Tempo and Pattern of Avian Brain Size Evolution.

Relative brain sizes in birds can rival those of primates, but large-scale patterns and drivers of avian brain evolution remain elusive. Here, we explore the evolution of the fundamental brain-body scaling relationship across the origin and evolution of birds. Using a comprehensive dataset sampling> 2,000 modern birds, fossil birds, and theropod dinosaurs, we infer patterns of brain-body co-variation in deep time. Our study confirms that no significant increase in relative brain size accompanied the trend toward miniaturization or evolution of flight during the theropod-bird transition. Critically, however, theropods and basal birds show weaker integration between brain size and body size, allowing for rapid changes in the brain-body relationship that set the stage for dramatic shifts in early crown birds. We infer that major shifts occurred rapidly in the aftermath of the Cretaceous-Paleogene mass extinction within Neoaves, in which multiple clades achieved higher relative brain sizes because of a reduction in body size. Parrots and corvids achieved the largest brains observed in birds via markedly different patterns. Parrots primarily reduced their body size, whereas corvids increased body and brain size simultaneously (with rates of brain size evolution outpacing rates of body size evolution). Collectively, these patterns suggest that an early adaptive radiation in brain size laid the foundation for subsequent selection and stabilization.

Bony pseudoteeth of extinct pelagic birds (Aves, Odontopterygiformes) formed through a response of bone cells to tooth-specific epithelial signals under unique conditions.

Modern birds (crown group birds, called Neornithes) are toothless; however, the extinct neornithine Odontopterygiformes possessed bone excrescences (pseudoteeth) which resembled teeth, distributed sequentially by size along jaws. The origin of pseudoteeth is enigmatic, but based on recent evidence, including microanatomical and histological analyses, we propose that conserved odontogenetic pathways most probably regulated the development of pseudodentition. The delayed pseudoteeth growth and epithelium keratinization allowed for the existence of a temporal window during which competent osteoblasts could respond to oral epithelial signaling, in place of the no longer present odontoblasts; thus, bony pseudoteeth developed instead of true teeth. Dynamic morphogenetic fields can explain the particular, sequential size distribution of pseudoteeth along the jaws of these birds. Hence, this appears as a new kind of deep homology, by which ancient odontogenetic developmental processes would have controlled the evolution of pseudodentition, structurally different from a true dentition, but morphologically and functionally similar.

A roller-like bird (Coracii) from the Early Eocene of Denmark.

The fossil record of crown group birds (Neornithes) prior to the Cretaceous-Paleogene boundary is scarce and fragmentary. Early Cenozoic bird fossils are more abundant, but are typically disarticulated and/or flattened. Here we report the oldest roller (Coracii), Septencoracias morsensis gen. et sp. nov. (Primobucconidae), based on a new specimen from the Early Eocene (about 54 million years ago) Fur Formation of Denmark. The new fossil is a nearly complete, three-dimensionally preserved and articulated skeleton. It lies at the lower end of the size range for extant rollers. Salient diagnostic features of Septencoracias relative to other Coracii include the proportionally larger skull and the small, ovoid and dorsally positioned narial openings. Our discovery adds to the evidence that the Coracii had a widespread northern hemisphere distribution in the Eocene. Septencoracias is the oldest substantial record of the Picocoraciae and provides a reliable calibration point for molecular phylogenetic studies.

A reappraisal of Cerebavis cenomanica (Aves, Ornithurae), from Melovatka, Russia.

The evolution of the avian brain is of crucial importance to studies of the transition from non-avian dinosaurs to modern birds, but very few avian fossils provide information on brain morphological development during the Mesozoic. An isolated specimen from the Cenomanian of Melovatka in Russia was described by Kurochkin and others as a fossilized brain, designated the holotype of Cerebavis cenomanica Kurochkin and Saveliev and tentatively referred to Enantiornithes. We have previously highlighted that this specimen is an incomplete skull, rendering the diagnostic characters invalid and Cerebavis cenomanica a nomen dubium. We provide here a revised diagnosis of Cerebavis cenomanica based on osteological characters, and a reconstruction of the endocranial morphology (= brain shape) based on µCT investigation of the braincase. Absence of temporal fenestrae indicates an ornithurine affinity for Cerebavis. The brain of this taxon was clearly closer to that of modern birds than to Archaeopteryx and does not represent a divergent evolutionary pathway as originally concluded by Kurochkin and others. No telencephalic wulst is present, suggesting that this advanced avian neurological feature was not recognizably developed 93 million years ago.

A redescription of Lithornis vulturinus (Aves, Palaeognathae) from the Early Eocene Fur Formation of Denmark.

The extinct Lithornithidae include several genera and species of flying palaeognathous birds of controversial affinities known from the Early Paleogene of North America and Europe. An almost complete, articulated skeleton from the Early Eocene marine deposits of the Fur Formation (Denmark) was recently assigned to Lithornis vulturinus Owen, 1840. This study provides a detailed redescription and comparison of this three-dimensionally preserved specimen (MGUH 26770), which is one of the best preserved representatives of the Lithornithidae yet known. We suggest that some new features might be diagnostic of Lithornis vulturinus, including a pterygoid fossa shallower than in other species of Lithornis and the presence of a small caudal process on the os palatinum. We propose that Lithornis nasi (Harrison, 1984) is a junior synonym of Lithornis vulturinus and we interpret minor differences in size and shape among the specimens as intraspecific variation. To date, Lithornis vulturinus is known with certainty from the latest Paleocene-earliest Eocene to Early Eocene of the North Sea Basin (Ølst, Fur and London Clay Formations). Among the four species of the genus Lithornis, the possibility that Lithornis plebius Houde, 1988 (Early Eocene of Wyoming) is conspecific with either Lithornis vulturinus or Lithornis promiscuus Houde, 1988 (Early Eocene of Wyoming) is discussed. The presence of closely related species of Lithornis on either side of the North Atlantic in the Early Eocene reflects the existence of a high-latitude land connection between Europe and North America at that time.

Avian cerebellar floccular fossa size is not a proxy for flying ability in birds.

Extinct animal behavior has often been inferred from qualitative assessments of relative brain region size in fossil endocranial casts. For instance, flight capability in pterosaurs and early birds has been inferred from the relative size of the cerebellar flocculus, which in life protrudes from the lateral surface of the cerebellum. A primary role of the flocculus is to integrate sensory information about head rotation and translation to stabilize visual gaze via the vestibulo-occular reflex (VOR). Because gaze stabilization is a critical aspect of flight, some authors have suggested that the flocculus is enlarged in flying species. Whether this can be further extended to a floccular expansion in highly maneuverable flying species or floccular reduction in flightless species is unknown. Here, we used micro computed-tomography to reconstruct "virtual" endocranial casts of 60 extant bird species, to extract the same level of anatomical information offered by fossils. Volumes of the floccular fossa and entire brain cavity were measured and these values correlated with four indices of flying behavior. Although a weak positive relationship was found between floccular fossa size and brachial index, no significant relationship was found between floccular fossa size and any other flight mode classification. These findings could be the result of the bony endocranium inaccurately reflecting the size of the neural flocculus, but might also reflect the importance of the flocculus for all modes of locomotion in birds. We therefore conclude that the relative size of the flocculus of endocranial casts is an unreliable predictor of locomotor behavior in extinct birds, and probably also pterosaurs and non-avian dinosaurs.

Bone growth marks reveal protracted growth in New Zealand kiwi (Aves, Apterygidae).

The presence of bone growth marks reflecting annual rhythms in the cortical bone of non-avian tetrapods is now established as a general phenomenon. In contrast, ornithurines (the theropod group including modern birds and their closest relatives) usually grow rapidly in less than a year, such that no annual rhythms are expressed in bone cortices, except scarce growth marks restricted to the outer cortical layer. So far, cyclical growth in modern birds has been restricted to the Eocene Diatryma, the extant parrot Amazona amazonica and the extinct New Zealand (NZ) moa (Dinornithidae). Here we show the presence of lines of arrested growth in the long bones of the living NZ kiwi (Apteryx spp., Apterygidae). Kiwis take 5-6 years to reach full adult body size, which indicates a delayed maturity and a slow reproductive cycle. Protracted growth probably evolved convergently in moa and kiwi sometime since the Middle Miocene, owing to the severe climatic cooling in the southwest Pacific and the absence of mammalian predators.

Osteological evidence for sister group relationship between pseudo-toothed birds (Aves: Odontopterygiformes) and waterfowls (Anseriformes).

The phylogenetic affinities of the extinct pseudo-toothed birds have remained controversial. Some authors noted that they resemble both pelicans and allies (Pelecaniformes) and tube-nosed birds (Procellariiformes), but assigned them to a distinct taxon, the Odontopterygiformes. In most recent studies, the pseudo-toothed birds are referred to the family Pelagornithidae inside the Pelecaniformes. Here, I perform a cladistic analysis with five taxa of the pseudo-toothed birds including two undescribed new species from the Early Tertiary of Morocco. The present hypothesis strongly supports a sister group relationship of pseudo-toothed birds (Odontopterygiformes) and waterfowls (Anseriformes). The Odontoanserae (Odontopterygiformes plus Anseriformes) are the sister group of Neoaves. The placement of the landfowls (Galliformes) as the sister taxon of all other neognathous birds does not support the consensus view that the Galloanserae (Galliformes plus Anseriformes) are monophyletic.