An exquisitely preserved troodontid theropod with new information on the palatal structure from the Upper Cretaceous of Mongolia

Troodontidae is a clade of small-bodied theropod dinosaurs. A new troodontid, Gobivenator mongoliensis gen. et sp. nov., is described based on the most complete skeleton of a Late Cretaceous member of this clade presently known, from the Campanian Djadokhta Formation in the central Gobi Desert. G. mongoliensis is different from other troodontids in possessing a pointed anterior end of the fused parietal and a fossa on the surangular in front of the posterior surangular foramen. The skull was superbly preserved in the specimen and provides detailed information of the entire configuration of the palate in Troodontidae. Overall morphology of the palate in Gobivenator resembles those of dromaeosaurids and Archaeopteryx, showing an apparent trend of elongation of the pterygoid process of the palatine and reduction of the pterygopalatine suture toward the basal Avialae. The palatal configuration suggests that the skull of Gobivenator would have been akinetic but had already acquired prerequisites for later evolution of cranial kinesis in birds, such as the loss of the epipterygoid and reduction in contact areas among bones.     

A new bolosaurid parareptile, Belebey chengi sp. nov., from the Middle Permian of China and its paleogeographic significance

A new bolosaurid parareptile, Belebey chengi sp. nov., is described from Dashankou, Gansu Province, China, a Middle Permian locality which is known mostly for its therapsid fauna. The material consists of well-preserved mandibular and anterior skull remains and currently represents the largest and latest surviving member of Bolosauridae. Phylogenetic analysis of bolosaurid interrelationships, the first analysis of any clade of Early Permian parareptiles, indicates that the new taxon groups consistently with the other (Russian) members of the genus Belebey, and forms the sister clade to the genus Bolosaurus from North America. The Early Permian Eudibamus cursoris from Germany falls into the basal most position within Bolosauridae. Our analysis also shows that the split between the main bolosaurid lineages must have occurred near or before the Permo-Carboniferous boundary and that the paleo-equatorial region of Laurasia probably served as the center of origination for these parareptiles. A similar pattern can be found in other clades of Paleozoic amniotes, suggesting that this may be the general trend in early amniote evolution.     

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.     

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 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.     

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.     

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.     

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.     

The earliest record of birch mice from the Early Miocene Nei Mongol,China

The earliest species of birch mouse, Sicista primus sp. nov., was recovered from the 17-Ma-old (Early Miocene) Gashunyinadege locality, central Nei Mongol, China. It is ~9 Ma older than the previous first appearance datum of Sicista in Eurasia. This study indicates that North American Macrognathomys is a synonym of Eurasian Sicista, having 12 shared dental characters. As a result, the biogeography of dipodids indicates that Asian Sicista dispersed to North America as opposed to the hypothesis that Sicista originated from the North American clade. Sicista is one of the few extant rodent genera that originated as early as the Early Miocene.     

Phylogeography of Australia’s king brown snake (Pseudechis australis) reveals Pliocene divergence and Pleistocene dispersal of a top predator

King brown snakes or mulga snakes (Pseudechis australis) are the largest and among the most dangerous and wide-ranging venomous snakes in Australia and New Guinea. They occur in diverse habitats, are important predators, and exhibit considerable morphological variation. We infer the relationships and historical biogeography of P. australis based on phylogenetic analysis of 1,249 base pairs from the mitochondrial cytochrome b, NADH dehydrogenase subunit 4 and three adjacent tRNA genes using Bayesian, maximum-likelihood, and maximum-parsimony methods. All methods reveal deep phylogenetic structure with four strongly supported clades comprising snakes from New Guinea (I), localities all over Australia (II), the Kimberleys of Western Australia (III), and north-central Australia (IV), suggesting a much more ancient radiation than previously believed. This conclusion is robust to different molecular clock estimations indicating divergence in Pliocene or Late Miocene, after landbridge dispersal to New Guinea had occurred. While members of clades I, III and IV are medium-sized, slender snakes, those of clade II attain large sizes and a robust build, rendering them top predators in their ecosystems. Genetic differentiation within clade II is low and haplotype distribution largely incongruent with geography or colour morphs, suggesting Pleistocene dispersal and recent ecomorph evolution. Significant haplotype diversity exists in clades III and IV, implying that clade IV comprises two species. Members of clade II are broadly sympatric with members of both northern Australian clades. Thus, our data support the recognition of at least five species from within P. australis (auct.) under various criteria. We discuss biogeographical, ecological and medical implications of our findings.Electronic Supplementary Material  Supplementary material is available in the online version of this article at     

Spatial and temporal variations of two cyanobacteria in the mesotrophic Miyun reservoir, China

Spatial variations in phytoplankton community within a large mesotrophic reservoir （Miyun reservoir, North China） were investigated in relation to variations in physico-chemical properties, nutrient concentrations, temperature and light conditions over a 5 month period in 2009. The dynamics of phytoplankton community was represented by the dominance of cyanobacteria through summer and fall, following with a short term dominance of chlorophyta in late fall, and a relatively high abundance of diatom in October; on the other hand, maximum phytoplankton biomass was recorded in the north shallow region of Miyun reservoir with a higher nutrients level. Particular attention was paid to the impacts of environmental conditions on the growth of two cyanobacteria genera, the toxin-producing Microcystis and the taste ＆ odor-producing Oscillatoria. Microcystis biomass was in general greatly affected by water temperature and mixing depth/local water depth ratio in this reservoir, while the Oscillatoria biomass in the surface and middle layers was greatly affected by total dissolved phosphorus, and that in the bottom layer was related with the Secchi depth/local water depth ratio. Abundant Oscillatoria biomass was observed only in late September when Microcystis biomass decreased and allowed sufficient light go through.     

Middle-Eocene artiodactyls from Shanghuang (Jiangsu Province,Coastal China) and the diversity of basal dichobunoids in Asia

A new assemblage of basal dichobunoid artiodactyls from the middle-Eocene Shanghuang fissure fillings includes the diacodexeid Jiangsudon shanghuangensis gen. and sp. nov., a new species of the lantianine dichobunoid Elaschitotherium, Elaschitotherium crepaturus sp. nov., and an indeterminate suoid which is presently the earliest record of this clade. Diacodexeids are also represented by two forms provisionally referred to cf. Diacodexis sp. and to an indeterminate Diacodexeidae, respectively. The occurrence of diacodexeids in Shanghuang contrasts with the early and earliest middle-Eocene chronological range of the family in Europe and North America and suggests that the stratigraphic range of the family in Asia extends up to the middle Eocene. This may reflect particular habitats in coastal China that may have been relatively stable during the early and middle Eocene, thus preserving forest-dwelling artiodactyls that became extinct in the other Holarctic regions. Compared to other supposedly coeval North American, European, and Asian faunas, the Shanghuang mammalian assemblage is most similar to early Uintan faunas of North America but is also remarkable in recording forms close to taxa that are characteristic of the Wasatchian and Bridgerian North American Land Mammal Ages. The Irdinmanhan age of the Shanghuang fauna is supported by the mammalian assemblage recovered from the fissure D, but an Arshantan age cannot be completely ruled out at this point. Although the Shanghuang assemblage is biased towards preservation of small components of the mammalian fauna, the Shanghuang fauna provide an important and unique window into the Eocene diversity and early evolution of cetartiodactyls in eastern Asia.     

The armoured dissorophid Cacops from the Early Permian of Oklahoma and the exploitation of the terrestrial realm by amphibians

Cacops, one of the most distinctive Paleozoic amphibians, is part of a clade of dissorophoid temnospondyls that diversified in the equatorial region of Pangea during the Late Carboniferous and Early Permian, persisting into the Late Permian in Central Russia and China. Dissorophids were a successful group of fully terrestrial, often spectacularly armoured predators, the only amphibians apparently able to coexist with amniotes when the latter started to dominate terrestrial ecosystems. In this paper, we describe excellent new skulls from the Early Permian of Oklahoma attributed to Cacops, Cacops morrisi sp. nov. and provide for the first time detailed information about this iconic dissorophid. These specimens show anatomical and ontogenetic features that will impact on future studies on the evolution of terrestriality in tetrapods. For example, the large, posteriorly closed tympanic embayment has fine striations on an otherwise smooth surface, documenting the oldest known clear evidence for the presence of a tympanic membrane in the fossil record, a structure that is used for hearing airborne sound in extant tetrapods. The skull of C. morrisi also has several features associated with predatory behaviour, indicating that this dissorophid may have been one of the top terrestrial predators of its time.     

A new dromaeosaurid (Dinosauria: Theropoda) with Asian affinities from the latest Cretaceous of North America

Dromaeosaurids from the Maastrichtian of North America have a poor fossil record and are known largely from isolated teeth, which have typically been referred to taxa based on more complete material from earlier Campanian strata. An almost complete maxilla with well-preserved dentition and an associated dentary from the Hell Creek Formation of Montana are used to establish a new dromaeosaurid taxon in the latest Maastrichtian, immediately prior to the end-Cretaceous extinction event. Acheroraptor temertyorum gen. et sp. nov. is differentiated from other dromaeosaurids on the basis of a hypertrophied postantral wall that projects posteriorly into the antorbital fenestra, a maxillary fenestra positioned low in the antorbital fossa and directly posterior to the promaxillary fenestra, and distinctive dentition with marked apicobasal ridges. The new material allows a dromaeosaurid from the Maastrichtian of North America to be placed within a phylogenetic framework for the first time. Phylogenetic analysis suggests Acheroraptor is a velociraptorine that is more closely related to Asian dromaeosaurids, including Tsaagan and Velociraptor, than it is to Dromaeosaurus, Saurornitholestes, or any other taxon from North America. As part of the Lancian Tyrannosaurus–Triceratops fauna, A. temertyorum is the latest occurring dromaeosaurid. Its relationships and occurrence suggest a complex historical biogeographic scenario that involved multiple, bi-directional faunal interchanges between Asia and North America during the Late Cretaceous.     

Turanoceratops tardabilis—the first ceratopsid dinosaur from Asia

Turanoceratops tardabilis from the Upper Cretaceous (Turonian) Bissekty Formation of Dzharakuduk, Uzbekistan, represents the first definite ceratopsid dinosaur recorded from Asia. Reexamination of the original and study of newly collected material indicate that Turanoceratops has well-developed supraorbital horns and apparently lacks a nasal horn. Turanoceratops is more derived than the more or less coeval Zuniceratops from the Moreno Hill Formation of New Mexico in the presence of double-rooted teeth and of two or three teeth in each vertical dental file.     

New Early Cretaceous fossil from China documents a novel trophic specialization for Mesozoic birds

We report on a new Mesozoic bird, Longirostravis hani, from the Early Cretaceous Jehol Biota of northeastern China. The new taxon has a long, slender rostrum and mandible, and a small number of rostralmost teeth. Postcranial characters such as a furcular ramus wider ventrally than dorsally, a centrally concave proximal margin of the humeral head, and a minor metacarpal that projects distally more than the major metacarpal, support the placement of Longirostravis within euenantiornithine Enantiornithes, the most diverse clade of Mesozoic birds. The morphology of the skull, however, suggests that Longirostravis had a probing feeding behavior, a specialization previously unknown for Enantiornithes. Indeed, this discovery provides the first evidence in support of the existence of such a foraging behavior among basal lineages of Mesozoic birds.     

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.     

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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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.