Audiogram, body mass, and basilar papilla length: correlations in birds and predictions for extinct archosaurs

The inner ear in the group of archosaurs (birds, crocodilians, and extinct dinosaurs) shows a high degree of structural similarity, enabling predictions of their function in extinct species based on relationships among similar variables in living birds. Behavioral audiograms and morphological data on the length of the auditory sensory epithelium (the basilar papilla) are available for many avian species. By bringing different data sets together, we show that body mass and the size of the basilar papilla are significantly correlated, and the most sensitive frequency in a given species is inversely related to the body mass and the length of the basilar papilla. We also demonstrate that the frequency of best hearing is correlated with the high-frequency limit of hearing. Small species with a short basilar papilla hear higher frequencies compared with larger species with a longer basilar papilla. Based on the regression analysis of two significant correlations in living archosaurs (best audiogram frequency vs body mass and best audiogram frequency vs papillar length), we suggest that hearing in large dinosaurs was restricted to low frequencies with a high-frequency limit below 3 kHz. Supplementary material is available in the online version of this article at and accessible for authorised users.     

Earliest and first Northern Hemispheric hoatzin fossils substantiate Old World origin of a “Neotropic endemic”

The recent identification of hoatzins (Opisthocomiformes) in the Miocene of Africa showed part of the evolution of these birds, which are now only found in South America, to have taken place outside the Neotropic region. Here, we describe a new fossil species from the late Eocene of France, which constitutes the earliest fossil record of hoatzins and the first one from the Northern Hemisphere. Protoazin parisiensis gen. et sp. nov. is more closely related to South American Opisthocomiformes than the African taxon Namibiavis and substantiates an Old World origin of hoatzins, as well as a relictual distribution of the single extant species. Although recognition of hoatzins in Europe may challenge their presumed transatlantic dispersal, there are still no North American fossils in support of an alternative, Northern Hemispheric, dispersal route. In addition to Opisthocomiformes, other avian taxa are known from the Cenozoic of Europe, the extant representatives of which are only found in South America. Recognition of hoatzins in the early Cenozoic of Europe is of particular significance because Opisthocomiformes have a fossil record in sub-Saharan Africa, which supports the hypothesis that extinction of at least some of these “South American” groups outside the Neotropic region was not primarily due to climatic factors.     

Behavioral and faunal implications of Early Cretaceous deinonychosaur trackways from China

Deinonychosaurian theropods, the dinosaurian sister group of birds, are characterized by a large raptorial claw borne on a highly modified second digit that was thought to be held in a retracted position during locomotion. In this study, we present new trackway evidence for two coeval deinonychosaurian taxa from the Early Cretaceous of Shandong, China that indicate a hitherto unrecognized body size diversity for this period and continent. These fossil tracks confirm diversity and locomotory patterns implied by phylogeny and biogeography, but not yet manifested in the body fossil record. Multiple parallel and closely spaced trackways generated by the larger track maker provide the best evidence yet discovered for gregarious behavior in deinonychosaurian theropods.     

Associated skeletons of a new middle Triassic "Rauisuchia" from Brazil

For more than 30 million years, in early Mesozoic Pangea, ??rauisuchian?? archosaurs were the apex predators in most terrestrial ecosystems, but their biology and evolutionary history remain poorly understood. We describe a new ??rauisuchian?? based on ten individuals found in a single locality from the Middle Triassic (Ladinian) Santa Maria Formation of southern Brazil. Nine articulated and associated skeletons were discovered, three of which have nearly complete skulls. Along with sedimentological and taphonomic data, this suggests that those highly successful predators exhibited some kind of intraspecific interaction. Other monotaxic assemblages of Triassic archosaurs are Late Triassic (Norian-Rhaetian) in age, approximately 10 million years younger than the material described here. Indeed, the studied assemblage may represent the earliest evidence of gregariousness among archosaurs, adding to our knowledge on the origin of a behavior pattern typical of extant taxa.     

Survival of Theriosuchus (Mesoeucrocodylia: Atoposauridae) in a Late Cretaceous archipelago: a new species from the Maastrichtian of Romania

Small terrestrial non-eusuchian mesoeucrocodylians are common components of Cretaceous assemblages of Gondwanan provinces with notosuchians and araripesuchids as flagship taxa in South America, Africa and Madagascar, well into the Late Cretaceous. On the other hand, these are exceedingly rare in Laurasian landmasses during the Late Cretaceous. Small terrestrial mesoeucrocodylians from Europe were often referred to the genus Theriosuchus, a taxon with stratigraphic range extending from the Late Jurassic to the late Early Cretaceous. Theriosuchus is abundantly reported from various European localities, although Asiatic and possibly North American members are also known. It has often been closely associated with the first modern crocodilians, members of the Eusuchia, because of the presence of procoelous vertebrae, a widespread key character diagnosing the Eusuchia. Nevertheless, the relationships of Theriosuchus have not been explored in detail although one species, Theriosuchus pusillus, has been extensively described and referred in numerous works. Here, we describe a new basal mesoeucrocodylian, Theriosuchus sympiestodon sp. nov. from the Maastrichtian of the Haţeg Basin, Romania, suggesting a large temporal gap (about 58 myr) in the fossil record of the genus. Inclusion of the new taxon, along with Theriosuchus guimarotae, in a phylogenetic analysis confirms its referral to the genus Theriosuchus, within a monophyletic atoposaurid clade. Although phylogenetic resolution within this clade is still poor, the new taxon appears, on morphological grounds, to be most closely related to T. pusillus. The relationships of Atoposauridae within Mesoeucrocodylia and especially to Neosuchia are discussed in light of the results of the present contribution as well as from recent work. Our results raise the possibility that Atoposauridae might not be regarded as a derived neosuchian clade anymore, although further investigation of the neosuchian interrelationships is needed. Reports of isolated teeth referable to a closely related taxon from the Upper Cretaceous of Romania and France, together with the presence of Doratodon and Ischyrochampsa, indicate a previously unsuspected diverse assemblage of non-eusuchian mesoeucrocodylians in the Late Cretaceous European archipelago.     

Europe's last Mesozoic bird

Birds known from more than isolated skeletal elements are rare in the fossil record, especially from the European Mesozoic. This paucity has hindered interpretations of avian evolution immediately prior to, and in the aftermath of, the Cretaceous-Tertiary (K-T) extinction event. We report on a specimen of a large ornithurine bird (closely related to Ichthyornis) from the uppermost Cretaceous (Maastricht Formation) of Belgium. This is the first record of a bird from these historic strata and the only phylogenetically informative ornithurine to be recovered from the Mesozoic of Europe. Because this new specimen was collected from 40 m below the K-T boundary (approximate age of 65.8 Ma), it is also the youngest non-neornithine (=non-modern) bird known from anywhere in the world.     

No evidence for an afrotherian-like delayed dental eruption in South American notoungulates

The fossil South American ungulates are of great interest relative to the new phylogenetic framework elaborated for living placental mammals. In particular, studies on these endemic taxa can allow for testing congruence between southern placental phylogeny and plate tectonics, beyond what has already been suggested in the Atlantogenata hypothesis based on extant afrotherians and xenarthrans. The presence of delayed dental eruption relative to skull growth is one feature characterizing the extant afrotherians and possibly the xenarthrans. Late dental eruption has been mentioned previously in South American notoungulates, thus suggesting possible resemblance with afrotherians and perhaps xenarthrans. We provide here a detailed study of the dental eruption pattern relative to the skull growth in the notoungulates. In contrast to previous statements, our results demonstrate that there is currently no evidence for an afrotherian-like delayed dental eruption in this group. For now, the inferred absence of a delayed dental eruption in notoungulates does not support atlantogenatan/afrotherian affinities for the Notoungulata, but other atlantogenatan/afrotherian characteristics remain to be explored in more detail in this group and other South American ungulates.     

Bizarre tubercles on the vertebrae of Eocene fossil birds indicate an avian disease without modern counterpart

Remains of fossil birds with numerous bony tubercles on the cervical vertebrae are reported from the Middle Eocene of Messel in Germany and the Late Eocene of the Quercy fissure fillings in France. These structures, which are unknown from extant birds and other vertebrates, were previously described for an avian skeleton from Messel but considered a singular feature of this specimen. The new fossils are from a different species of uncertain phylogenetic affinities and show that tuberculated vertebrae have a wider taxonomic, temporal, and geographic distribution. In contrast to previous assumptions, they are no ontogenetic feature and arise from the vertebral surface. It is concluded that they are most likely of pathologic origin and the first record of a Paleogene avian disease. Their regular and symmetrical arrangement over most of the external vertebral surface indicates a systemic disorder caused by factors that do not affect extant birds, such as especially high-dosed phytohormones or extinct pathogens.     

The evolutionary position of turtles revised

Consensus on the evolutionary position of turtles within the amniote phylogeny has eluded evolutionary biologists for more than a century. This phylogenetic problem has remained unsolved partly because turtles have such a unique morphology that only few characters can be used to link them with any other group of amniotes. Among the many alternative hypotheses that have been postulated to explain the origin and phylogenetic relationships of turtles, a general agreement among paleontologists emerged in favoring the placement of turtles as the only living survivors of the anapsid reptiles (those that lack temporal fenestrae in the skull). However, recent morphological and molecular studies have radically changed our view of amniote phylogenetic relationships, and evidence is accumulating that supports the diapsid affinities of turtles. Molecular studies favor archosaurs (crocodiles and birds) as the living sister group of turtles, whereas morphological studies support lepidosaurs (tuatara, lizards, and snakes) as the closest living relatives of turtles. Accepting these hypotheses implies that turtles cannot be viewed any longer as primitive reptiles, and that they might have lost the temporal holes in the skull secondarily rather than never having had them.     

A Phororhacoid bird from the Eocene of Africa

The bird fossil record is globally scarce in Africa. The early Tertiary evolution of terrestrial birds is virtually unknown in that continent. Here, we report on a femur of a large terrestrial new genus discovered from the early or early middle Eocene (between ～52 and 46?Ma) of south-western Algeria. This femur shows all the morphological features of the Phororhacoidea, the so-called Terror Birds. Most of the phororhacoids were indeed large, or even gigantic, flightless predators or scavengers with no close modern analogs. It is likely that this extinct group originated in South America, where they are known from the late Paleocene to the late Pleistocene (～59 to 0.01?Ma). The presence of a phororhacoid bird in Africa cannot be explained by a vicariant mechanism because these birds first appeared in South America well after the onset of the mid-Cretaceous Gondwana break up (～100?million years old). Here, we propose two hypotheses to account for this occurrence, either an early dispersal of small members of this group, which were still able of a limited flight, or a transoceanic migration of flightless birds from South America to Africa during the Paleocene or earliest Eocene. Paleogeographic reconstructions of the South Atlantic Ocean suggest the existence of several islands of considerable size between South America and Africa during the early Tertiary, which could have helped a transatlantic dispersal of phororhacoids.     

Skeletal remains of a diminutive primate from the Paleocene of Germany

Most living mammal orders, including our own, started their career during the first 10 million years of the Cenozoic, the Age of Mammals. The fossil record documents that early Paleogene adaptive radiations of various clades included tiny species of the size of living shrews. Remains of particularly diminutive limb bones are described from the late Paleocene site of Walbeck, Sachsen-Anhalt. Discovered in 1939, it has remained the only known Paleocene mammal-bearing locality from Germany. The remains are referred to the family Adapisoriculidae, which is considered on the basis of the present postcranial evidence to represent plesiadapiform primates rather than alleged lipotyphlan insectivores as previously proposed. The Walbeck fossils compete with the Early Eocene species Toliapina vinealis from Europe and Picromomys petersonorum from North America for the status of the smallest known primate, fossil and living. Their estimated body weights are as small as 10 g. The limb bones show features related to enhanced flexion at the elbow and hip joint, suggesting arboreal habits and environments such as terminal branches. The diminutive size and tooth morphology suggest feeding on small insects and other invertebrates. Postcranials are important to assess early radiations, such tiny specimens as the present ones are extremely scarce in the fossil record, however.     

The origin and early evolution of birds: discoveries,disputes, and perspectives from fossil evidence

The study of the origin and early evolution of birds has never produced as much excitement and public attention as in the past decade. Well preserved and abundant new fossils of birds and dinosaurs have provided unprecedented new evidence on the dinosaurian origin of birds, the arboreal origin of avian flight, and the origin of feathers prior to flapping flight. The Mesozoic avian assemblage mainly comprises two major lineages: the prevalent extinct group Enantiornithes, and the Ornithurae, which gave rise to all modern birds, as well as several more basal taxa. Cretaceous birds radiated into various paleoecological niches that included fish- and seed-eating. Significant size and morphological differences and variation in flight capabilities, ranging from gliding to powerful flight among early birds, highlight the diversification of birds in the Early Cretaceous. There is little evidence, however, to support a Mesozoic origin of modern avian groups. Controversy and debate, nevertheless, surround many of these findings, and more details are needed to give a better appreciation of the significance of these new discoveries.     

Post-dispersal weed seed predation by avian and non-avian predators

Seed predation by avian and non-avian predators was quantified in the boundaries and cropped areas of cereal fields by presenting known quantities of seed with and without exclusion cages. Predator encounter-rates with the dishes exceeded 99%. Birds removed on average 6.7% seed from the dishes during the seven-day trials compared to 51% by non-avian predators. A comparison was made of the causal factors responsible for predation of Avena fatua, Chenopodium album and Cirsium arvense seeds. A. fatua seeds were preyed most heavily by both avian and non-avian predators. Seed removal by birds was greater in the cropped area than in the field boundary, non-avian predators being generally more active in the field boundary. Seed predation by birds was greater in spring than in any other season, whilst losses to other animals were greater during autumn and winter. Although, birds were not the main seed predators in cereal fields, they may contribute to weed seed depletion, of relevance to reduced-input farming systems where herbicides use is restricted.     

Minute theropod eggs and embryo from the Lower Cretaceous of Thailand and the dinosaur-bird transition

We report on very small fossil eggs from the Lower Cretaceous of Thailand, one of them containing a theropod embryo, which display a remarkable mosaic of characters. While the surficial ornamentation is typical of non-avian saurischian dinosaurs, the three-layered prismatic structure of the eggshell is currently known only in extant and fossil eggs associated with birds. These eggs, about the size of a goldfinch's, mirror at the reproductive level the retention of small body size that was paramount in the transition from non-avian theropods to birds. The egg-layer may have been a small feathered theropod similar to those recently found in China.     

Direct and indirect fossil records of megachilid bees from the Paleogene of Central Europe (Hymenoptera: Megachilidae)

Aside from pollen and nectar, bees of the subfamily Megachilinae are closely associated with plants as a source of materials for nest construction. Megachilines use resins, masticated leaves, trichomes and other plant materials sometimes along with mud to construct nests in cavities or in soil. Among these, the leafcutter bees (Megachile s.l.) are the most famous for their behaviour to line their brood cells with discs cut from various plants. We report on fossil records of one body fossil of a new non-leafcutting megachiline and of 12 leafcuttings from three European sites—Eckfeld and Messel, both in Germany (Eocene), and Menat, France (Paleocene). The excisions include the currently earliest record of probable Megachile activity and suggest the presence of such bees in the Paleocene European fauna. Comparison with extant leafcuttings permits the interpretation of a minimal number of species that produced these excisions. The wide range of size for the leafcuttings indirectly might suggest at least two species of Megachile for the fauna of Messel in addition to the other megachiline bee described here. The presence of several cuttings on most leaves from Eckfeld implies that the preferential foraging behaviour of extant Megachile arose early in megachiline evolution. These results demonstrate that combined investigation of body and trace fossils complement each other in understanding past biodiversity, the latter permitting the detection of taxa not otherwise directly sampled and inferences on behavioural evolution.     

Postnatal nutrition influences male attractiveness and promotes plasticity in male mating preferences

The fossil record of Late Cretaceous–Paleogene modern birds in the Southern Hemisphere includes the Maastrichtian Neogaeornis wetzeli from Chile, Polarornis gregorii and Vegavis iaai from Antarctica, and Australornis lovei from the Paleogene of New Zealand. The recent finding of a new and nearly complete Vegavis skeleton constitutes the most informative source for anatomical comparisons among Australornis, Polarornis, and Vegavis. The present contribution includes, for the first time, Vegavis, Polarornis, and Australornis in a comprehensive phylogenetic analysis. This analysis resulted in the recognition of these taxa as a clade of basal Anseriformes that we call Vegaviidae. Vegaviids share a combination of characters related to diving adaptations, including compact and thickened cortex of hindlimb bones, femur with anteroposteriorly compressed and bowed shaft, deep and wide popliteal fossa delimited by a medial ridge, tibiotarsus showing notably proximally expanded cnemial crests, expanded fibular crest, anteroposterior compression of the tibial shaft, and a tarsometatarsus with a strong transverse compression of the shaft. Isolated bones coming from the Cretaceous and Paleogene of South America, Antarctica, and New Zealand are also referred to here to Vegaviidae and support the view that these basal anseriforms were abundant and diverse at high southern latitudes. Moreover, vegaviids represent the first avian lineage to have definitely crossed the K–Pg boundary, supporting the idea that some avian clades were not affected by the end Mesozoic mass extinction event, countering previous interpretations. Recognition of Vegaviidae indicates that modern birds were diversified in southern continents by the Cretaceous and reinforces the hypothesis indicating the important role of Gondwana for the evolutionary history of Anseriformes and Neornithes as a whole.

     

Interphyletic relationships in the use of nesting cavities: mutualism,competition and amensalism among hymenopterans and vertebrates

Although competition is usually assumed to be the most common interaction between closely related organisms that share limiting resources, the relationships linking distant taxa that use the same nesting sites are poorly understood. In the present study, we examine the interactions among social hymenopterans (honeybees and wasps) and vertebrates in tropical ecosystems of East Africa. By analysing the preferences of these three groups for nest boxes that were empty or previously occupied by a different taxon, we try to establish whether the relationships among them are commensal, mutualistic, competitive or amensal. Vertebrates and honeybees selected nest boxes that had previously been occupied by the other, which suggests that each obtains some benefit from the other. This relationship can be considered mutualistic, although a mutual preference for each others’ nests does not exclude a competitive interaction. Vertebrates and wasps preferred nest boxes not previously occupied by the other, which suggests that they compete for tree cavities. Finally, wasps seemed to completely refuse cavities previously used by honeybees, while the bees occupied cavities regardless of whether they had been previously used by wasps, an apparently amensal relationship. These results indicate that the interdependence between distantly related taxa is stronger and more complex than previously described, which may have important implications for population dynamics and community structure.

     

Earliest zygodactyl bird feet: evidence from Early Cretaceous roadrunner-like tracks

Fossil footprints are important in understanding Cretaceous avian diversity because they constitute evidence of paleodiversity and paleoecology that is not always apparent from skeletal remains. Early Cretaceous bird tracks have demonstrated the existence of wading birds in East Asia, but some pedal morphotypes, such as zygodactyly, common in modern and earlier Cenozoic birds (Neornithes) were unknown in the Cretaceous. We, herein, discuss the implications of a recently reported, Early Cretaceous (120–110 million years old) trackway of a large, zygodactyl bird from China that predates skeletal evidence of this foot morphology by at least 50 million years and includes the only known fossil zygodactyl footprints. The tracks demonstrate the existence of a Cretaceous bird not currently represented in the body fossil record that occupied a roadrunner (Geococcyx)-like niche, indicating a previously unknown degree of Cretaceous avian morphological and behavioral diversity that presaged later Cenozoic patterns. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.

A drowned Mesozoic bird breeding colony from the Late Cretaceous of Transylvania

Despite a rapidly improving fossil record, the reproductive biology of Mesozoic birds remains poorly known: only a handful of undisputed, isolated Cretaceous eggs (some containing embryonic remains) are known. We report here the first fossil evidence for a breeding colony of Mesozoic birds, preserved at the Late Cretaceous (Maastrichtian) Oarda de Jos (Od) site in the Sebe? area of Transylvania, Romania. A lens of calcareous mudstone with minimum dimensions of 80?cm length, 50?cm width and 20?cm depth contains thousands of tightly packed, morphologically homogenous eggshell fragments, seven near-complete eggs and neonatal and adult avialan skeletal elements. Eggshell forms 70–80?% of the matrix, and other fossils are entirely absent. The bones exhibit clear characters of the Cretaceous avialan clade Enantiornithes, and the eggshell morphology is also consistent with this identification. Both taphonomy and lithology show that the components of this lens were deposited in a single flood event, and we conclude that it represents the drowned remains of a larger enantiornithine breeding colony, swamped by rising water, washed a short distance and deposited in a shallow, low-energy pond. The same fate often befalls modern bird colonies. Such a large concentration of breeding birds suggests aquatic feeding in this species, augments our understanding of enantiornithine biology and shows that colonial nesting was not unique to crown birds.