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The fossil record of mammaliaforms (mammals and their closest relatives) of the Mesozoic era from the southern supercontinent Gondwana is far less extensive than that from its northern counterpart, Laurasia 1 , 2 . Among Mesozoic mammaliaforms, Gondwanatheria is one of the most poorly known clades, previously represented by only a single cranium and isolated jaws and teeth 1 – 5 . As a result, the anatomy, palaeobiology and phylogenetic relationships of gondwanatherians remain unclear. Here we report the discovery of an articulated and very well-preserved skeleton of a gondwanatherian of the latest age (72.1–66 million years ago) of the Cretaceous period from Madagascar that we assign to a new genus and species, Adalatherium hui . To our knowledge, the specimen is the most complete skeleton of a Gondwanan Mesozoic mammaliaform that has been found, and includes the only postcranial material and ascending ramus of the dentary known for any gondwanatherian. A phylogenetic analysis including the new taxon recovers Gondwanatheria as the sister group to Multituberculata. The skeleton, which represents one of the largest of the Gondwanan Mesozoic mammaliaforms, is particularly notable for exhibiting many unique features in combination with features that are convergent on those of therian mammals. This uniqueness is consistent with a lineage history for A. hui of isolation on Madagascar for more than 20 million years. Adalatherium hui , a newly discovered gondwanatherian mammal from Madagascar dated to near the end of the Cretaceous period, shows features consistent with a long evolutionary trajectory of isolation in an insular environment.

Aim: To evaluate the Gunnerus Ridge land-bridge hypothesis, which postulates a Late Cretaceous causeway between eastern Antarctica and southern Madagascar allowing the passage of terrestrial vertebrates. Location: Eastern Antarctica, southern Indian Ocean, Madagascar. Methods: The review involves palaeogeographical modelling, which draws upon geological and geophysical data, bathymetrie charts, and plate tectonic reconstructions, and the evaluation of stratigraphically calibrated phylogenetic analyses to document ghost lineages of select taxa. Results: The available geological and geophysical evidence indicates that eastern Antarctica's Gunnerus Ridge and southern Madagascar were separated for the entire Late Cretaceous by a vast marine expanse. In the mid-Late Cretaceous, the gap was probably punctuated by land on two intervening physiographical highs, the northern Madagascar Plateau and Conrad Rise, the latter of which, although probably large, was still separated from Antarctica's Riiser-Larsen Peninsula by c. 1600 km. Recent, stratigraphically calibrated phylogenies including large, terrestrial end-Cretaceous vertebrate taxa of Madagascar and the Indian subcontinent reveal long ghost lineages that extended into the Early Cretaceous. Main conclusions: The view that Antarctica and Madagascar were connected by a long causeway between the Gunnerus Ridge and southern Madagascar in the Late Cretaceous, and that terrestrial vertebrates were able to colonize new frontiers using this physiographical feature, is almost certainly incorrect, as was previously demonstrated for the purported causeway between Antarctica and the Indian subcontinent across the Kerguelen Plateau. Connection across mainland Africa to account for the close relationships of several fossil and extant vertebrate taxa of Indo-Madagascar and South America is another option, although this too lacks credibility. We conclude that (1) throughout the Late Cretaceous there was no intervening, continuous causeway through Antarctica and associated land bridges between South America to the west and Indo-Madagascar to the east; and (2) mid-to large-sized, obligate terrestrial forms (e.g. abelisauroid theropod and titanosaurian sauropod dinosaurs and notosuchian crocodyliforms) gained broad distribution across Gondwanan land masses prior to fragmentation and were isolated on Indo-Madagascar before the end of the Early Cretaceous.

The extant anuran fauna of Madagascar is exceptionally rich and almost completely endemic. In recent years, many new species have been described and understanding of the history and relationships of this fauna has been greatly advanced by molecular studies, but very little is known of the fossil history of frogs on the island. Beelzebufo ampinga, the first named pre-Holocene frog from Madagascar, was described in 2008 on the basis of numerous disarticulated cranial and postcranial elements from the Upper Cretaceous (Maastrichtian) Maevarano Formation of Madagascar. These specimens documented the presence of a hyperossified taxon that differed strikingly from extant Malagasy frogs in its large size and heavy coarse cranial exostosis. Here we describe and analyse new, articulated, and more complete material of the skull, vertebral column, and hind limb, as well as additional isolated elements discovered since 2008. μCT scans allow a detailed understanding of both internal and external morphology and permit a more accurate reconstruction. The new material shows Beelzebufo to have been even more bizarre than originally interpreted, with large posterolateral skull flanges and sculptured vertebral spine tables. The apparent absence of a tympanic membrane, the strong cranial exostosis, and vertebral morphology suggest it may have burrowed during seasonally arid conditions, which have been interpreted for the Maevarano Formation from independent sedimentological and taphonomic evidence. New phylogenetic analyses, incorporating both morphological and molecular data, continue to place Beelzebufo with hyloid rather than ranoid frogs. Within Hyloidea, Beelzebufo still groups with the South American Ceratophryidae thus continuing to pose difficulties with both biogeographic interpretations and prior molecular divergence dates.

How, when, and from where Madagascar's vertebrates arrived on the island is poorly known, and a comprehensive explanation for the distribution of its organisms has yet to emerge. We begin to break that impasse by analyzing vertebrate arrival patterns implied by currently existing taxa. For each of 81 clades, we compiled arrival date, source, and ancestor type (obligate freshwater, terrestrial, facultative swimmer, or volant). We analyzed changes in arrival rates, with and without adjusting for clade extinction. Probability of successful transoceanic dispersal is negatively correlated with distance traveled and influenced by ocean currents and ancestor type. Obligate rafters show a decrease in probability of successful transoceanic dispersal from the Paleocene onward, reaching the lowest levels after the mid-Miocene. This finding is consistent with a paleoceanographic model [Ali JR, Huber M (2010) Nature 463:653–656] that predicts Early Cenozoic surface currents periodically conducive to rafting or swimming from Africa, followed by a reconfiguration to present-day flow 15–20 million years ago that significantly diminished the ability for transoceanic dispersal to Madagascar from the adjacent mainland.

Previously known only from isolated teeth and lower jaw fragments recovered from the Cretaceous and Palaeogene of the Southern Hemisphere, the Gondwanatheria constitute the most poorly known of all major mammaliaform radiations. Here we report the discovery of the first skull material of a gondwanatherian, a complete and well-preserved cranium from Upper Cretaceous strata in Madagascar that we assign to a new genus and species. Phylogenetic analysis strongly supports its placement within Gondwanatheria, which are recognized as monophyletic and closely related to multituberculates, an evolutionarily successful clade of Mesozoic mammals known almost exclusively from the Northern Hemisphere. The new taxon is the largest known mammaliaform from the Mesozoic of Gondwana. Its craniofacial anatomy reveals that it was herbivorous, large-eyed and agile, with well-developed high-frequency hearing and a keen sense of smell. The cranium exhibits a mosaic of primitive and derived features, the disparity of which is extreme and probably reflective of a long evolutionary history in geographic isolation. The gondwanatherians were mammals known only from teeth and some jaw fragments that lived in the southern continents alongside dinosaurs; here the entire cranium of a bizarre and badger-sized fossil mammal from the Cretaceous of Madagascar shows that gondwanatherians were related to the better-known multituberculates, a long-lived and successful group of now-extinct rodent-like mammals. Anatomy of a Gondwana mammal The gondwanatheres were mammals that lived the southern continents alongside the dinosaurs during the Late Cretaceous and early Paleocene. Known only from a few teeth and some jaw fragments, their appearance and evolutionary relationships remained obscure. The entire skull of a bizarre and badger-sized fossil mammal from the Cretaceous of Madagascar changes all that. Although almost certainly highly derived — as one would expect from a member of the unique endemic island fauna of Madagascar at that time — Vintana is clearly a gondwanathere. The anatomy of the herbivorous, large-eyed and agile creature shows that gondwanatheres were related to the better-known multituberculates, a long-lived and successful group of (now also extinct) rodent-like mammals.

Analyses of phylogenetic topology and estimates of divergence timing have facilitated a reconstruction of Madagascar's colonization events by vertebrate animals, but that information alone does not reveal the major factors shaping the island's biogeographic history. Here, we examine profiles of Malagasy vertebrate clades through time within the context of the island's paleogeographical evolution to determine how particular events influenced the arrival of the island's extant groups. First we compare vertebrate profiles on Madagascar before and after selected events; then we compare tetrapod profiles on Madagascar to contemporary tetrapod compositions globally. We show that changes from the Mesozoic to the Cenozoic in the proportions of Madagascar's tetrapod clades (particularly its increase in the representation of birds and mammals) are tied to changes in their relative proportions elsewhere on the globe. Differences in the representation of vertebrate classes from the Mesozoic to the Cenozoic reflect the effects of extinction (i.e., the non-random susceptibility of the different vertebrate clades to purported catastrophic global events 65 million years ago), and new evolutionary opportunities for a subset of vertebrates with the relatively high potential for transoceanic dispersal potential. In comparison, changes in vertebrate class representation during the Cenozoic are minor. Despite the fact that the island's isolation has resulted in high vertebrate endemism and a unique and taxonomically imbalanced extant vertebrate assemblage (both hailed as testimony to its long isolation), that isolation was never complete. Indeed, Madagascar's extant tetrapod fauna owes more to colonization during the Cenozoic than to earlier arrivals. Madagascar's unusual vertebrate assemblage needs to be understood with reference to the basal character of clades originating prior to the K-T extinction, as well as to the differential transoceanic dispersal advantage of other, more recently arriving clades. Thus, the composition of Madagascar's endemic vertebrate assemblage itself provides evidence of the island's paleogeographic history.

Madagascar has a diverse but mainly endemic frog fauna, the biogeographic history of which has generated intense debate, fueled by recent molecular phylogenetic analyses and the near absence of a fossil record. Here, we describe a recently discovered Late Cretaceous anuran that differs strikingly in size and morphology from extant Malagasy taxa and is unrelated either to them or to the predicted occupants of the Madagascar-Seychelles-India landmass when it separated from Africa 160 million years ago (Mya). Instead, the previously undescribed anuran is attributed to the Ceratophryinae, a clade previously considered endemic to South America. The discovery offers a rare glimpse of the anuran assemblage that occupied Madagascar before the Tertiary radiation of mantellids and microhylids that now dominate the anuran fauna. In addition, the presence of a ceratophryine provides support for a controversial paleobiogeographical model that posits physical and biotic links among Madagascar, the Indian subcontinent, and South America that persisted well into the Late Cretaceous. It also suggests that the initial radiation of hyloid anurans began earlier than proposed by some recent estimates.

Fossil evidence reveals how jaws, ears, and tongues evolved in mammals and their relatives The early embryos of mammals and other vertebrates typically have six pharyngeal arches, tissue bands under the early brain that develop into structures of the head and neck. The first arch gives rise to the mammalian malleus and incus (middle ear bones), mandible (part of the lower jaw), and tympanic bone (which supports the ear drum); the second, the stapes (middle ear bone) and part of the hyoid bone (a horseshoe-shaped structure in the neck); and the third, the remainder of the hyoid. Although the evolutionary transition of the first pharyngeal arch is well documented by fossil evidence, those of the second and third arches have received little attention in the developmental and paleontological literature. On page 276 of this issue, Zhou et al. ( 1 ) report on a newly discovered 165-million-year-old fossil from China in which the bones of the first three pharyngeal arches are preserved. The fossil defines a new taxon named Microdocodon.

The unexpected discovery of a nearly complete skull from the Early Cretaceous epoch that has been preserved in three dimensions provides profound insights into the evolution and biogeography of early mammals. The unexpected discovery of a nearly complete skull from the Early Cretaceous epoch that has been preserved in three dimensions provides profound insights into the evolution and biogeography of early mammals.

Mesozoic birds display considerable diversity in size, flight adaptations and feather organization 1 – 4 , but exhibit relatively conserved patterns of beak shape and development 5 – 7 . Although Neornithine (that is, crown group) birds also exhibit constraint on facial development 8 , 9 , they have comparatively diverse beak morphologies associated with a range of feeding and behavioural ecologies, in contrast to Mesozoic birds. Here we describe a crow-sized stem bird, Falcatakely forsterae gen. et sp. nov., from the Late Cretaceous epoch of Madagascar that possesses a long and deep rostrum, an expression of beak morphology that was previously unknown among Mesozoic birds and is superficially similar to that of a variety of crown-group birds (for example, toucans). The rostrum of Falcatakely is composed of an expansive edentulous maxilla and a small tooth-bearing premaxilla. Morphometric analyses of individual bony elements and three-dimensional rostrum shape reveal the development of a neornithine-like facial anatomy despite the retention of a maxilla–premaxilla organization that is similar to that of nonavialan theropods. The patterning and increased height of the rostrum in Falcatakely reveals a degree of developmental lability and increased morphological disparity that was previously unknown in early branching avialans. Expression of this phenotype (and presumed ecology) in a stem bird underscores that consolidation to the neornithine-like, premaxilla-dominated rostrum was not an evolutionary prerequisite for beak enlargement. A crow-sized stem bird, Falcatakely forsterae , possesses a long and deep rostrum—a beak morphology that was previously unknown among Mesozoic birds and is similar to that of some crown-group birds, such as toucans.