The evolution of giant flightless birds and novel phylogenetic relationships for extinct fowl (Aves, Galloanseres).
ABSTRACT: The extinct dromornithids, gastornithids and phorusrhacids are among the most spectacular birds to have ever lived, with some giants exceeding 500?kg. The affinities and evolution of these and other related extinct birds remain contentious, with previous phylogenetic analyses being affected by widespread convergence and limited taxon sampling. We address these problems using both parsimony and tip-dated Bayesian approaches on an expansive taxon set that includes all key extinct flightless and flighted (e.g. Vegavis and lithornithids) forms, an extensive array of extant fowl (Galloanseres), representative Neoaves and palaeognaths. The Paleogene volant Lithornithidae are recovered as stem palaeognaths in the Bayesian analyses. The Galloanseres comprise four clades inferred to have diverged in the Late Cretaceous on Gondwana. In addition to Anseriformes and Galliformes, we recognize a robust new clade (Gastornithiformes) for the giant flightless Dromornithidae (Australia) and Gastornithidae (Eurasia, North America). This clade exhibits parallels to ratite palaeognaths in that flight presumably was lost and giant size attained multiple times. A fourth clade is represented by the Cretaceous Vegavis (Antarctica), which was strongly excluded from Anseriformes; thus, a crucial molecular calibration point needs to be reconsidered. The presbyornithids Wilaru (Australia) and Presbyornis (Northern Hemisphere) are robustly found to be the sister group to Anatoidea (Anseranatidae?+?Anatidae), a relatively more basal position than hitherto recognized. South America's largest bird, Brontornis, is not a galloansere, but a member of Neoaves related to Cariamiformes; therefore, giant Galloanseres remain unknown from this continent. Trait analyses showed that while gigantism and flightlessness evolved repeatedly in groups, diet is constrained by phylogeny: all giant Galloanseres and palaeognaths are herbivores or mainly herbivorous, and giant neoavians are zoophagous or omnivorous.
Project description:The recently extinct Malagasy elephant birds (Palaeognathae, Aepyornithiformes) included the largest birds that ever lived. Elephant bird neuroanatomy is understudied but can shed light on the lifestyle of these enigmatic birds. Palaeoneurological studies can provide clues to the ecologies and behaviours of extinct birds because avian brain shape is correlated with neurological function. We digitally reconstruct endocasts of two elephant bird species, Aepyornis maximus and A. hildebrandti, and compare them with representatives of all major extant and recently extinct palaeognath lineages. Among palaeognaths, we find large olfactory bulbs in taxa generally occupying forested environments where visual cues used in foraging are likely to be limited. We detected variation in olfactory bulb size among elephant bird species, possibly indicating interspecific variation in habitat. Elephant birds exhibited extremely reduced optic lobes, a condition also observed in the nocturnal kiwi. Kiwi, the sister taxon of elephant birds, have effectively replaced their visual systems with hyperdeveloped olfactory, somatosensory and auditory systems useful for foraging. We interpret these results as evidence for nocturnality among elephant birds. Vision was likely deemphasized in the ancestor of elephant birds and kiwi. These results show a previously unreported trend towards decreased visual capacity apparently exclusive to flightless, nocturnal taxa endemic to predator-depauperate islands.
Project description:The origin and timing of the diversification of modern birds remains controversial, primarily because phylogenetic relationships are incompletely resolved and uncertainty persists in molecular estimates of lineage ages. Here, we present a species tree for the major palaeognath lineages using 27 nuclear genes and 27 archaic retroposon insertions. We show that rheas are sister to the kiwis, emu and cassowaries, and confirm ratite paraphyly because tinamous are sister to moas. Divergence dating using 10 genes with broader taxon sampling, including emu, cassowary, ostrich, five kiwis, two rheas, three tinamous, three extinct moas and 15 neognath lineages, suggests that three vicariant events and possibly two dispersals are required to explain their historical biogeography. The age of crown group birds was estimated at 131 Ma (95% highest posterior density 122-138 Ma), similar to previous molecular estimates. Problems associated with gene tree discordance and incomplete lineage sorting in birds will require much larger gene sets to increase species tree accuracy and improve error in divergence times. The relatively rapid branching within neoaves pre-dates the extinction of dinosaurs, suggesting that the genesis of the radiation within this diverse clade of birds was not in response to the Cretaceous-Paleogene extinction event.
Project description:In 2006, a partial avian femur (South Dakota School of Mines and Technology (SDSM) 78247) from the Upper Cretaceous (Maastrichtian) Sandwich Bluff Member of the López de Bertodano Formation of Sandwich Bluff on Vega Island of the northern Antarctic Peninsula was briefly reported as that of a cariamiform-a clade that includes extant and volant South American species and many extinct flightless and cursorial species. Although other authors have since rejected this taxonomic assignment, SDSM 78247 had never been the subject of a detailed description, hindering a definitive assessment of its affinities. Here we provide the first comprehensive description, illustration, and comparative study of this specimen. Comparison of characters that may be assessed in this femur with those of avian taxa scored in published character matrices refutes the inclusion of SDSM 78247 within Cariamiformes, instead supporting its assignment to a new, as-yet unnamed large-bodied species within the genus Vegavis, and therefore its referral to a clade of semiaquatic anseriforms. Important character states diagnostic of Vegavis + Polarornis include strong craniocaudal bowing of the femoral shaft, the presence of a distinct fossa just proximal to the fibular trochlea, and the broad and flat shape of the patellar sulcus. Referral to Vegavis is based on the presence of a distinctive proximocaudal fossa and distolateral scar. This genus was previously known only from Vegavis iaai, a smaller-bodied taxon from the same locality and stratigraphic unit. Our reassignment of SDSM 78247 to Vegavis sp. removes the record of cariamiform landbirds from the Antarctic Cretaceous.
Project description:Just as skeletal characteristics provide clues regarding behavior of extinct vertebrates, phylogenetically-informed evaluation of endocranial morphology facilitates comparisons among extinct taxa and extant taxa with known behavioral characteristics. Previous research has established that endocranial morphology varies across Aves; however, variation of those systems among closely related species remains largely unexplored. The Charadriiformes (shorebirds and allies) are an ecologically diverse clade with a comparatively rich fossil record, and therefore, are well suited for investigating interspecies variation, and potential links between endocranial morphology, phylogeny, ecology and other life history attributes. Endocranial endocasts were rendered from high resolution X-ray computed tomography data for 17 charadriiforms (15 extant and two flightless extinct species). Evaluation of endocranial character state changes on a phylogeny for Charadriiformes resulted in identification of characters that vary in taxa with distinct feeding and locomotor ecologies. In comparison with all other charadriiforms, stem and crown clade wing-propelled diving Pan-Alcidae displayed compressed semicircular canals, and indistinct occipital sinuses and cerebellar fissures. Flightless wing-propelled divers have relatively smaller brains for their body mass and smaller optic lobes than volant pan-alcids. Observed differences between volant and flightless wing-propelled sister taxa are striking given that flightless pan-alcids continue to rely on the flight stroke for underwater propulsion. Additionally, the brain of the Black Skimmer Rynchops niger, a taxon with a unique feeding ecology that involves continuous forward aerial motion and touch-based prey detection used both at day and night, is discovered to be unlike that of any other sampled charadriiform in having an extremely large wulst as well as a small optic lobe and distinct occipital sinus. Notably, the differences between the Black Skimmer and other charadriiforms are more pronounced than between wing-propelled divers and other charadriiforms. Finally, aspects of endosseous labyrinth morphology are remarkably similar between divers and non-divers, and may deserve further evaluation.
Project description:Although flightless alcids from the Miocene and Pliocene of the eastern Pacific Ocean have been known for over 100 years, there is no detailed evaluation of diversity and systematic placement of these taxa. This is the first combined analysis of morphological and molecular data to include all extant alcids, the recently extinct Great Auk Pinguinus impennis, the mancalline auks, and a large outgroup sampling of 29 additional non-alcid charadriiforms. Based on the systematic placement of Mancallinae outside of crown clade Alcidae, the clade name Pan-Alcidae is proposed to include all known alcids. An extensive review of the Mancallinae fossil record resulted in taxonomic revision of the clade, and identification of three new species. In addition to positing the first hypothesis of inter-relationships between Mancallinae species, phylogenetic results support placement of Mancallinae as the sister taxon to all other Alcidae, indicating that flightlessness evolved at least twice in the alcid lineage. Convergent osteological characteristics of Mancallinae, the flightless Great Auk, and Spheniscidae are summarized, and implications of Mancallinae diversity, radiation, and extinction in the context of paleoclimatic changes are discussed.
Project description:Crocodyliforms were one of the most successful groups of Mesozoic tetrapods, radiating into terrestrial, semiaquatic and marine environments, while occupying numerous trophic niches, including carnivorous, insectivorous, herbivorous, and piscivorous species. Among these taxa were the enigmatic, poorly represented flat-headed crocodyliforms from the late Cretaceous of northern Africa. Here we report a new, giant crocodyliform from the early Late Cretaceous (Cenomanian) Kem Kem Formation of Morocco. Represented by a partial braincase, the taxon has an extremely long, flat skull with large jaw and craniocervical muscles. The skull roof is ridged and ornamented with a broad, rough boss surrounded by significant vascular impressions, likely forming an integumentary structure unique among crocodyliforms. Size estimates using endocranial volume indicate the specimen was very large. The taxon possesses robust laterosphenoids with laterally oriented capitate processes and isolated epipterygoids, features allying it with derived eusuchians. Phylogenetic analysis finds the taxon to be a derived eusuchian and sister taxon to Aegyptosuchus, a poorly understood, early Late Cretaceous taxon from the Bahariya formation. This clade forms the sister clade of crown-group Crocodylia, making these taxa the earliest eusuchian crocodyliforms known from Africa. These results shift phylogenetic and biogeographical hypotheses on the origin of modern crocodylians towards the circum-Tethyean region and provide important new data on eusuchian morphology and evolution.
Project description:Insular avifaunas have repeatedly spawned evolutionary novelties in the form of unusually large, often flightless species. We report fossils from the Early Miocene St Bathans Fauna of New Zealand that attests to the former existence of a giant psittaciform, which is described as a new genus and species. The fossils are two incomplete tibiotarsi from a bird with an estimated mass of 7 kg, double that of the heaviest known parrot, the kakapo Strigops habroptila. These psittaciform fossils show that parrots join the growing group of avian taxa prone to giantism in insular species, currently restricted to palaeognaths, anatids, sylviornithids, columbids, aptornithids, ciconiids, tytonids, falconids and accipitrids.
Project description:BACKGROUND:Bothremydidae is a clade of extinct pleurodiran turtles known from the Cretaceous to Paleogene of Africa, Europe, India, Madagascar, and North and South America. The group is most diverse during the Late Cretaceous to Paleogene of Africa. Little is known, however, about the early evolution of the group. METHODS:We here figure and describe a fossil turtle from early Late Cretaceous deposits exposed at MacFarlane Mine in Cedar Canyon, southwestern Utah, USA. The sediments associated with the new turtle are utilized to infer its stratigraphic provenience and the depositional settings in which it was deposited. The fossil is compared to previously described fossil pleurodires, integrated into a modified phylogenetic analysis of pelomedusoid turtles, and the biogeography of bothremydid turtles is reassessed. In light of the novel phylogenetic hypotheses, six previously established taxon names are converted to phylogenetically defined clade names to aid communication. RESULTS:The new fossil turtle can be inferred with confidence to have originated from a brackish water facies within the late Cenomanian Culver Coal Zone of the Naturita Formation. The fossil can be distinguished from all other previously described pleurodires and is therefore designated as a new taxon, Paiutemys tibert gen. et. sp. nov. Phylogenetic analysis places the new taxon as sister to the European Polysternon provinciale, Foxemys trabanti and Foxemys mechinorum at the base of Bothremydinae. Biogeographic analysis suggests that bothremydids originated as continental turtles in Gondwana, but that bothremydines adapted to near-shore marine conditions and therefore should be seen as having a circum-Atlantic distribution.
Project description:The giant flightless bird Sylviornis neocaledoniae (Aves: Sylviornithidae) existed on La Grande Terre and Ile des Pins, New Caledonia, until the late Holocene when it went extinct shortly after human arrival on these islands. The species was generally considered to be a megapode (Megapodiidae) until the family Sylviornithidae was erected for it in 2005 to reflect multiple cranial autapomorphies. However, despite thousands of bones having been reported for this unique and enigmatic taxon, the postcranial anatomy has remained largely unknown. We rectify this deficiency and describe the postcranial skeleton of S. neocaledoniae based on ~600 fossils and use data from this and its cranial anatomy to make a comprehensive assessment of its phylogenetic affinities. Sylviornis neocaledoniae is found to be a stem galliform, distant from megapodiids, and the sister taxon to the extinct flightless Megavitiornis altirostris from Fiji, which we transfer to the family Sylviornithidae. These two species form the sister group to extant crown-group galliforms. Several other fossil galloanseres also included in the phylogenetic analysis reveal novel hypotheses of their relationships as follows: Dromornis planei (Dromornithidae) is recovered as a stem galliform rather than a stem anseriform; Presbyornis pervetus (Presbyornithidae) is the sister group to Anseranatidae, not to Anatidae; Vegavis iaai is a crown anseriform but remains unresolved relative to Presbyornis pervetus, Anseranatidae and Anatidae. Sylviornis neocaledoniae was reconstructed herein to be 0.8 m tall in a resting stance and weigh 27-34 kg. The postcranial anatomy of S. neocaledoniae shows no indication of the specialised adaptation to digging seen in megapodiids, with for example, its ungual morphology differing little from that of chicken Gallus gallus. These observations and its phylogenetic placement as stem galliforms makes it improbable that this species employed ectothermic incubation or was a mound-builder. Sylviornis neocaledoniae can therefore be excluded as the constructor of tumuli in New Caledonia.
Project description:Patterns of diversification and timing of evolution within Neoaves, which includes almost 95% of all bird species, are virtually unknown. On the other hand, molecular data consistently indicate a Cretaceous origin of many neoavian lineages and the fossil record seems to support an Early Tertiary diversification. Here, we present the first well-resolved molecular phylogeny for Neoaves, together with divergence time estimates calibrated with a large number of stratigraphically and phylogenetically well-documented fossils. Our study defines several well-supported clades within Neoaves. The calibration results suggest that Neoaves, after an initial split from Galloanseres in Mid-Cretaceous, diversified around or soon after the K/T boundary. Our results thus do not contradict palaeontological data and show that there is no solid molecular evidence for an extensive pre-Tertiary radiation of Neoaves.