Assembly of modern mammal community structure driven by Late Cretaceous dental evolution, rise of flowering plants, and dinosaur demise.

The long-standing view that Mesozoic mammaliaforms living in dinosaur-dominated ecosystems were ecologically constrained to small size and insectivory has been challenged by astonishing fossil discoveries over the last three decades. By studying these well-preserved early mammaliaform specimens, paleontologists now agree that mammaliaforms underwent ecomorphological diversification during the Mesozoic Era. This implies that Mesozoic mammaliaform communities had ecological structure and breadth that were comparable to today's small-bodied mammalian communities. However, this hypothesis remains untested in part because the primary focus of most studies is on individual taxa. Here, we present a study quantifying the ecological structure of Mesozoic mammaliaform communities with the aim of identifying evolutionary and ecological drivers that influenced the deep-time assembly of small-bodied mammaliaform communities. We used body size, dietary preference, and locomotor mode to establish the ecospace occupation of 98 extant, small-bodied mammalian communities from diverse biomes around the world. We calculated ecological disparity and ecological richness to measure the magnitude of ecological differences among species in a community and the number of different eco-cells occupied by species of a community, respectively. This modern dataset served as a reference for analyzing five exceptionally preserved, extinct mammaliaform communities (two Jurassic, two Cretaceous, one Eocene) from Konservat-Lagerstätten. Our results indicate that the interplay of at least three factors, namely the evolution of the tribosphenic molar, the ecological rise of angiosperms, and potential competition with other vertebrates, may have been critical in shaping the ecological structure of small-bodied mammaliaform communities through time.

Adaptive radiation of multituberculate mammals before the extinction of dinosaurs.

The Cretaceous-Paleogene mass extinction approximately 66 million years ago is conventionally thought to have been a turning point in mammalian evolution. Prior to that event and for the first two-thirds of their evolutionary history, mammals were mostly confined to roles as generalized, small-bodied, nocturnal insectivores, presumably under selection pressures from dinosaurs. Release from these pressures, by extinction of non-avian dinosaurs at the Cretaceous-Paleogene boundary, triggered ecological diversification of mammals. Although recent individual fossil discoveries have shown that some mammalian lineages diversified ecologically during the Mesozoic era, comprehensive ecological analyses of mammalian groups crossing the Cretaceous-Paleogene boundary are lacking. Such analyses are needed because diversification analyses of living taxa allow only indirect inferences of past ecosystems. Here we show that in arguably the most evolutionarily successful clade of Mesozoic mammals, the Multituberculata, an adaptive radiation began at least 20 million years before the extinction of non-avian dinosaurs and continued across the Cretaceous-Paleogene boundary. Disparity in dental complexity, which relates to the range of diets, rose sharply in step with generic richness and disparity in body size. Moreover, maximum dental complexity and body size demonstrate an adaptive shift towards increased herbivory. This dietary expansion tracked the ecological rise of angiosperms and suggests that the resources that were available to multituberculates were relatively unaffected by the Cretaceous-Paleogene mass extinction. Taken together, our results indicate that mammals were able to take advantage of new ecological opportunities in the Mesozoic and that at least some of these opportunities persisted through the Cretaceous-Paleogene mass extinction. Similar broad-scale ecomorphological inventories of other radiations may help to constrain the possible causes of mass extinctions.

A new Late Cretaceous iguanomorph from North America and the origin of New World Pleurodonta (Squamata, Iguania).

Iguanomorpha (stem + crown Iguania) is a diverse squamate clade with members that predominate many modern American lizard ecosystems. However, the temporal and palaeobiogeographic origins of its constituent crown clades (e.g. Pleurodonta (basilisks, iguanas, and their relatives)) are poorly constrained, mainly due to a meagre Mesozoic-age fossil record. Here, we report on two nearly complete skeletons from the Late Cretaceous (Campanian) of North America that represent a new and relatively large-bodied and possibly herbivorous iguanomorph that inhabited a semi-arid environment. The new taxon exhibits a mosaic of anatomical features traditionally used in diagnosing Iguania and non-iguanian squamates (i.e. Scleroglossa; e.g. parietal foramen at the frontoparietal suture, astragalocalcaneal notch in the tibia, respectively). Our cladistic analysis of Squamata revealed a phylogenetic link between Campanian-age North American and East Asian stem iguanomorphs (i.e. the new taxon + Temujiniidae). These results and our evaluation of the squamate fossil record suggest that crown pleurodontans were restricted to the low-latitude Neotropics prior to their early Palaeogene first appearances in the mid-latitudes of North America.

High-level similarity of dentitions in carnivorans and rodents.

The study of mammalian evolution depends greatly on understanding the evolution of teeth and the relationship of tooth shape to diet. Links between gross tooth shape, function and diet have been proposed since antiquity, stretching from Aristotle to Cuvier, Owen and Osborn. So far, however, the possibilities for exhaustive, quantitative comparisons between greatly different tooth shapes have been limited. Cat teeth and mouse teeth, for example, are fundamentally distinct in shape and structure as a result of independent evolutionary change over tens of millions of years. There is difficulty in establishing homology between their tooth components or in summarizing their tooth shapes, yet both carnivorans and rodents possess a comparable spectrum of dietary specializations from animals to plants. Here we introduce homology-free techniques to measure the phenotypic complexity of the three-dimensional shape of tooth crowns. In our geographic information systems (GIS) analysis of 441 teeth from 81 species of carnivorans and rodents, we show that the surface complexity of tooth crowns directly reflects the foods they consume. Moreover, the absolute values of dental complexity for individual dietary classes correspond between carnivorans and rodents, illustrating a high-level similarity between overall tooth shapes despite a lack of low-level similarity of specific tooth components. These results suggest that scale-independent forces have determined the high-level dental shape in lineages that are widely divergent in size, ecology and life history. This link between diet and phenotype will be useful for inferring the ecology of extinct species and illustrates the potential of fast-throughput, high-level analysis of the phenotype.

A new mammal skull from the Lower Cretaceous of China with implications for the evolution of obtuse-angled molars and 'amphilestid' eutriconodonts.

We report the discovery of Juchilestes liaoningensis, a new genus and species of eutriconodont mammal from the Lujiatun Site of the Lower Cretaceous Yixian Formation (123.2 +/- 1.0 Ma; Lower Aptian). The holotype preserves a partial skull and full dentition. Among eutriconodonts, its lower dentition is similar to taxa formerly assigned to the paraphyletic group of 'amphilestids'. Some have considered 'amphilestid' molars to represent the structural intermediate between the lower molars of the 'triconodont' pattern of cusps in alignment and the fully triangulate and more derived therian molars. However, 'amphilestid' taxa were previously represented only by the lower dentition. Our study reveals, for the first time, the upper dentition and skull structure of an 'amphilestid', and shows that at least some eutriconodonts have an obtuse-angled cusp pattern on molars in middle positions of the long molar series. Its petrosal is similar to those of other eutriconodonts and spalacotheroid 'symmetrodonts'. Our phylogenetic analyses suggest that (i) Juchilestes is most closely related to the Early Cretaceous Hakusanodon from Japan, in the same Eastern Asiatic geographic region; (ii) 'amphilestids' are not monophyletic; and (iii) eutriconodonts might not be a monophyletic group, although this hypothesis must be further tested.

Untangling the Multiple Ecological Radiations of Early Mammals.

The ecological diversification of early mammals is one of the most globally transformative events in Earth's history and the Cretaceous Terrestrial Revolution (KTR) and end-Cretaceous mass extinction are commonly hailed as catalysts. However, a confounding issue when examining this diversification is that it comprised nested radiations of mammalian subclades within the broader scope of mammalian evolution. In the past 200 million years, various independent groups experienced large-scale radiations, each involving ecological diversification from ancestral lineages of small insectivores; examples include Jurassic mammaliaforms, Late Cretaceous metatherians, and Cenozoic placentals. Here, we review these ecological radiations, highlighting the nuanced complexity of early mammal evolution, the value of ecomorphological fossil data, and the importance of phylogenetic context in macroevolutionary studies.

A large carnivorous mammal from the Late Cretaceous and the North American origin of marsupials.

Marsupial mammal relatives (stem metatherians) from the Mesozoic Era (252-66 million years ago) are mostly known from isolated teeth and fragmentary jaws. Here we report on the first near-complete skull remains of a North American Late Cretaceous metatherian, the stagodontid Didelphodon vorax. Our phylogenetic analysis indicates that marsupials or their closest relatives evolved in North America, as part of a Late Cretaceous diversification of metatherians, and later dispersed to South America. In addition to being the largest known Mesozoic therian mammal (node-based clade of eutherians and metatherians), Didelphodon vorax has a high estimated bite force and other craniomandibular and dental features that suggest it is the earliest known therian to invade a durophagous predator-scavenger niche. Our results broaden the scope of the ecomorphological diversification of Mesozoic mammals to include therian lineages that, in this case, were linked to the origin and evolution of marsupials.

The origin and early evolution of metatherian mammals: the Cretaceous record.

Metatherians, which comprise marsupials and their closest fossil relatives, were one of the most dominant clades of mammals during the Cretaceous and are the most diverse clade of living mammals after Placentalia. Our understanding of this group has increased greatly over the past 20 years, with the discovery of new specimens and the application of new analytical tools. Here we provide a review of the phylogenetic relationships of metatherians with respect to other mammals, discuss the taxonomic definition and diagnosis of Metatheria, outline the Cretaceous history of major metatherian clades, describe the paleobiology, biogeography, and macroevolution of Cretaceous metatherians, and provide a physical and climatic background of Cretaceous metatherian faunas. Metatherians are a clade of boreosphendian mammals that must have originated by the Late Jurassic, but the first unequivocal metatherian fossil is from the Early Cretaceous of Asia. Metatherians have the distinctive tightly interlocking occlusal molar pattern of tribosphenic mammals, but differ from Eutheria in their dental formula and tooth replacement pattern, which may be related to the metatherian reproductive process which includes an extended period of lactation followed by birth of extremely altricial young. Metatherians were widespread over Laurasia during the Cretaceous, with members present in Asia, Europe, and North America by the early Late Cretaceous. In particular, they were taxonomically and morphologically diverse and relatively abundant in the Late Cretaceous of western North America, where they have been used to examine patterns of biogeography, macroevolution, diversification, and extinction through the Late Cretaceous and across the Cretaceous-Paleogene (K-Pg) boundary. Metatherian diversification patterns suggest that they were not strongly affected by a Cretaceous Terrestrial Revolution, but they clearly underwent a severe extinction across the K-Pg boundary.