Survival of Theriosuchus (Mesoeucrocodylia: Atoposauridae) in a Late Cretaceous archipelago: a new species from the Maastrichtian of Romania

Small terrestrial non-eusuchian mesoeucrocodylians are common components of Cretaceous assemblages of Gondwanan provinces with notosuchians and araripesuchids as flagship taxa in South America, Africa and Madagascar, well into the Late Cretaceous. On the other hand, these are exceedingly rare in Laurasian landmasses during the Late Cretaceous. Small terrestrial mesoeucrocodylians from Europe were often referred to the genus Theriosuchus, a taxon with stratigraphic range extending from the Late Jurassic to the late Early Cretaceous. Theriosuchus is abundantly reported from various European localities, although Asiatic and possibly North American members are also known. It has often been closely associated with the first modern crocodilians, members of the Eusuchia, because of the presence of procoelous vertebrae, a widespread key character diagnosing the Eusuchia. Nevertheless, the relationships of Theriosuchus have not been explored in detail although one species, Theriosuchus pusillus, has been extensively described and referred in numerous works. Here, we describe a new basal mesoeucrocodylian, Theriosuchus sympiestodon sp. nov. from the Maastrichtian of the Haţeg Basin, Romania, suggesting a large temporal gap (about 58 myr) in the fossil record of the genus. Inclusion of the new taxon, along with Theriosuchus guimarotae, in a phylogenetic analysis confirms its referral to the genus Theriosuchus, within a monophyletic atoposaurid clade. Although phylogenetic resolution within this clade is still poor, the new taxon appears, on morphological grounds, to be most closely related to T. pusillus. The relationships of Atoposauridae within Mesoeucrocodylia and especially to Neosuchia are discussed in light of the results of the present contribution as well as from recent work. Our results raise the possibility that Atoposauridae might not be regarded as a derived neosuchian clade anymore, although further investigation of the neosuchian interrelationships is needed. Reports of isolated teeth referable to a closely related taxon from the Upper Cretaceous of Romania and France, together with the presence of Doratodon and Ischyrochampsa, indicate a previously unsuspected diverse assemblage of non-eusuchian mesoeucrocodylians in the Late Cretaceous European archipelago.     

The first definitive Asian spinosaurid (Dinosauria: Theropoda) from the early cretaceous of Laos

Spinosaurids are among the largest and most specialized carnivorous dinosaurs. The morphology of their crocodile-like skull, stomach contents, and oxygen isotopic composition of the bones suggest they had a predominantly piscivorous diet. Even if close relationships between spinosaurids and Middle Jurassic megalosaurs seem well established, very little is known about the transition from a generalized large basal tetanuran to the specialized morphology of spinosaurids. Spinosaurid remains were previously known from the Early to Late Cretaceous of North Africa, Europe, and South America. Here, we report the discovery of a new spinosaurid theropod from the late Early Cretaceous Savannakhet Basin in Laos, which is distinguished by an autapomorphic sinusoidal dorsosacral sail. This new taxon, Ichthyovenator laosensis gen. et sp. nov., includes well-preserved and partially articulated postcranial remains. Although possible spinosaurid teeth have been reported from various Early Cretaceous localities in Asia, the new taxon I. laosensis is the first definite record of Spinosauridae from Asia. Cladistic analysis identifies Ichthyovenator as a member of the sub-clade Baryonychinae and suggests a widespread distribution of this clade at the end of the Early Cretaceous. Chilantaisaurus tashouikensis from the Cretaceous of Inner Mongolia, and an ungual phalanx from the Upper Jurassic of Colorado are also referred to spinosaurids, extending both the stratigraphical and geographical range of this clade.     

A new phylogeny for basal Trechnotheria and Cladotheria and affinities of South American endemic Late Cretaceous mammals

The endemic South American mammals Meridiolestida, considered previously as dryolestoid cladotherians, are found to be non-cladotherian trechnotherians related to spalacotheriid symmetrodontans based on a parsimony analysis of 137 morphological characters among 44 taxa. Spalacotheriidae is the sister taxon to Meridiolestida, and the latter clade is derived from a primitive spalacolestine that migrated to South America from North America at the beginning of the Late Cretaceous. Meridiolestida survived until the early Paleocene (Peligrotherium) and early Miocene (Necrolestes) in South America, and their extinction is probably linked to the increasing competition with metatherian and eutherian tribosphenic mammals. The clade Meridiolestida plus Spalacotheriidae is the sister taxon to Cladotheria and forms a new clade Alethinotheria. Alethinotheria and its sister taxon Zhangheotheria, new clade (Zhangheotheriidae plus basal taxa), comprise Trechnotheria. Cladotheria is divided into Zatheria (plus stem taxa, including Amphitherium) and Dryolestida, including Dryolestidae and a paraphyletic array of basal dryolestidans (formerly classified as “Paurodontidae”). The South American Vincelestes and Groebertherium are basal dryolestidans.     

The oldest known tracks of web-footed birds from the Lower Cretaceous of South Korea

 We describe the oldest tracks of web-footed birds from the Early Cretaceous in South Korea. The tracks are characterized by a wide divarication angle and a long reversed hallux. The web is semipalmate and restricted to the proximal portion of the three forward digits. The tracks from the Early Cretaceous in South Korea are smaller than those of the Late Cretaceous, therefore confirming the trend of size increasing in the early evolution of birds as shown by skeletal fossils. The discovery of web-footed tracks with abundant non-web-footed tracks indicates that there was a considerable diversification of shore birds as early as the Early Cretaceous. Received: 4 November 1999 / Accepted in revised form: 28 February 2000     

First complete sauropod dinosaur skull from the Cretaceous of the Americas and the evolution of sauropod dentition

Sauropod dinosaur bones are common in Mesozoic terrestrial sediments, but sauropod skulls are exceedingly rare—cranial materials are known for less than one third of sauropod genera and even fewer are known from complete skulls. Here we describe the first complete sauropod skull from the Cretaceous of the Americas, Abydosaurus mcintoshi, n. gen., n. sp., known from 104.46 ± 0.95 Ma (megannum) sediments from Dinosaur National Monument, USA. Abydosaurus shares close ancestry with Brachiosaurus, which appeared in the fossil record ca. 45 million years earlier and had substantially broader teeth. A survey of tooth shape in sauropodomorphs demonstrates that sauropods evolved broad crowns during the Early Jurassic but did not evolve narrow crowns until the Late Jurassic, when they occupied their greatest range of crown breadths. During the Cretaceous, brachiosaurids and other lineages independently underwent a marked diminution in tooth breadth, and before the latest Cretaceous broad-crowned sauropods were extinct on all continental landmasses. Differential survival and diversification of narrow-crowned sauropods in the Late Cretaceous appears to be a directed trend that was not correlated with changes in plant diversity or abundance, but may signal a shift towards elevated tooth replacement rates and high-wear dentition. Sauropods lacked many of the complex herbivorous adaptations present within contemporaneous ornithischian herbivores, such as beaks, cheeks, kinesis, and heterodonty. The spartan design of sauropod skulls may be related to their remarkably small size—sauropod skulls account for only 1/200th of total body volume compared to 1/30th body volume in ornithopod dinosaurs.     

An abelisaurid from the Late Cretaceous of Egypt: implications for theropod biogeography

Recent paleogeographic scenarios postulate the isolation of continental Africa during the Late Cretaceous. The absence of abelisaurid theropods from Upper Cretaceous African strata was offered as support of hypothesized African isolation with the acknowledgement that the paucity of African abelisaurids may be mostly an issue of sampling. Here we report on a shed theropod tooth from the Upper Cretaceous (Maastrichtian, ∼70 Ma) Duwi Formation of Egypt. The tooth was referred to the Malagasy abelisaurid “Megalosaurus” crenatissimus (=Majungasaurus crenatissimus) in 1921. A discriminant function analysis was run to test for morphological congruence between the Egyptian tooth and the dentitions of 24 theropod taxa. The analysis correctly classified 96.6% of the teeth in the sample and assigned the tooth to Majungasaurus. As current paleogeographic reconstructions posit Madagascar had attained its current position relative to Africa before the Late Cretaceous, it is unlikely that the Egyptian tooth actually pertains to Majungasaurus. Nevertheless, its classification as an abelisaurid supports its referral to the clade. This tooth thus constitutes defensible evidence of an abelisaurid from the post-Cenomanian Cretaceous of mainland Africa. Combined with recent discoveries of abelisaurids in Niger and Morocco, the result indicates that Abelisauridae was a diverse group in Africa during the Cretaceous, existing in multiple places for at least ∼25 Ma and weakens support for hypotheses of an isolated Africa during the Late Cretaceous.Electronic Supplementary Material  Supplementary material is available for this article at

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The last polar dinosaurs: high diversity of latest Cretaceous arctic dinosaurs in Russia

A latest Cretaceous (68 to 65 million years ago) vertebrate microfossil assemblage discovered at Kakanaut in northeastern Russia reveals that dinosaurs were still highly diversified in Arctic regions just before the Cretaceous–Tertiary mass extinction event. Dinosaur eggshell fragments, belonging to hadrosaurids and non-avian theropods, indicate that at least several latest Cretaceous dinosaur taxa could reproduce in polar region and were probably year-round residents of high latitudes. Palaeobotanical data suggest that these polar dinosaurs lived in a temperate climate (mean annual temperature about 10°C), but the climate was apparently too cold for amphibians and ectothermic reptiles. The high diversity of Late Maastrichtian dinosaurs in high latitudes, where ectotherms are absent, strongly questions hypotheses according to which dinosaur extinction was a result of temperature decline, caused or not by the Chicxulub impact.     

The last “pelycosaur”: a varanopid synapsid from the <Emphasis Type="Italic">Pristerognathus</Emphasis> Assemblage Zone,Middle Permian of South Africa

We report on a partial varanopid skull and mandible from the Pristerognathus Assemblage Zone of the Beaufort Group, in the South African Karoo Basin, which is probably latest Middle Permian (Capitanian) in age. This mycterosaurine is not only the youngest known varanopid from the Southern Hemisphere, but it is also the youngest known “pelycosaur” (i.e., non-therapsid synapsid). Like all other members of this clade of hypercarnivores, the teeth are strongly flattened, recurved, and have finely serrated cutting edges. The anterior dentary teeth form a caniniform region, and the splenial features a foramen intermandibularis oralis, the first ever to be described in a “pelycosaur.” The last varanopids were the smallest carnivores of latest Middle Permian continental faunas. Occupation of the small carnivore guild appears to have allowed varanopids to achieve a nearly cosmopolitan distribution throughout the Middle Permian, between the great Early Permian radiation of basal synapsids and the spectacular diversification of therapsid synapsids in the Late Permian and Early Triassic.     

Passive and active margin history of the northern Tatricum (Western Carpathians, Slovakia)

The Tatricum, an upper crustal thrust sheet of the Central Western Carpathians, comprises pre-Alpine crystalline basement and a Late Paleozoic–Mesozoic sedimentary cover. The sedimentary record indicates gradual subsidence during the Triassic, Early Jurassic initial rifting, a Jurassic–Early Cretaceous extensional tectonic regime with episodic rifting events and thermal subsidence periods, and Middle Cretaceous overall flexural subsidence in front of the orogenic wedge prograding from the hinterland. Passive rifting led to the separation of the Central Carpathian realm from the North European Platform. A passive margin, rimmed by peripheral half-graben, was formed along the northern Tatric edge, facing the Vahic (South Penninic) oceanic domain. The passive versus active margin inversion occurred during the Senonian, when the Vahic ocean began to be consumed southwards below the Tatricum. It is argued that passive to active margin conversion is an integral part of the general shortening polarity of the Western Carpathians during the Mesozoic that lacks features of an independent Wilson cycle. An attempt is presented to explain all the crustal deformation by one principal driving force – the south-eastward slab pull generated by the subduction of the Meliatic (Triassic–Jurassic Tethys) oceanic lithosphere followed by the subcrustal subduction of the continental mantle lithosphere.     

The first multituberculate mammal from India

Mesozoic deposits of the former Gondwanaland are depauperate in early mammals, in general, and multituberculate mammals, in particular. Until now, the oldest multituberculate mammals known from the Gondwanan continents come from the Early Cretaceous of Morocco, NW Africa. Here, we report the presence of a new multituberculate mammal, Indobaatar zofiae gen. et sp. nov., from the Lower/Middle Jurassic Kota Formation, Pranhita-Godavari valley in peninsular India. This is the first record of a multituberculate from the Mesozoic rocks of India and possibly predates the oldest known multituberculates from Gondwanan continents. The new specimen, representing an upper premolar (P⁴), compares well with the upper premolar morphology of Eobaatariinae multituberculates known from the Early Cretaceous of Mongolia, China, England, and Spain. Together with the recent findings of cimolodontan multituberculates from the Early Cretaceous of Australia and Late Cretaceous of South America, the new discovery indicates a wide temporal and spatial distribution for multituberculate mammals in the former Gondwanaland.     

Digital preparation of a probable neoceratopsian preserved within an egg, with comments on microstructural anatomy of ornithischian eggshells

We describe the first known embryo of a neoceratopsian dinosaur, perhaps the most ubiquitous Laurasian group of Cretaceous dinosaurs, which is preserved completely enclosed within an egg. This specimen was collected from Late Cretaceous beds of southern Mongolia, which commonly preserve fossils of the neoceratopsian, Yamaceratops dorngobiensis. The small egg was scanned using high-resolution X-ray computed tomography and digitally prepared from the matrix. The preserved and imaged elements support a diagnosis of the embryo to Neoceratopsia and allow preliminary observations of ontogenetic transformations within this group. The addition of an embryo also adds another important data point to the already impressive postnatal ontogenetic series that are available for this clade.     

The first definitive carcharodontosaurid (Dinosauria: Theropoda) from Asia and the delayed ascent of tyrannosaurids

Little is known about the evolution of large-bodied theropod dinosaurs during the Early to mid Cretaceous in Asia. Prior to this time, Asia was home to an endemic fauna of basal tetanurans, whereas terminal Cretaceous ecosystems were dominated by tyrannosaurids, but the intervening 60 million years left a sparse fossil record. Here, we redescribe the enigmatic large-bodied Chilantaisaurus maortuensis from the Turonian of Inner Mongolia, China. We refer this species to a new genus, Shaochilong, and analyze its systematic affinities. Although Shaochilong has previously been allied with several disparate theropod groups (Megalosauridae, Allosauridae, Tyrannosauroidea, Maniraptora), we find strong support for a derived carcharodontosaurid placement. As such, Shaochilong is the first unequivocal Asian member of Carcharodontosauridae, which was once thought to be restricted to Gondwana. The discovery of an Asian carcharodontosaurid indicates that this clade was cosmopolitan in the Early to mid Cretaceous and that Asian large-bodied theropod faunas were no longer endemic at this time. It may also suggest that the ascent of tyrannosaurids into the large-bodied dinosaurian predator niche was a late event that occurred towards the end of the Cretaceous, between the Turonian and the Campanian. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Adaptations for marine habitat and the effect of Triassic and Jurassic predator pressure on development of decompression syndrome in ichthyosaurs

Decompression syndrome (caisson disease or the “the bends”) resulting in avascular necrosis has been documented in mosasaurs, sauropterygians, ichthyosaurs, and turtles from the Middle Jurassic to Late Cretaceous, but it was unclear that this disease occurred as far back as the Triassic. We have examined a large Triassic sample of ichthyosaurs and compared it with an equally large post-Triassic sample. Avascular necrosis was observed in over 15?% of Late Middle Jurassic to Cretaceous ichthyosaurs with the highest occurrence (18?%) in the Early Cretaceous, but was rare or absent in geologically older specimens. Triassic reptiles that dive were either physiologically protected, or rapid changes of their position in the water column rare and insignificant enough to prevent being recorded in the skeleton. Emergency surfacing due to a threat from an underwater predator may be the most important cause of avascular necrosis for air-breathing divers, with relative frequency of such events documented in the skeleton. Diving in the Triassic appears to have been a “leisurely” behavior until the evolution of large predators in the Late Jurassic that forced sudden depth alterations contributed to a higher occurrence of bends.     

鄂尔多斯盆地上古生界天然气的运移与聚集

通过对上古生界烃源岩天然气排气史、天然气运移方向和沉积相带的展布等综合分析 ,可将上古生界天然气的运移聚集分为三叠纪—早白垩世和早白垩世后两个阶段。在三叠纪—早白垩世阶段 ,天然气运移方向主要是向北 ,而在早白垩世后阶段 ,天然气主要是就近运移聚集而成藏 ,这一阶段是鄂尔多斯盆地上古生界天然气重要的运移与聚集期 ,故上古生界煤成气藏的成藏期较晚。鄂尔多斯盆地北部和中部的靖边古三角洲是天然气聚集的有利地区 ,具有较好的天然气勘探前景     

New carnivorous dinosaur from the Late Cretaceous of NW Patagonia and the evolution of abelisaurid theropods

A nearly complete skeleton of the new abelisaurid Skorpiovenator bustingorryi is reported here. The holotype was found in Late Cenomanian–Early Turonian outcrops of NW Patagonia, Argentina. This new taxon is deeply nested within a new clade of South American abelisaurids, named Brachyrostra. Within brachyrostrans, the skull shortening and hyperossification of the skull roof appear to be correlated with a progressive enclosure of the orbit, a set of features possibly related to shock-absorbing capabilities. Moreover, the development of horn-like structures and differential cranial thickening appear to be convergently acquired within Abelisauridae. Based on the similarities between Skorpiovenator and carcharodontosaurid tooth morphology, we suggest that isolated teeth originally referred as post-Cenomanian Carcharodontosauridae most probably belong to abelisaurids. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

A new clade of archaic large-bodied predatory dinosaurs (Theropoda: Allosauroidea) that survived to the latest Mesozoic

Non-avian theropod dinosaurs attained large body sizes, monopolising terrestrial apex predator niches in the Jurassic–Cretaceous. From the Middle Jurassic onwards, Allosauroidea and Megalosauroidea comprised almost all large-bodied predators for 85 million years. Despite their enormous success, however, they are usually considered absent from terminal Cretaceous ecosystems, replaced by tyrannosaurids and abelisaurids. We demonstrate that the problematic allosauroids Aerosteon, Australovenator, Fukuiraptor and Neovenator form a previously unrecognised but ecologically diverse and globally distributed clade (Neovenatoridae, new clade) with the hitherto enigmatic theropods Chilantaisaurus, Megaraptor and the Maastrichtian Orkoraptor. This refutes the notion that allosauroid extinction pre-dated the end of the Mesozoic. Neovenatoridae includes a derived group (Megaraptora, new clade) that developed long, raptorial forelimbs, cursorial hind limbs, appendicular pneumaticity and small size, features acquired convergently in bird-line theropods. Neovenatorids thus occupied a 14-fold adult size range from 175 kg (Fukuiraptor) to approximately 2,500 kg (Chilantaisaurus). Recognition of this major allosauroid radiation has implications for Gondwanan paleobiogeography: The distribution of early Cretaceous allosauroids does not strongly support the vicariant hypothesis of southern dinosaur evolution or any particular continental breakup sequence or dispersal scenario. Instead, clades were nearly cosmopolitan in their early history, and later distributions are explained by sampling failure or local extinction.     

A new Jurassic theropod from China documents a transitional step in the macrostructure of feathers

Genuine fossils with exquisitely preserved plumage from the Late Jurassic and Early Cretaceous of northeastern China have recently revealed that bird-like theropod dinosaurs had long pennaceous feathers along their hindlimbs and may have used their four wings to glide or fly. Thus, it has been postulated that early bird flight might initially have involved four wings (Xu et al. Nature 421:335–340, 2003; Hu et al. Nature 461:640–643, 2009; Han et al. Nat Commun 5:4382, 2014). Here, we describe Serikornis sungei gen. et sp. nov., a new feathered theropod from the Tiaojishan Fm (Late Jurassic) of Liaoning Province, China. Its skeletal morphology suggests a ground-dwelling ecology with no flying adaptations. Our phylogenetic analysis places Serikornis, together with other Late Jurassic paravians from China, as a basal paravians, outside the Eumaniraptora clade. The tail of Serikornis is covered proximally by filaments and distally by slender rectrices. Thin symmetrical remiges lacking barbules are attached along its forelimbs and elongate hindlimb feathers extend up to its toes, suggesting that hindlimb remiges evolved in ground-dwelling maniraptorans before being co-opted to an arboreal lifestyle or flight.     

New postcrania of Deccanolestes from the Late Cretaceous of India and their bearing on the evolutionary and biogeographic history of euarchontan mammals

Extant species of the supraordinal mammal clade Euarchonta belong to the orders Primates, Scandentia, or Dermoptera. The fossil record of euarchontans suggests that they underwent their initial radiation during the Paleocene (65–55 million years ago) in North America, Eurasia, and Africa. The time and place of origin is poorly resolved due to lack of definitive fossils of euarchontan stem taxa. We describe a fragmentary humerus and two fragmentary ulnae from the latest Cretaceous of India that bear significantly on this issue. The fossils are tentatively referred to Deccanolestes cf. hislopi due to their small size and the fact that Deccanolestes is the only eutherian dental taxon to have been recovered from the same locality. The new fossils are used to evaluate the existing behavioral hypothesis that Deccanolestes was arboreal, and the competing phylogenetic hypotheses that Deccanolestes is a stem eutherian versus a stem euarchontan. The humerus resembles those of euarchontans in possessing a laterally keeled ulnar trochlea, a distinct zona conoidea, and a spherical capitulum. These features also suggest an arboreal lifestyle. The ulnar morphology is consistent with that of the humerus in reflecting an arboreal/scansorial animal. Detailed quantitative comparisons indicate that, despite morphological correlates to euarchontan-like arboreality, the humerus of Deccanolestes is morphologically intermediate between those of Cretaceous “condylarthran” mammals and definitive Cenozoic euarchontans. Additionally, humeri attributed to adapisoriculids are morphologically intermediate between those of Deccanolestes and definitive euarchontans. If adapisoriculids are euarchontans, as recently proposed, our results suggest that Deccanolestes is more basal. The tentative identification of Deccanolestes as a basal stem euarchontan suggests that (1) Placentalia began to diversify and Euarchonta originated before the Cretaceous–Tertiary boundary and (2) the Indian subcontinent, Eurasia, and Africa are more likely places of origin for Euarchonta than is North America.     

A new captorhinid reptile, Gansurhinus qingtoushanensis, gen. et sp. nov., from the Permian of China

Captorhinids, a clade of Paleozoic reptiles, are represented by a rich fossil record that extends from the Late Carboniferous into the Late Permian. Representatives of this clade dispersed from the equatorial regions of Laurasia into the temperate regions of Pangea during the Middle and Late Permian. This rich fossil record shows that there was an evolutionary trend from faunivorous to omnivorous and herbivorous feeding habits within this clade. The discovery of well-preserved captorhinid materials in the Middle Permian of China allows us to determine that the new taxon, Gansurhinus qingtoushanensis, gen. et sp. nov, is a member of Moradisaurinae, a clade of captorhinids with multiple tooth rows arranged in parallel. The presence of this moradisaurine in the Middle Permian of south central Asia leads us to suggest that paleogeographic changes during the Permian, with part of what is today China becoming a large peninsula of Pangea, allowed these early reptiles as well as other terrestrial vertebrates to extend their geographic ranges to this region of the Late Paleozoic supercontinent.     

Postnatal nutrition influences male attractiveness and promotes plasticity in male mating preferences

The fossil record of Late Cretaceous–Paleogene modern birds in the Southern Hemisphere includes the Maastrichtian Neogaeornis wetzeli from Chile, Polarornis gregorii and Vegavis iaai from Antarctica, and Australornis lovei from the Paleogene of New Zealand. The recent finding of a new and nearly complete Vegavis skeleton constitutes the most informative source for anatomical comparisons among Australornis, Polarornis, and Vegavis. The present contribution includes, for the first time, Vegavis, Polarornis, and Australornis in a comprehensive phylogenetic analysis. This analysis resulted in the recognition of these taxa as a clade of basal Anseriformes that we call Vegaviidae. Vegaviids share a combination of characters related to diving adaptations, including compact and thickened cortex of hindlimb bones, femur with anteroposteriorly compressed and bowed shaft, deep and wide popliteal fossa delimited by a medial ridge, tibiotarsus showing notably proximally expanded cnemial crests, expanded fibular crest, anteroposterior compression of the tibial shaft, and a tarsometatarsus with a strong transverse compression of the shaft. Isolated bones coming from the Cretaceous and Paleogene of South America, Antarctica, and New Zealand are also referred to here to Vegaviidae and support the view that these basal anseriforms were abundant and diverse at high southern latitudes. Moreover, vegaviids represent the first avian lineage to have definitely crossed the K–Pg boundary, supporting the idea that some avian clades were not affected by the end Mesozoic mass extinction event, countering previous interpretations. Recognition of Vegaviidae indicates that modern birds were diversified in southern continents by the Cretaceous and reinforces the hypothesis indicating the important role of Gondwana for the evolutionary history of Anseriformes and Neornithes as a whole.