition, potentially due to developmental modifications involving differential forelimbs/hindlimbs embryological growth rates, secondarily associated with post-natal growth rates leading to large and gigantic sizes; a converging state with Tyrannosauridae. Nevertheless, whereas developmental growth rates are also considered in the shortened condition of Gualicho, there is no association with post-natal gigantism. Finally, the digit III reduction likely followed the same evolutionary pathways as Tyrannosauridae, potentially involving BMPs, Fgfs and Shh signalling.

Time lapse: A glimpse into prehistoric genomics.

For the purpose of this review, 'time-lapse' refers to the reconstruction of ancestral (in this case dinosaur) karyotypes using genome assemblies of extant species. Such reconstructions are only usually possible when genomes are assembled to 'chromosome level' i.e. a complete representation of all the sequences, correctly ordered contiguously on each of the chromosomes. Recent paleontological evidence is very clear that birds are living dinosaurs, the latest example of dinosaurs emerging from a catastrophic extinction event. Non-avian dinosaurs (ever present in the public imagination through art, and broadcast media) emerged some 240 million years ago and have displayed incredible phenotypic diversity. Here we report on our recent studies to infer the overall karyotype of the Theropod dinosaur lineage from extant avian chromosome level genome assemblies. Our work first focused on determining the likely karyotype of the avian ancestor (most likely a chicken-sized, two-legged, feathered, land dinosaur from the Jurassic period) finding karyotypic similarity to the chicken. We then took the work further to determine the likely karyotype of the bird-lizard ancestor and the chromosomal changes (chiefly translocations and inversions) that occurred between then and modern birds. A combination of bioinformatics and cross-species fluorescence in situ hybridization (zoo-FISH) uncovered a considerable number of translocations and fissions from a 'lizard-like' genome structure of 2n = 36-46 to one similar to that of soft-shelled turtles (2n = 66) from 275 to 255 million years ago (mya). Remarkable karyotypic similarities between some soft-shelled turtles and chicken suggests that there were few translocations from the bird-turtle ancestor (plus ∼7 fissions) through the dawn of the dinosaurs and pterosaurs, through the theropod linage and on to most to modern birds. In other words, an avian-like karyotype was in place about 240mya when the dinosaurs and pterosaurs first emerged. We mapped 49 chromosome inversions from then to the present day, uncovering some gene ontology enrichment in evolutionary breakpoint regions. This avian-like karyotype with its many (micro)chromosomes provides the basis for variation (the driver of natural selection) through increased random segregation and recombination. It may therefore contribute to the ability of dinosaurs to survive multiple extinction events, emerging each time as speciose and diverse.

Birdlike growth and mixed-age flocks in avimimids (Theropoda, Oviraptorosauria).

Avimimids were unusual, birdlike oviraptorosaurs from the Late Cretaceous of Asia. Initially enigmatic, new information has ameliorated the understanding of their anatomy, phylogenetic position, and behaviour. A monodominant bonebed from the Nemegt Formation of Mongolia showed that some avimimids were gregarious, but the site is unusual in the apparent absence of juveniles. Here, a second monodominant avimimid bonebed is described from the Iren Dabasu Formation of northern China. Elements recovered include numerous vertebrae and portions of the forelimbs and hindlimbs, representing a minimum of six individuals. Histological sampling of two tibiotarsi from the bonebed reveals rapid growth early in ontogeny followed by unexpectedly early onset of fusion and limited subsequent growth. This indicates that avimimids grew rapidly to adult size, like most extant birds but contrasting other small theropod dinosaurs. The combination of adults and juveniles in the Iren Dabasu bonebed assemblage provides evidence of mixed-age flocking in avimimids and the onset of fusion in young individuals suggests that some of the individuals in the Nemegt Formation bonebed may have been juveniles. Regardless, these individuals were likely functionally analogous to adults, and this probably facilitated mixed-age flocking by reducing ontogenetic niche shift in avimimids.

Reassessment of a juvenile Daspletosaurus from the Late Cretaceous of Alberta, Canada with implications for the identification of immature tyrannosaurids.

Daspletosaurus is a large tyrannosaurine found in upper Campanian deposits of Alberta and Montana. Although several large subadult and adult individuals of this taxon are known, only one juvenile individual, TMP 1994.143.1, has been identified. This specimen has played a key role in the idea that juvenile tyrannosaurid individuals are difficult to differentiate among species. Here the taxonomic affinity of TMP 1994.143.1 is reassessed in light of a juvenile tyrannosaurine postorbital recently discovered in the Dinosaur Park Formation of Alberta. Anatomical comparisons and phylogenetic analyses reveal that TMP 1994.143.1 is referable to the albertosaurine Gorgosaurus libratus, whereas the new postorbital belongs to a small juvenile Daspletosaurus. This taxonomic reassignment of TMP 1994.143.1 results in the juvenile ontogenetic stage of Daspletosaurus being known only from two isolated cranial elements. The new postorbital provides insights into early Daspletosaurus ontogeny, revealing that the cornual process developed earlier or faster than in other tyrannosaurids. Although some ontogenetic changes in the postorbital are found to be unique to Daspletosaurus, overall changes are most consistent with those of other large tyrannosaurines. Our results also show that diagnostic features develop early in ontogeny, such that juveniles of different tyrannosaurid species are easier to differentiate than previously thought.

A new alvarezsaurian theropod from the Upper Jurassic Shishugou Formation of western China.

Alvarezsaurian dinosaurs, a group of bizarre theropods with greatly shortened and modified forelimbs, are known mostly from the Cretaceous of Asia and South America. Here we report a new alvarezsaurian, Shishugounykus inexpectus gen. et sp. nov, based on a specimen recovered from the Middle-Upper Jurassic Shishugou Formation of the Junggar Basin, western China. Together with two other alvarezsaurians from this formation, i.e., Haplocheirus sollers and Aorun zhaoi, these Shishugou forms represent the only known Jurassic alvarezsaurians worldwide. Similar to the two other Shishugou alvarezsaurians, this new alvarezsaurian displays early stages in the development of the highly modified alvarezsaurian forelimb, but it possesses a number of manual features closer to the typical coelurosaurian theropod condition. Combining morphological and histological features, our analysis indicates that the earliest known alvarezsaurians are variable in size and other important morphological features, and in particular display a mosaic distribution of forelimb features.

Olfactory receptor repertoire size in dinosaurs.

The olfactory bulb (OB) ratio is the size of the OB relative to the cerebral hemisphere, and is used to estimate the proportion of the forebrain devoted to smell. In birds, OB ratio correlates with the number of olfactory receptor (OR) genes and therefore has been used as a proxy for olfactory acuity. By coupling OB ratios with known OR gene repertoires in birds, we infer minimum repertoire sizes for extinct taxa, including non-avian dinosaurs, using phylogenetic modelling, ancestral state reconstruction and comparative genomics. We highlight a shift in the scaling of OB ratio to body size along the lineage leading to modern birds, demonstrating variable OR repertoires present in different dinosaur and crown-bird lineages, with varying factors potentially influencing sensory evolution in theropods. We investigate the ancestral sensory space available to extinct taxa, highlighting potential adaptations to ecological niches. Through combining morphological and genomic data, we show that, while genetic information for extinct taxa is forever lost, it is potentially feasible to investigate evolutionary trajectories in extinct genomes.

A new home for microbes.

Modern microorganisms growing in fossils provide major challenges for researchers trying to detect ancient molecules in the same fossils.

Ontogenetic changes in the body plan of the sauropodomorph dinosaur Mussaurus patagonicus reveal shifts of locomotor stance during growth.

Ontogenetic information is crucial to understand life histories and represents a true challenge in dinosaurs due to the scarcity of growth series available. Mussaurus patagonicus was a sauropodomorph dinosaur close to the origin of Sauropoda known from hatchling, juvenile and mature specimens, providing a sufficiently complete ontogenetic series to reconstruct general patterns of ontogeny. Here, in order to quantify how body shape and its relationship with locomotor stance (quadruped/biped) changed in ontogeny, hatchling, juvenile (~1 year old) and adult (8+ years old) individuals were studied using digital models. Our results show that Mussaurus rapidly grew from about 60 g at hatching to ~7 kg at one year old, reaching >1000 kg at adulthood. During this time, the body's centre of mass moved from a position in the mid-thorax to a more caudal position nearer to the pelvis. We infer that these changes of body shape and centre of mass reflect a shift from quadrupedalism to bipedalism occurred early in ontogeny in Mussaurus. Our study indicates that relative development of the tail and neck was more influential in determining the locomotor stance in Sauropodomorpha during ontogeny, challenging previous studies, which have emphasized the influence of hindlimb vs. forelimb lengths on sauropodomorph stance.

The Evolution of Pneumatic Foramina in Pterosaur Vertebrae.

Pterosaurs possess skeletal pneumatization, which can be assessed externally through the observation of pneumatic foramina. These foramina vary in position, shape, and number among taxa. Here we propose new characters referring to the position and number of pneumatic foramina present in cervical and dorsal vertebrae of pterosaurs. A cladistic analysis was performed after first-hand study of material in collections and using data from the literature in order to test these new characters for homology, and they were subsequently mapped in a strict consensus tree. The analysis resulted in nine equally parsimonious trees with 215 steps each. The strict consensus tree obtained does not present significant differences in relation to trees obtained in previous studies. The mapping performed identified that most pneumatic foramina evolved independently in several lineages, and only two of the eight added characters appeared once in the evolution of pterosaurs. In general, pneumatic foramina were more common in the Dsungaripteroidea, however, the flattened preservation of non-pterodactyloids and in the Archaeopterodactyloidea often precludes their visualization, and therefore the presence of vertebral pneumatization may be broaden with the discovery of new, better preserved specimens.

A new hadrosauroid (Dinosauria: Ornithopoda) from the Late Cretaceous Baynshire Formation of the Gobi Desert (Mongolia).

A new genus and species of non-hadrosaurid hadrosauroid, Gobihadros mongoliensis, is described from a virtually complete and undeformed skull and postcranial skeleton, as well as extensive referred material, collected from the Baynshire Formation (Cenomanian-Santonian) of the central and eastern Gobi Desert, Mongolia. Gobihadros mongoliensis is the first non-hadrosaurid hadrosauroid from the Late Cretaceous of central Asia known from a complete, articulated skull and skeleton. The material reveals the skeletal anatomy of a proximate sister taxon to Hadrosauridae in remarkable detail. Gobihadros is similar to Bactrosaurus johnsoni and Gilmoreosaurus mongoliensis, but can be distinguished from them in several autapomorphic traits, including the maximum number (three) of functional dentary teeth per tooth position, a premaxillary oral margin with a 'double-layer morphology', and a sigmoidal dorsal outline of the ilium with a well-developed, fan-shaped posterior process. All of these characters in Gobihadros are inferred to be convergent in Hadrosauridae. Phylogenetic analysis positions Gobihadros mongoliensis as a Bactrosaurus-grade hadrosauromorph hadrosauroid. Its relationship with Maastrichtian hadrosaurids from Asia (e.g., Saurolophus angustirostris, Kerberosaurus manakini, Wulagasaurus dongi, Kundurosaurus nagornyi) are sufficiently distant to indicate that these latter taxa owe their distribution to migration from North America across Beringia, rather than having a common Asian origin with Go. mongoliensis.

A new transitional therizinosaurian theropod from the Early Cretaceous Jehol Biota of China.

Therizinosaurian theropods evolved many highly specialized osteological features in association with their bulky proportions, which were unusual in the context of the generally gracile Theropoda. Here we report a new therizinosaur, Lingyuanosaurus sihedangensis gen. et sp. nov., based on a specimen recovered from the Lower Cretaceous Jehol Group of Lingyuan, Liaoning Province, China, which displays a combination of plesiomorphic and derived features. Most notably, the specimen is characterized by posterior dorsal vertebrae with a complex and unusual laminar structure; an ilium with a highly dorsoventrally expanded preacetabular process showing only slight lateral flaring of the ventral margin, a strongly anterodorsally inclined iliac blade, a small postacetabular process with a strongly concave dorsal margin, and a relatively robust pubic peduncle with a posteroventrally facing distal articular surface; a straight and robust femur with a small lesser trochanter; and a tibia that is longer than the femur. Phylogenetic analysis places Lingyuanosaurus in an intermediate position within Therizinosauria, i.e., between the early-branching therizinosaurs such as Falcarius, Jianchangosaurus, and Beipiaosaurus and the late-branching ones such as Alxasaurus and Therizinosaurus. This new therizinosaur sheds additional light on the evolution of major therizinosaurian characteristics, including particularly the distinctive pelvic girdle and hindlimb morphology seen in this group.