A Sea Scorpion's Strike: New Evidence of Extreme Lateral Flexibility in the Opisthosoma of Eurypterids.

Among the largest and most abundant aquatic predators during much of the early evolution of vertebrates, eurypterids have long been an iconic and intensely studied group of Paleozoic arthropods. We report a new specimen of the eurypterid Slimonia acuminata, which includes a fully articulated series of tail (postabdominal and telsonal) segments preserved in a tight lateral curve. Such a high degree of apparent lateral tail flexibility has not been previously recognized in eurypterids. From the perspective of hydrodynamics, the dorsoventrally flattened body plan of eurypterids would have limited the effectiveness of lateral tail motion as a means of propulsion. However, the long and serrated terminal tail spine of S. acuminata (and other eurypterids) would have made lateral tail strikes-which would have met a minimum of hydraulic resistance-an effective means of predatory attack and self-defense. Thus, many eurypterids are reinterpreted as substantially better-armed predators than previously supposed.

The functional origin of dinosaur bipedalism: Cumulative evidence from bipedally inclined reptiles and disinclined mammals.

Bipedalism is a trait basal to, and widespread among, dinosaurs. It has been previously argued that bipedalism arose in the ancestors of dinosaurs for the function of freeing the forelimbs to serve as predatory weapons. However, this argument does not explain why bipedalism was retained among numerous herbivorous groups of dinosaurs. We argue that bipedalism arose in the dinosaur line for the purpose of enhanced cursoriality. Modern facultatively bipedal lizards offer an analog for the first stages in the evolution of dinosaurian bipedalism. Many extant lizards assume a bipedal stance while attempting to flee predators at maximum speed. Bipedalism, when combined with a caudofemoralis musculature, has cursorial advantages because the caudofemoralis provides a greater source of propulsion to the hindlimbs than is generally available to the forelimbs. That cursorial advantage explains the relative abundance of cursorial facultative bipeds and obligate bipeds among fossil diapsids and the relative scarcity of either among mammals. Having lost their caudofemoralis in the Permian, perhaps in the context of adapting to a fossorial lifestyle, the mammalian line has been disinclined towards bipedalism, but, having never lost the caudofemoralis of their ancestors, cursorial avemetatarsalians (bird-line archosaurs) were naturally inclined towards bipedalism.

Tyrannosauroid integument reveals conflicting patterns of gigantism and feather evolution.

Recent evidence for feathers in theropods has led to speculations that the largest tyrannosaurids, including Tyrannosaurus rex, were extensively feathered. We describe fossil integument from Tyrannosaurus and other tyrannosaurids (Albertosaurus, Daspletosaurus, Gorgosaurus and Tarbosaurus), confirming that these large-bodied forms possessed scaly, reptilian-like skin. Body size evolution in tyrannosauroids reveals two independent occurrences of gigantism; specifically, the large sizes in Yutyrannus and tyrannosaurids were independently derived. These new findings demonstrate that extensive feather coverings observed in some early tyrannosauroids were lost by the Albian, basal to Tyrannosauridae. This loss is unrelated to palaeoclimate but possibly tied to the evolution of gigantism, although other mechanisms exist.

Deinonychosaur trackways in southeastern China record a possible giant troodontid.

The Longxiang tracksite (lower Upper Cretaceous, Shanghang Basin) includes twelve didactyl deinonychosaur tracks that fall into two morphologies, differentiated by both size and form. The smaller tracks (∼11 cm long) are referable to the ichnogenus Velociraptorichnus. The larger tracks (∼36 cm long) establish the ichnotaxon Fujianipus yingliangi. Based on the size of the tracks, F. yingliangi has an estimated hip height of over 1.8 m, a size comparable to that of the largest known deinonychosaurs, i.e., Austroraptor and Utahraptor. The reduced form of digit IV, relative to digit III, indicates that F. yingliangi is a probable troodontid. Gigantism evidently evolved independently at least four times within the Deinonychosauria and within at least three major lineages: the Eudromaeosauria, Unenlagiidae, and Troodontidae. In the mid-Cretaceous of Asia, the evolution of F. yingliangi overlapped with that of early large-bodied tyrannosauroids and with previously established large allosaurids (although the latter may have been in decline).

New data on tail lengths and variation along the caudal series in the non-avialan dinosaurs.

The tails of non-avialan dinosaurs varied considerably in terms of overall length, total number of vertebrae, and gross form and function. A new dataset confirms that there is little or no consistent relationship between tail length and snout-sacrum length. Consequently, attempts to estimate one from the other are likely to be very error-prone. Patterns of changes in centra lengths across the caudal series vary among non-avian dinosaurs. However, some overarching patterns do emerge. A number of taxa show (anterior to posterior) a series of short centra, followed by a series of longer centra, with the remainder of the tail consisting of a long series of centra tapering in length. This pattern is consistent with functional constraints, and the anterior series of longer centra are coincident with the major attachments of femoral musculature. This pattern is not present in many basal taxa and may have evolved independently in different dinosaurian groups, further suggesting functional importance.

Lower cretaceous avian-dominated, theropod, thyreophoran, pterosaur and turtle track assemblages from the Tugulu Group, Xinjiang, China: ichnotaxonomy and palaeoecology.

Rich tetrapod ichnofaunas, known for more than a decade, from the Huangyangquan Reservoir (Wuerhe District, Karamay City, Xinjiang) have been an abundant source of some of the largest Lower Cretaceous track collections from China. They originate from inland lacustrine clastic exposures of the 581-877 m thick Tugulu Group, variously divided into four formations and subgroups in the northwestern margin of the Junggar Basin. The large Huangyangquan track assemblages occur in the Lower layer/Subgroup II. Similarly-composed track assemblages also occur at the smaller Asphaltite site in the Upper Layer/Subgroup III. The Huangyangquan assemblages have yielded more than 1,500 identified tracks including abundant tracks of avian and non-avian theropods, pterosaurs and turtles and less abundant tracks of stegosaurs. Previous avian track identifications have been reassessed to conclude that Moguiornipes robustus is a taphotaxon and Koreanaornis dodsoni might be better accommodated in the ichnogenus Aquatilavipes which appears to be the dominant avian ichnotaxon. The avian track Ignotornis is also recognized and represents the first occurrence of this ichnogenus in China. Although the Huangyangquan assemblages lack some of the larger components (e.g., sauropodan and ornithopodan tracks) known from other Lower Cretaceous localities, the association of abundant tracks of smaller tetrapods (avian and non-avian theropods, pterosaurs and turtles) appears to be representative of lacustrine basin faunas of this region, and are an excellent example of the shorebird ichnocoenosis/ichnofacies concept. This is the first comprehensive review and re-analysis of an important Lower Cretaceous ecosystem.

An Older and Exceptionally Large Adult Specimen of Tyrannosaurus rex.

Here we describe an extremely large and relatively complete (roughly 65%) skeleton of Tyrannosaurus rex (RSM P2523.8). Multiple measurements (including those of the skull, hip, and limbs) show that RSM P2523.8 was a robust individual with an estimated body mass exceeding all other known T. rex specimens and representatives of all other gigantic terrestrial theropods. Histological sampling of the fibula confirms that RSM P2523.8 is skeletally mature. The prevalence of incompletely coossified elements contradicts previous assertions that such unfused elements can be taken as indicators of somatic immaturity. As an extreme example of both ontogenetic maturity and osteological robustness, RSM P2523.8 offers support for prior hypotheses that a sampling bias occurs throughout the Dinosauria, making it likely that most taxa grew to significantly greater size than current known specimens indicate. Anat Rec, 303:656-672, 2020. © 2019 Wiley Periodicals, Inc.

Feather evolution exemplifies sexually selected bridges across the adaptive landscape.

Over the last two decades, paleontologists have pieced together the early evolutionary history of feathers. Simple hair-like feathers served as insulating pelage, but the first feathers with complex branching structures and a plainer form evolved for the purpose of sexual display. The evolution of these complex display feathers was essential to the later evolution of flight. Feathers illustrate how sexual selection can generate complex novel phenotypes, which are then available for natural selection to modify and direct toward novel functions. In the longstanding metaphor of the adaptive landscape, sexual selection is a means by which lineages resting on one adaptive peak may gradually bridge a gap to another peak, without the landscape itself being first altered by environmental changes.

Response to: Phylogenetic placement, developmental trajectories and evolutionary implications of a feathered dinosaur tail in Mid-Cretaceous amber.

In his correspondence, Markus Lambertz [1] raises some concerns about the phylogenetic placement and feather development of DIP-V-15103, the amber-entombed tail section that we recently reported [2] as fragmentary remains of a non-pygostylian coelurosaur (likely within the basal part of Coelurosauria). We here would like to respond to these concerns.

An approach to scoring cursorial limb proportions in carnivorous dinosaurs and an attempt to account for allometry.

From an initial dataset of 53 theropod species, the general relationship between theropod lower-leg length and body mass is identified. After factoring out this allometric relationship, theropod hindlimb proportions are assessed irrespective of body mass. Cursorial-limb-proportion (CLP) scores derived for each of the considered theropod taxa offer a measure of the extent to which a particular species deviates in favour of higher or lower running speeds. Within the same theropod species, these CLP scores are found to be consistent across multiple adult specimens and across disparate ontogenetic stages. Early theropods are found to have low CLP scores, while the coelurosaurian tyrannosauroids and compsognathids are found to have high CLP scores. Among deinonychosaurs, troodontids have consistently high CLP scores, while many dromaeosaur taxa, including Velociraptor and Deinonychus, have low CLP scores. This indicates that dromaeosaurs were not, overall, a particularly cursorily adapted group. Comparisons between the CLP scores of Tyrannosaurus and specimens referred to the controversial genus Nanotyrannus indicate a strong discrepancy in cursorial adaptations, which supports the legitimacy of Nanotyrannus and the previous suggestions of ecological partitioning between Nanotyrannus and the contemporaneous Tyrannosaurus.

Digit-only sauropod pes trackways from China--evidence of swimming or a preservational phenomenon?

For more than 70 years unusual sauropod trackways have played a pivotal role in debates about the swimming ability of sauropods. Most claims that sauropods could swim have been based on manus-only or manus-dominated trackways. However none of these incomplete trackways has been entirely convincing, and most have proved to be taphonomic artifacts, either undertracks or the result of differential depth of penetration of manus and pes tracks, but otherwise showed the typical pattern of normal walking trackways. Here we report an assemblage of unusual sauropod tracks from the Lower Cretaceous Hekou Group of Gansu Province, northern China, characterized by the preservation of only the pes claw traces, that we interpret as having been left by walking, not buoyant or swimming, individuals. They are interpreted as the result of animals moving on a soft mud-silt substrate, projecting their claws deeply to register their traces on an underlying sand layer where they gained more grip during progression. Other sauropod walking trackways on the same surface with both pes and manus traces preserved, were probably left earlier on relatively firm substrates that predated the deposition of soft mud and silt . Presently, there is no convincing evidence of swimming sauropods from their trackways, which is not to say that sauropods did not swim at all.

A Feathered Dinosaur Tail with Primitive Plumage Trapped in Mid-Cretaceous Amber.

In the two decades since the discovery of feathered dinosaurs [1-3], the range of plumage known from non-avialan theropods has expanded significantly, confirming several features predicted by developmentally informed models of feather evolution [4-10]. However, three-dimensional feather morphology and evolutionary patterns remain difficult to interpret, due to compression in sedimentary rocks [9, 11]. Recent discoveries in Cretaceous amber from Canada, France, Japan, Lebanon, Myanmar, and the United States [12-18] reveal much finer levels of structural detail, but taxonomic placement is uncertain because plumage is rarely associated with identifiable skeletal material [14]. Here we describe the feathered tail of a non-avialan theropod preserved in mid-Cretaceous (∼99 Ma) amber from Kachin State, Myanmar [17], with plumage structure that directly informs the evolutionary developmental pathway of feathers. This specimen provides an opportunity to document pristine feathers in direct association with a putative juvenile coelurosaur, preserving fine morphological details, including the spatial arrangement of follicles and feathers on the body, and micrometer-scale features of the plumage. Many feathers exhibit a short, slender rachis with alternating barbs and a uniform series of contiguous barbules, supporting the developmental hypothesis that barbs already possessed barbules when they fused to form the rachis [19]. Beneath the feathers, carbonized soft tissues offer a glimpse of preservational potential and history for the inclusion; abundant Fe2+ suggests that vestiges of primary hemoglobin and ferritin remain trapped within the tail. The new finding highlights the unique preservation potential of amber for understanding the morphology and evolution of coelurosaurian integumentary structures.

A possible instance of sexual dimorphism in the tails of two oviraptorosaur dinosaurs.

The hypothesis that oviraptorosaurs used tail-feather displays in courtship behavior previously predicted that oviraptorosaurs would be found to display sexually dimorphic caudal osteology. MPC-D 100/1002 and MPC-D 100/1127 are two specimens of the oviraptorosaur Khaan mckennai. Although similar in absolute size and in virtually all other anatomical details, the anterior haemal spines of MPC-D 100/1002 exceed those of MPC-D 100/1127 in ventral depth and develop a hitherto unreported "spearhead" shape. This dissimilarity cannot be readily explained as pathologic and is too extreme to be reasonably attributed to the amount of individual variation expected among con-specifics. Instead, this discrepancy in haemal spine morphology may be attributable to sexual dimorphism. The haemal spine form of MPC-D 100/1002 offers greater surface area for caudal muscle insertions. On this basis, MPC-D 100/1002 is regarded as most probably male, and MPC-D 100/1127 is regarded as most probably female.

An Ornithopod-Dominated Tracksite from the Lower Cretaceous Jiaguan Formation (Barremian-Albian) of Qijiang, South-Central China: New Discoveries, Ichnotaxonomy, Preservation and Palaeoecology.

The historically-famous Lotus Fortress site, a deep 1.5-3.0-meter-high, 200-meter-long horizonal notch high up in near-vertical sandstone cliffs comprising the Cretaceous Jiaguan Formation, has been known since the 13th Century as an impregnable defensive position. The site is also extraordinary for having multiple tetrapod track-bearing levels, of which the lower two form the floor of part of the notch, and yield very well preserved asseamblages of ornithopod, bird (avian theropod) and pterosaur tracks. Trackway counts indicate that ornithopods dominate (69%) accounting for at least 165 trackmakers, followed by bird (18%), sauropod (10%), and pterosaur (3%). Previous studies designated Lotus Fortress as the type locality of Caririchnium lotus and Wupus agilis both of which are recognized here as valid ichnotaxa. On the basis of multiple parallel trackways both are interpreted as representing the trackways of gregarious species. C. lotus is redescribed here in detail and interpreted to indicate two age cohorts representing subadults that were sometimes bipedal and larger quadrupedal adults. Two other previously described dinosaurian ichnospecies, are here reinterpreted as underprints and considered nomina dubia. Like a growing number of significant tetrapod tracksites in China the Lotus Fortress site reveals new information about the composition of tetrapod faunas from formations in which the skeletal record is sparse. In particular, the site shows the relatively high abundance of Caririchium in a region where saurischian ichnofaunas are often dominant. It is also the only site known to have yielded Wupus agilis. In combination with information from other tracksites from the Jiaguan formation and other Cretaceous formations in the region, the track record is proving increasingly impotant as a major source of information on the vertebrate faunas of the region. The Lotus Fortress site has been developed as a spectacular, geologically-, paleontologically- and a culturally-significant destination within Qijiang National Geological Park.

Piscivory in the feathered dinosaur Microraptor.

The largest specimen of the four-winged dromaeosaurid dinosaur Microraptor gui includes preserved gut contents. Previous reports of gut contents and considerations of functional morphology have indicated that Microraptor hunted in an arboreal environment. The new specimen demonstrates that this was not strictly the case, and offers unique insights into the ecology of nonavian dinosaurs early in the evolution of flight. The preserved gut contents are composed of teleost fish remains. Several morphological adaptations of Microraptor are identified as consistent with a partially piscivorous diet, including dentition with reduced serrations and forward projecting teeth on the anterior of the dentary. The feeding habits of Microraptor can now be understood better than that of any other carnivorous nonavian dinosaur, and Microraptor appears to have been an opportunistic and generalist feeder, able to exploit the most common prey in both the arboreal and aquatic microhabitats of the Early Cretaceous Jehol ecosystem.

Abdominal contents from two large early cretaceous compsognathids (Dinosauria: Theropoda) demonstrate feeding on confuciusornithids and dromaeosaurids.

Two skeletons of the large compsognathid Sinocalliopteryx gigas include intact abdominal contents. Both specimens come from the Jianshangou Beds of the lower Yixian Formation (Neocomian), Liaoning, China. The holotype of S. gigas preserves a partial dromaeosaurid leg in the abdominal cavity, here attributed to Sinornithosaurus. A second, newly-discovered specimen preserves the remains of at least two individuals of the primitive avialan, Confuciusornis sanctus, in addition to acid-etched bones from a possible ornithischian. Although it cannot be stated whether such prey items were scavenged or actively hunted, the presence of two Confuciusornis in a grossly similar state of digestion suggests they were consumed in rapid succession. Given the lack of clear arboreal adaptations in Sinocalliopteryx, we suggest it may have been an adept stealth hunter.

The tail of Tyrannosaurus: reassessing the size and locomotive importance of the M. caudofemoralis in non-avian theropods.

Unlike extant birds and mammals, most non-avian theropods had large muscular tails, with muscle arrangements similar to those of modern reptiles. Examination of ornithomimid and tyrannosaurid tails revealed sequential diagonal scarring on the lateral faces of four or more hemal spines that consistently correlates with the zone of the tail just anterior to the disappearance of the vertebral transverse processes. This sequential scarring is interpreted as the tapering boundary between the insertions of the M. caudofemoralis and the M. ilioischiocaudalis. Digital muscle reconstructions based on measurements of fossil specimens and dissections of modern reptiles showed that the M. caudofemoralis of many non-avian theropods was exceptionally large. These high caudofemoral mass estimates are consistent with the elevation of the transverse processes of the caudal vertebra above the centrum, which creates an enlarged hypaxial region. Dorsally elevated transverse processes are characteristic of even primitive theropods and suggest that a large M. caudofemoralis is a basal characteristic of the group. In the genus Tyrannosaurus, the mass of the M. caudofemoralis was further increased by dorsoventrally lengthening the hemal arches. The expanded M. caudofemoralis of Tyrannosaurus may have evolved as compensation for the animal's immense size. Because the M. caudofemoralis is the primary hind limb retractor, large M. caudofemoralis masses and the resulting contractile force and torque estimates presented here indicate a sizable investment in locomotive muscle among theropods with a range of body sizes and give new evidence in favor of greater athleticism, in terms of overall cursoriality, balance, and turning agility.

Dinosaur speed demon: the caudal musculature of Carnotaurus sastrei and implications for the evolution of South American abelisaurids.

In the South American abelisaurids Carnotaurus sastrei, Aucasaurus garridoi, and, to a lesser extent Skorpiovenator bustingorryi, the anterior caudal ribs project at a high dorsolateral inclination and have interlocking lateral tips. This unique morphology facilitated the expansion of the caudal hypaxial musculature at the expense of the epaxial musculature. Distinct ridges on the ventrolateral surfaces of the caudal ribs of Aucasaurus garridoi are interpreted as attachment scars from the intra caudofemoralis/ilio-ischiocaudalis septa, and confirm that the M. caudofemoralis of advanced South American abelisaurids originated from a portion of the caudal ribs. Digital muscle models indicate that, relative to its overall body size, Carnotaurus sastrei had a substantially larger M. caudofemoralis than any other theropod yet studied. In most non-avian theropods, as in many extant sauropsids, the M. caudofemoralis served as the primary femoral retractor muscle during the locomotive power stroke. This large investment in the M. caudofemoralis suggests that Carnotaurus sastrei had the potential for great cursorial abilities, particularly short-burst sprinting. However, the tightly interlocking morphology of the anterior caudal vertebrae implies a reduced ability to make tight turns. Examination of these vertebral traits in evolutionary context reveals a progressive sequence of increasing caudofemoral mass and tail rigidity among the Abelisauridae of South America.