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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.

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.

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.

Adalatherium hui is a latest Cretaceous (Maastrichtian) gondwanatherian mammal based on only a single specimen, a virtually complete, articulated, and well-preserved skull and postcranial skeleton. The specimen is the most complete and best preserved of any mammaliaform from the Mesozoic of the southern supercontinent Gondwana. It was discovered in the Anembalemba Member of the Maevarano Formation in the Mahajanga Basin of northwestern Madagascar. The holotype specimen includes the only complete lower jaw and the only postcranial remains known for Gondwanatheria, which, other than the cranium of Vintana sertichi (also from the latest Cretaceous of Madagascar), are represented only by isolated teeth and fragmentary dentaries. Despite being represented by an immature individual, A. hui is third only to V. sertichi and Coloniatherium cilinskii as the largest Mesozoic mammaliaform (based on body fossils) from Gondwana. Here, we (1) review the paucity of mammaliamorph skull and postcranial skeletal material from the Mesozoic of Gondwana relative to the record from Laurasia; (2) review the systematic paleontology of A. hui; (3) provide an overview of the history of discovery of the holotype specimen; (4) detail the preservation of the holotype, its preparation history, and the imaging techniques used to study it; (5) provide an overview of the geological context of A. hui, which indicates that the species lived close to the end-Cretaceous extinction event in a highly seasonal, semiarid climate; and (6) estimate the body mass of A. hui in the context of other Mesozoic mammaliaforms.

Vintana sertichi is a sudamericid gondwanatherian mammal known only from the Upper Cretaceous (Maastrichtian) Maevarano Formation in the Mahajanga Basin of northwestern Madagascar. It is based on a single specimen, a well-preserved and virtually complete cranium discovered in 2010 near Lac Kinkony. The cranium is superficially bizarre and constitutes the only cranial remains of the poorly known, phylogenetically enigmatic Gondwanatheria, which are otherwise known only from isolated teeth and fragmentary dentaries. Gondwanatheria are represented by seven other monotypic genera assigned to two families (Ferugliotheriidae and Sudamericidae). Historically, this clade was assigned to Xenarthra, Paratheria (as a sister group to Xenarthra), Multituberculata, Allotheria (as a sister group to Multituberculata), Mammalia incertae sedis, and, most recently, back to Multituberculata or a close relative of Multituberculata. The craniodental evidence provided by Vintana supports inclusion in Sudamericidae, the monophyly of Gondwanatheria, and the position of Gondwanatheria as nested within or sister to Multituberculata. In addition to briefly reviewing the taxonomic composition and phylogenetic history of Gondwanatheria, this introductory chapter sets the stage for the other chapters in the volume by (1) briefly summarizing the inferred life habits of gondwanatherians; (2) reviewing the systematic paleontology of V. sertichi; and (3) providing overviews of the discovery of the holotypic specimen, its preservation, its preparation, and the imaging and measurement techniques used to study it. The chapter closes with an overview of the geological context of V. sertichi, which indicates that the species lived in a coastal floodplain environment and in a highly seasonal, semiarid climate.

Adalatherium hui is a latest Cretaceous (Maastrichtian) gondwanatherian mammal based on only a single specimen, a virtually complete, articulated, and well-preserved skull and postcranial skeleton. The specimen is the most complete and best preserved of any mammaliaform from the Mesozoic of the southern supercontinent Gondwana. It was discovered in the Anembalemba Member of the Maevarano Formation in the Mahajanga Basin of northwestern Madagascar. The holotype specimen includes the only complete lower jaw and the only postcranial remains known for Gondwanatheria, which, other than the cranium of Vintana sertichi (also from the latest Cretaceous of Madagascar), are represented only by isolated teeth and fragmentary dentaries. Despite being represented by an immature individual, A. hui is third only to V. sertichi and Coloniatherium cilinskii as the largest Mesozoic mammaliaform (based on body fossils) from Gondwana. Here, we (1) review the paucity of mammaliamorph skull and postcranial skeletal material from the Mesozoic of Gondwana relative to the record from Laurasia; (2) review the systematic paleontology of A. hui; (3) provide an overview of the history of discovery of the holotype specimen; (4) detail the preservation of the holotype, its preparation history, and the imaging techniques used to study it; (5) provide an overview of the geological context of A. hui, which indicates that the species lived close to the end-Cretaceous extinction event in a highly seasonal, semiarid climate; and (6) estimate the body mass of A. hui in the context of other Mesozoic mammaliaforms.

Vintana sertichi is a sudamericid gondwanatherian mammal known only from the Upper Cretaceous (Maastrichtian) Maevarano Formation in the Mahajanga Basin of northwestern Madagascar. It is based on a single specimen, a well-preserved and virtually complete cranium discovered in 2010 near Lac Kinkony. The cranium is superficially bizarre and constitutes the only cranial remains of the poorly known, phylogenetically enigmatic Gondwanatheria, which are otherwise known only from isolated teeth and fragmentary dentaries. Gondwanatheria are represented by seven other monotypic genera assigned to two families (Ferugliotheriidae and Sudamericidae). Historically, this clade was assigned to Xenarthra, Paratheria (as a sister group to Xenarthra), Multituberculata, Allotheria (as a sister group to Multituberculata), Mammalia incertae sedis, and, most recently, back to Multituberculata or a close relative of Multituberculata. The craniodental evidence provided by Vintana supports inclusion in Sudamericidae, the monophyly of Gondwanatheria, and the position of Gondwanatheria as nested within or sister to Multituberculata. In addition to briefly reviewing the taxonomic composition and phylogenetic history of Gondwanatheria, this introductory chapter sets the stage for the other chapters in the volume by (1) briefly summarizing the inferred life habits of gondwanatherians; (2) reviewing the systematic paleontology of V. sertichi; and (3) providing overviews of the discovery of the holotypic specimen, its preservation, its preparation, and the imaging and measurement techniques used to study it. The chapter closes with an overview of the geological context of V. sertichi, which indicates that the species lived in a coastal floodplain environment and in a highly seasonal, semiarid climate.