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      Late Maastrichtian pterosaurs from North Africa and mass extinction of Pterosauria at the Cretaceous-Paleogene boundary

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      PLoS Biology
      Public Library of Science

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          Abstract

          Pterosaurs were the first vertebrates to evolve powered flight and the largest animals to ever take wing. The pterosaurs persisted for over 150 million years before disappearing at the end of the Cretaceous, but the patterns of and processes driving their extinction remain unclear. Only a single family, Azhdarchidae, is definitively known from the late Maastrichtian, suggesting a gradual decline in diversity in the Late Cretaceous, with the Cretaceous–Paleogene (K-Pg) extinction eliminating a few late-surviving species. However, this apparent pattern may simply reflect poor sampling of fossils. Here, we describe a diverse pterosaur assemblage from the late Maastrichtian of Morocco that includes not only Azhdarchidae but the youngest known Pteranodontidae and Nyctosauridae. With 3 families and at least 7 species present, the assemblage represents the most diverse known Late Cretaceous pterosaur assemblage and dramatically increases the diversity of Maastrichtian pterosaurs. At least 3 families—Pteranodontidae, Nyctosauridae, and Azhdarchidae—persisted into the late Maastrichtian. Late Maastrichtian pterosaurs show increased niche occupation relative to earlier, Santonian-Campanian faunas and successfully outcompeted birds at large sizes. These patterns suggest an abrupt mass extinction of pterosaurs at the K-Pg boundary.

          Author summary

          Pterosaurs were winged cousins of the dinosaurs and lived from around 200 million years ago to 66 million years ago, when the last pterosaurs disappeared during the Cretaceous-Paleogene extinction that wiped out the dinosaurs. The pterosaurs are thought to have declined in diversity before their final extinction, suggesting that gradual processes played a major role in their demise. However, pterosaur fossils are very rare, and thus, it is unclear whether pterosaurs were really low in diversity at this time or whether these patterns merely result from a paucity of fossils. We describe new pterosaur fossils from the end of the Cretaceous in Morocco, including as many as 7 species. They represent 3 different families and show a large range of variation in size and skeletal proportions, suggesting that they occupied a wide range of ecological niches.

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          Paleogene Fossil Birds

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            A pre-Archaeopteryx troodontid theropod from China with long feathers on the metatarsus.

            The early evolution of the major groups of derived non-avialan theropods is still not well understood, mainly because of their poor fossil record in the Jurassic. A well-known result of this problem is the 'temporal paradox' argument that is sometimes made against the theropod hypothesis of avian origins. Here we report on an exceptionally well-preserved small theropod specimen collected from the earliest Late Jurassic Tiaojishan Formation of western Liaoning, China. The specimen is referable to the Troodontidae, which are among the theropods most closely related to birds. This new find refutes the 'temporal paradox'1 and provides significant information on the temporal framework of theropod divergence. Furthermore, the extensive feathering of this specimen, particularly the attachment of long pennaceous feathers to the pes, sheds new light on the early evolution of feathers and demonstrates the complex distribution of skeletal and integumentary features close to the dinosaur-bird transition.
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              How many dimensions are needed to accurately assess functional diversity? A pragmatic approach for assessing the quality of functional spaces

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                Author and article information

                Contributors
                Role: ConceptualizationRole: Funding acquisitionRole: InvestigationRole: MethodologyRole: Project administrationRole: ResourcesRole: SupervisionRole: VisualizationRole: Writing – original draftRole: Writing – review & editing
                Role: InvestigationRole: Writing – original draftRole: Writing – review & editing
                Role: InvestigationRole: MethodologyRole: VisualizationRole: Writing – original draftRole: Writing – review & editing
                Role: Academic Editor
                Journal
                PLoS Biol
                PLoS Biol
                plos
                plosbiol
                PLoS Biology
                Public Library of Science (San Francisco, CA USA )
                1544-9173
                1545-7885
                13 March 2018
                March 2018
                13 March 2018
                : 16
                : 3
                : e2001663
                Affiliations
                [1 ] Department of Biology and Biochemistry and Milner Centre for Evolution, University of Bath, Bath, United Kingdom
                [2 ] School of Earth and Environmental Sciences, University of Portsmouth, Portsmouth, United Kingdom
                [3 ] Vertebrate Paleontology Laboratory, The University of Texas at Austin, Austin, Texas, United States of America
                Massey University, New Zealand
                Author notes

                The authors have declared that no competing interests exist.

                Author information
                http://orcid.org/0000-0002-9586-8907
                Article
                pbio.2001663
                10.1371/journal.pbio.2001663
                5849296
                29534059
                8202e0eb-79a7-497b-8e5d-399a2b51a1ec
                © 2018 Longrich et al

                This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                : 28 November 2016
                : 7 February 2018
                Page count
                Figures: 20, Tables: 0, Pages: 38
                Funding
                Research was supported by a Leverhulme Trust Research Leadership award to NRL. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
                Categories
                Research Article
                Biology and Life Sciences
                Organisms
                Eukaryota
                Animals
                Prehistoric Animals
                Archosauria
                Pterosauria
                Biology and Life Sciences
                Paleontology
                Prehistoric Animals
                Archosauria
                Pterosauria
                Earth Sciences
                Paleontology
                Prehistoric Animals
                Archosauria
                Pterosauria
                Biology and Life Sciences
                Paleontology
                Paleobiology
                Paleozoology
                Vertebrate Paleontology
                Prehistoric Animals
                Archosauria
                Pterosauria
                Earth Sciences
                Paleontology
                Paleobiology
                Paleozoology
                Vertebrate Paleontology
                Prehistoric Animals
                Archosauria
                Pterosauria
                Biology and Life Sciences
                Anatomy
                Musculoskeletal System
                Skeleton
                Humerus
                Medicine and Health Sciences
                Anatomy
                Musculoskeletal System
                Skeleton
                Humerus
                Earth Sciences
                Geology
                Geologic Time
                Mesozoic Era
                Cretaceous Period
                Biology and Life Sciences
                Organisms
                Eukaryota
                Animals
                Vertebrates
                Amniotes
                Birds
                Biology and Life Sciences
                Conservation Biology
                Species Extinction
                Ecology and Environmental Sciences
                Conservation Science
                Conservation Biology
                Species Extinction
                Biology and Life Sciences
                Evolutionary Biology
                Evolutionary Processes
                Species Extinction
                Biology and Life Sciences
                Anatomy
                Musculoskeletal System
                Skeleton
                Femur
                Medicine and Health Sciences
                Anatomy
                Musculoskeletal System
                Skeleton
                Femur
                Biology and Life Sciences
                Zoology
                Animal Anatomy
                Wings
                Biology and Life Sciences
                Ecology
                Biodiversity
                Ecology and Environmental Sciences
                Ecology
                Biodiversity
                Custom metadata
                All relevant data are included in the paper and its Supporting Information files. The fossil material described here is permanently accessioned at the Faculté des Sciences Aïn Chock, Université Hassan II, Casablanca, Morocco, and is available to qualified researchers.

                Life sciences
                Life sciences

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