Uncovering widespread Anthropocene dietary shifts in Chinese large mammalian herbivores.

The Anthropocene's human-dominated habitat expansion endangers global biodiversity. However, large mammalian herbivores experienced few extinctions during the 20th century, hinting at potentially overlooked ecological responses of a group sensitive to global change. Using dental microwear as a proxy, we studied large herbivore dietary niches over a century across mainland China before (1880s-1910s) and after (1970s-1990s) the human population explosion. We uncovered widespread and significant shifts (interspecific microwear differences increased and intraspecific microwear dispersion expanded) within dietary niches linked to geographical areas with rapid industrialization and population growth in eastern China. By contrast, in western China, where human population growth was slower, we found no indications of shifts in herbivore dietary niches. Further regression analysis links the intensity of microwear changes to human land-use expansion. These analyses highlight dietary adjustments of large herbivores as a likely key factor in their adaptation across a century of large-scale human-driven changes.

Intra-gastric phytoliths provide evidence for folivory in basal avialans of the Early Cretaceous Jehol Biota.

Angiosperms became the dominant plant group in early to middle Cretaceous terrestrial ecosystems, coincident with the timing of the earliest pulse of bird diversification. While living birds and angiosperms exhibit strong interactions across pollination/nectivory, seed dispersal/frugivory, and folivory, documentation of the evolutionary origins and construction of that ecological complexity remains scarce in the Mesozoic. Through the first study of preserved in situ dietary derived phytoliths in a nearly complete skeleton of the early diverging avialan clade Jeholornithidae, we provide direct dietary evidence that Jeholornis consumed leaves likely from the magnoliid angiosperm clade, and these results lend further support for early ecological connections among the earliest birds and angiosperms. The broad diet of the early diverging avialan Jeholornis including at least fruits and leaves marks a clear transition in the early evolution of birds in the establishment of an arboreal (angiosperm) herbivore niche in the Early Cretaceous occupied largely by birds today. Morphometric reanalysis of the lower jaw of Jeholornis further supports a generalized morphology shared with other herbivorous birds, including an extant avian folivore, the hoatzin.

Decoupling the skull and skeleton in a Cretaceous bird with unique appendicular morphologies.

The Cretaceous is a critical time interval that encompasses explosive diversifications of terrestrial vertebrates, particularly the period when the earliest-branching birds, after divergence from their theropod ancestors, evolved the characteristic avian Bauplan that led eventually to their global radiation. This early phylogenetic diversity is overwhelmed by the Ornithothoraces, consisting of the Enantiornithes and Ornithuromorpha, whose members evolved key derived features of crown birds. This disparity consequently circumscribes a large morphological gap between these derived clades and the oldest bird Archaeopteryx. The non-ornithothoracine pygostylians, with an intermediate phylogenetic position, are key to deciphering those evolutionary transformations, but progress in their study has been hampered by the limited diversity of known fossils. Here we report an Early Cetaceous non-ornithothoracine pygostylian, Cratonavis zhui gen. et sp. nov., that exhibits a unique combination of a non-avialan dinosaurian akinetic skull with an avialan post-cranial skeleton, revealing the key role of evolutionary mosaicism in early bird diversification. The unusually elongated scapular and metatarsal one preserved in Cratonavis highlights a breadth of skeletal plasticity, stemming from their distinct developmental modules and selection for possibly raptorial behaviour. Mapped changes in these two elements across theropod phylogeny demonstrate clade-specific evolutionary lability.

Insight into the evolutionary assemblage of cranial kinesis from a Cretaceous bird.

The independent movements and flexibility of various parts of the skull, called cranial kinesis, are an evolutionary innovation that is found in living vertebrates only in some squamates and crown birds and is considered to be a major factor underpinning much of the enormous phenotypic and ecological diversity of living birds, the most diverse group of extant amniotes. Compared to the postcranium, our understanding of the evolutionary assemblage of the characteristic modern bird skull has been hampered by sparse fossil records of early cranial materials, with competing hypotheses regarding the evolutionary development of cranial kinesis among early members of the avialans. Here, a detailed three-dimensional reconstruction of the skull of the Early Cretaceous enantiornithine Yuanchuavis kompsosoura allows for its in-depth description, including elements that are poorly known among early-diverging avialans but are central to deciphering the mosaic assembly of features required for modern avian cranial kinesis. Our reconstruction of the skull shows evolutionary and functional conservation of the temporal and palatal regions by retaining the ancestral theropod dinosaurian configuration within the skull of this otherwise derived and volant bird. Geometric morphometric analysis of the palatine suggests that loss of the jugal process represents the first step in the structural modifications of this element leading to the kinetic crown bird condition. The mixture of plesiomorphic temporal and palatal structures together with a derived avialan rostrum and postcranial skeleton encapsulated in Yuanchuavis manifests the key role of evolutionary mosaicism and experimentation in early bird diversification.

Low dinosaur biodiversity in central China 2 million years prior to the end-Cretaceous mass extinction.

Whether or not nonavian dinosaur biodiversity declined prior to the end-Cretaceous mass extinction remains controversial as the result of sampling biases in the fossil record, differences in the analytical approaches used, and the rarity of high-precision geochronological dating of dinosaur fossils. Using magnetostratigraphy, cyclostratigraphy, and biostratigraphy, we establish a high-resolution geochronological framework for the fossil-rich Late Cretaceous sedimentary sequence in the Shanyang Basin of central China. We have found only three dinosaurian eggshell taxa (Macroolithus yaotunensis, Elongatoolithus elongatus, and Stromatoolithus pinglingensis) representing two clades (Oviraptoridae and Hadrosauridae) in sediments deposited between ∼68.2 and ∼66.4 million y ago, indicating sustained low dinosaur biodiversity, and that assessment is consistent with the known skeletal remains in the Shanyang and surrounding basins of central China. Along with the dinosaur eggshell records from eastern and southern China, we find a decline in dinosaur biodiversity from the Campanian to the Maastrichtian. Our results support a long-term decline in global dinosaur biodiversity prior to 66 million y ago, which likely set the stage for the end-Cretaceous nonavian dinosaur mass extinction.

Early evolution of diurnal habits in owls (Aves, Strigiformes) documented by a new and exquisitely preserved Miocene owl fossil from China.

SignificanceOwls, with their largely nocturnal habits, contrast strikingly with the vast majority of diurnal birds. A new spectacular late Miocene owl skeleton from China unexpectedly preserves the oldest evidence for daytime behavior in owls. The extinct owl is a member of the clade Surniini, which contains most living diurnal owl species. Analysis of the preserved eye bones documents them as consistent with diurnal birds, and phylogenetically constrained character mapping coincides with a reconstruction of an early evolutionary reversal away from nocturnal habits in this owl group. These results support a potential Miocene origin of nonnocturnal habits in a globally distributed owl group, which may be linked to steppe habitat expansion and climatic cooling in the late Miocene.

Novel evolution of a hyper-elongated tongue in a Cretaceous enantiornithine from China and the evolution of the hyolingual apparatus and feeding in birds.

The globally distributed extinct clade Enantiornithes comprises the most diverse early radiation of birds in the Mesozoic with species exhibiting a wide range of body sizes, morphologies, and ecologies. The fossil of a new enantiornithine bird, Brevirostruavis macrohyoideus gen. et sp. nov., from the Lower Cretaceous Jiufotang Formation in Liaoning Province, northeastern China, preserves a few important skeletal features previously unknown among early stem and extant birds, including an extremely elongate bony hyoid element (only slightly shorter than the skull), combined with a short cranial rostrum. The long hyoid provides direct evidence for the evolution of specialized feeding in this extinct species, and appears similar to the highly mobile tongue that is mobilized by the paired epibranchials present in living hummingbirds, honeyeaters, and woodpeckers. The likely linkage between food acquisition and tongue protrusion might have been a key factor in the independent evolution of particularly elongate hyobranchials in early birds.

The evolutionary and functional implications of the unusual quadrate of Longipteryx chaoyangensis (Avialae: Enantiornithes) from the Cretaceous Jehol Biota of China.

While the morphology and evolution of the quadrate among early birds and through the evolutionary origin of birds is not well known, we add to knowledge about that past diversity through description of the morphology of the quadrate in the unusually elongate skull of the Cretaceous enantiornithine bird Longipteryx chaoyangensis. The lateral and caudal surfaces of the quadrate are well exposed in two specimens revealing morphologies typical of early birds and their dinosaurian close relatives like a small otic head and two mandibular condyles. However, both skeletons exhibit quadrates with a unique, enlarged lateral crest that has not been previously described among Mesozoic birds. It is possible that the rostral surface of this lateral expansion served as the origination site for enlarged jaw musculature in a manner similar to the enlarged subcapitular tubercle in extant galloanserine birds. The caudally concave surface of the quadrate likely reflects some aspect of cranial pneumaticity, with its shape and position reminiscent of quadrates found in close non-avialan maniraptoran relatives. It is possible that this lateral crest has a wider distribution among enantiornithines and other early birds and that the crest has been misidentified as the orbital process in some more damaged specimens. In addition, the enlarged lateral mandibular condyle (relative to the medial condyle) differs from the condition typically reported among enantiornithines and could indicate a difference in jaw function or mechanics in this bird with an elongated rostrum, or simply misinterpretations of morphology. Further examination of the quadrate in temporally early and phylogenetically stemward birds, along with their close outgroups, could greatly impact the study of several different aspects of bird biology including assessment of phylogenetic relationships, interpretation of the function and kinematics of the skull, reconstruction of foraging paleoecology, and evolution of skull morphological diversity among Mesozoic birds.

Cretaceous bird with dinosaur skull sheds light on avian cranial evolution.

The transformation of the bird skull from an ancestral akinetic, heavy, and toothed dinosaurian morphology to a highly derived, lightweight, edentulous, and kinetic skull is an innovation as significant as powered flight and feathers. Our understanding of evolutionary assembly of the modern form and function of avian cranium has been impeded by the rarity of early bird fossils with well-preserved skulls. Here, we describe a new enantiornithine bird from the Early Cretaceous of China that preserves a nearly complete skull including the palatal elements, exposing the components of cranial kinesis. Our three-dimensional reconstruction of the entire enantiornithine skull demonstrates that this bird has an akinetic skull indicated by the unexpected retention of the plesiomorphic dinosaurian palate and diapsid temporal configurations, capped with a derived avialan rostrum and cranial roof, highlighting the highly modular and mosaic evolution of the avialan skull.

Earliest fossils of giant-sized bony-toothed birds (Aves: Pelagornithidae) from the Eocene of Seymour Island, Antarctica.

While pelagornithid or 'bony-toothed' bird fossils representing multiple species are known from Antarctica, a new dentary fragment of a pelagornithid bird from the middle Eocene Submeseta Formation on Seymour Island, Antarctica represents a species with a body size on par with the largest known species in the clade. Measurements from the partial 'toothed' dentary point to a giant body size for the species, although the spacing among the pseudoteeth differs from that published for other pelagornithids. The discrepancy might suggest that previous techniques are not adequate for examination of incomplete material or that another factor such as phylogeny might impact size estimates and comparisons. Combined with a revised stratigraphic position in the early Eocene La Meseta Formation on Seymour Island for the largest pelagornithid tarsometatarsus known, these Antarctic fossils demonstrate the early evolution of giant body size in the clade (by ~ 50 Ma), and they likely represent not only the largest flying birds of the Eocene but also some of the largest volant birds that ever lived (with an estimated 5-6 m wingspan). Furthermore, the distribution of giant-sized pelagornithid fossils across more than 10 million years of Antarctic geological deposits points to a prolonged survival of giant-sized pelagornithids within the southern seas, and their success as a pelagic predatory component of marine and coastal ecosystems alongside early penguins.

A new clade of basal Early Cretaceous pygostylian birds and developmental plasticity of the avian shoulder girdle.

Early members of the clade Pygostylia (birds with a short tail ending in a compound bone termed "pygostyle") are critical for understanding how the modern avian bauplan evolved from long-tailed basal birds like Archaeopteryx However, the currently limited known diversity of early branching pygostylians obscures our understanding of this major transition in avian evolution. Here, we describe a basal pygostylian, Jinguofortis perplexus gen. et sp. nov., from the Early Cretaceous of China that adds important information about early members of the short-tailed bird group. Phylogenetic analysis recovers a clade (Jinguofortisidae fam. nov.) uniting Jinguofortis and the enigmatic basal avian taxon Chongmingia that represents the second earliest diverging group of the Pygostylia. Jinguofortisids preserve a mosaic combination of plesiomorphic nonavian theropod features such as a fused scapulocoracoid (a major component of the flight apparatus) and more derived flight-related morphologies including the earliest evidence of reduction in manual digits among birds. The presence of a fused scapulocoracoid in adult individuals independently evolved in Jinguofortisidae and Confuciusornithiformes may relate to an accelerated osteogenesis during chondrogenesis and likely formed through the heterochronic process of peramorphosis by which these basal taxa retain the scapulocoracoid of the nonavian theropod ancestors with the addition of flight-related modifications. With wings having a low aspect ratio and wing loading, Jinguofortis may have been adapted particularly to dense forest environments. The discovery of Jinguofortis increases the known ecomorphological diversity of basal pygostylians and highlights the importance of developmental plasticity for understanding mosaic evolution in early birds.

Vocal specialization through tracheal elongation in an extinct Miocene pheasant from China.

Modifications to the upper vocal tract involving hyper-elongated tracheae have evolved many times within crown birds, and their evolution has been linked to a 'size exaggeration' hypothesis in acoustic signaling and communication, whereby smaller-sized birds can produce louder sounds. A fossil skeleton of a new extinct species of wildfowl (Galliformes: Phasianidae) from the late Miocene of China, preserves an elongated, coiled trachea that represents the oldest fossil record of this vocal modification in birds and the first documentation of its evolution within pheasants. The phylogenetic position of this species within Phasianidae has not been fully resolved, but appears to document a separate independent origination of this vocal modification within Galliformes. The fossil preserves a coiled section of the trachea and other remains supporting a tracheal length longer than the bird's body. This extinct species likely produced vocalizations with a lower fundamental frequency and reduced harmonics compared to similarly-sized pheasants. The independent evolution of this vocal feature in galliforms living in both open and closed habitats does not appear to be correlated with other factors of biology or its open savanna-like habitat. Features present in the fossil that are typically associated with sexual dimorphism suggest that sexual selection may have resulted in the evolution of both the morphology and vocalization mechanism in this extinct species.

Early Paleocene landbird supports rapid phylogenetic and morphological diversification of crown birds after the K-Pg mass extinction.

Evidence is accumulating for a rapid diversification of birds following the K-Pg extinction. Recent molecular divergence dating studies suggest that birds radiated explosively during the first few million years of the Paleocene; however, fossils from this interval remain poorly represented, hindering our understanding of morphological and ecological specialization in early neoavian birds. Here we report a small fossil bird from the Nacimiento Formation of New Mexico, constrained to 62.221-62.517 Ma. This partial skeleton represents the oldest arboreal crown group bird known. Phylogenetic analyses recovered Tsidiiyazhi abini gen. et sp. nov. as a member of the Sandcoleidae, an extinct basal clade of stem mousebirds (Coliiformes). The discovery of Tsidiiyazhi pushes the minimum divergence ages of as many as nine additional major neoavian lineages into the earliest Paleocene, compressing the duration of the proposed explosive post-K-Pg radiation of modern birds into a very narrow temporal window parallel to that suggested for placental mammals. Simultaneously, Tsidiiyazhi provides evidence for the rapid morphological (and likely ecological) diversification of crown birds. Features of the foot indicate semizygodactyly (the ability to facultatively reverse the fourth pedal digit), and the arcuate arrangement of the pedal trochleae bears a striking resemblance to the conformation in owls (Strigiformes). Inclusion of fossil taxa and branch length estimates impacts ancestral state reconstructions, revealing support for the independent evolution of semizygodactyly in Coliiformes, Leptosomiformes, and Strigiformes, none of which is closely related to extant clades exhibiting full zygodactyly.

A late Paleocene probable metatherian (?deltatheroidan) survivor of the Cretaceous mass extinction.

Deltatheroidans are primitive metatherian mammals (relatives of marsupials), previously thought to have become extinct during the Cretaceous mass extinction. Here, we report a tiny new deltatheroidan mammal (Gurbanodelta kara gen. et sp. nov.) discovered at the South Gobi locality in China (Xinjiang Province) that is the first Cenozoic record of this clade and renders Deltatheroida a Lazarus taxon (with a new record 10 million years younger than their supposed extinction). The vertebrate fauna associated with Gurbanodelta is most similar to that from the slightly older late Paleocene Subeng locality in Inner Mongolia. The upper molars of Gurbanodelta exhibit a broad stylar shelf with one prominent cusp (stylocone), and a paracone that is sharp and significantly taller than the metacone. The lower molar tentatively assigned to Gurbanodelta has a very small talonid without an entoconid. This combination of these features is known only in deltatheroidans. Phylogenetic analysis places Gurbanodelta as the sister taxon of the North American latest Cretaceous Nanocuris. Gurbanodelta is the smallest-known deltatheroidan, and roughly the same size as the smallest living marsupial. It is likely that the Gurbanodelta lineage dispersed between Asia and North America as part of known intercontinental mammalian dispersals in the late Paleocene, or possibly earlier.

The palaeobiology of high latitude birds from the early Eocene greenhouse of Ellesmere Island, Arctic Canada.

Fossils attributable to the extinct waterfowl clade Presbyornithidae and the large flightless Gastornithidae from the early Eocene (~52-53 Ma) of Ellesmere Island, in northernmost Canada are the oldest Cenozoic avian fossils from the Arctic. Except for its slightly larger size, the Arctic presbyornithid humerus is not distinguishable from fossils of Presbyornis pervetus from the western United States, and the Gastornis phalanx is within the known size range of mid-latitude individuals. The occurrence of Presbyornis above the Arctic Circle in the Eocene could be the result of annual migration like that of its living duck and geese relatives, or it may have been a year-round resident similar to some Eocene mammals on Ellesmere and some extant species of sea ducks. Gastornis, along with some of the mammalian and reptilian members of the Eocene Arctic fauna, likely over-wintered in the Arctic. Despite the milder (above freezing) Eocene climate on Ellesmere Island, prolonged periods of darkness occurred during the winter. Presence of these extinct birds at both mid and high latitudes on the northern continents provides evidence that future increases in climatic warming (closer to Eocene levels) could lead to the establishment of new migratory or resident populations within the Arctic Circle.

A pelican tarsometatarsus (Aves: Pelecanidae) from the latest Pliocene Siwaliks of India.

We report a new fossil specimen of a pelican from the Tatrot Formation of the Siwalik Hills, India. It likely represents Pelecanus sivalensis Davies, 1880, the smaller of the two previously published species from the Siwalik Group stratigraphic sequence. This complete tarsometatarsus is the first fossil bone of a pelican collected in India for over 100 years. It is from the latest Pliocene (∼2.6 Ma), and is the youngest pelican fossil from the region. The new specimen exhibits a derived distoplantar 'slant' to the plantar margin of the medial crest of the hypotarsus, and a combination of features related to the morphology of the hypotarsus, the distal foramen, trochleae, and overall size that allow further differentiation from known tarsometatarsi of fossil and extant pelicans, including the three species of extant pelicans that occur in India (Pelecanus crispus, P. onocrotalus, and P. philippensis). It is of appropriate size for Pelecanus sivalensis, which to date has been known only by fragments of other skeletal elements of the wing, leg, and shoulder girdle. Thus, the observation that this tarsometatarsus is morphologically distinct from those of known pelicans provides further support for the distinctiveness of at least one extinct species of pelican from the Siwalik Group sediments. While the morphology of the tarsometatarsus allows for separation from other taxa known from tarsometatarsi, we found no clear shared derived states to place this taxon with any confidence in a phylogenetic context relative to any other pelican species, or even determine if it is part of the crown group of Pelecanidae. However, published molecular data are consistent with an origin of the crown clade prior to the Pleistocene, suggesting (along with one morphological character) the possibility that this species belongs to the Old World clade of pelican species.

Morphology of the quadrate in the Eocene anseriform Presbyornis and extant galloanserine birds.

Despite the notoriety, phylogenetic significance, and large number of available specimens of Presbyornis, its cranial anatomy has never been studied in detail, and its quadrate has been partly misinterpreted. We studied five quadrates of Presbyornis that reveal features hitherto unknown in the anseriforms but otherwise present in galliforms. As a result, we analyzed the variable quadrate characters among all extant galloanserine families and identified synapomorphies and other morphological variation among the major galloanserine clades. In terms of quadrate morphology, Presbyornis is more plesiomorphic than any extant anseriform (including the Anhimidae) and shares ancestral galloanserine characters with the Megapodiidae, the earliest branch of extant galliforms. The quadrate's morphology is inconsistent with the currently accepted anseriform phylogeny that nests Presbyornis within the crown-group as a close relative of the Anatidae. The presbyornithid quadrates exhibit an unusual variation in the presence of a caudomedial pneumatic foramen, which we interpret as a result of a discontinuous change in the growth path of the pneumatic diverticulum. Another episode of morphogenetic imbalance in the growth path of the pneumatic diverticulum may have accompanied the disappearance of the basiorbital pneumatic foramen (along with the pneumatization of the pterygoid) at the origin of the crown-group anseriforms. This episode is marked by the striking individual variation in the presence and location of pneumatic foramina in the mandibular part of the quadrate in the Anhimidae.