A new avialan theropod from an emerging Jurassic terrestrial fauna.

Birds are descended from non-avialan theropod dinosaurs of the Late Jurassic period, but the earliest phase of this evolutionary process remains unclear owing to the exceedingly sparse and spatio-temporally restricted fossil record1-5. Information about the early-diverging species along the avialan line is crucial to understand the evolution of the characteristic bird bauplan, and to reconcile phylogenetic controversies over the origin of birds3,4. Here we describe one of the stratigraphically youngest and geographically southernmost Jurassic avialans, Fujianvenator prodigiosus gen. et sp. nov., from the Tithonian age of China. This specimen exhibits an unusual set of morphological features that are shared with other stem avialans, troodontids and dromaeosaurids, showing the effects of evolutionary mosaicism in deep avialan phylogeny. F. prodigiosus is distinct from all other Mesozoic avialan and non-avialan theropods in having a particularly elongated hindlimb, suggestive of a terrestrial or wading lifestyle-in contrast with other early avialans, which exhibit morphological adaptations to arboreal or aerial environments. During our fieldwork in Zhenghe where F. prodigiosus was found, we discovered a diverse assemblage of vertebrates dominated by aquatic and semi-aquatic species, including teleosts, testudines and choristoderes. Using in situ radioisotopic dating and stratigraphic surveys, we were able to date the fossil-containing horizons in this locality-which we name the Zhenghe Fauna-to 148-150 million years ago. The diversity of the Zhenghe Fauna and its precise chronological framework will provide key insights into terrestrial ecosystems of the Late Jurassic.

Plant-hummingbird pollination networks exhibit limited rewiring after experimental removal of a locally abundant plant species.

Mutualistic relationships, such as those between plants and pollinators, may be vulnerable to the local extinctions predicted under global environmental change. However, network theory predicts that plant-pollinator networks can withstand species loss if pollinators switch to alternative floral resources (rewiring). Whether rewiring occurs following species loss in natural communities is poorly known because replicated species exclusions are difficult to implement at appropriate spatial scales. We experimentally removed a hummingbird-pollinated plant, Heliconia tortuosa, from within tropical forest fragments to investigate how hummingbirds respond to temporary loss of an abundant resource. Under the rewiring hypothesis, we expected that behavioural flexibility would allow hummingbirds to use alternative resources, leading to decreased ecological specialization and reorganization of the network structure (i.e. pairwise interactions). Alternatively, morphological or behavioural constraints-such as trait-matching or interspecific competition-might limit the extent to which hummingbirds alter their foraging behaviour. We employed a replicated Before-After-Control-Impact experimental design and quantified plant-hummingbird interactions using two parallel sampling methods: pollen collected from individual hummingbirds ('pollen networks', created from >300 pollen samples) and observations of hummingbirds visiting focal plants ('camera networks', created from >19,000 observation hours). To assess the extent of rewiring, we quantified ecological specialization at the individual, species and network levels and examined interaction turnover (i.e. gain/loss of pairwise interactions). H. tortuosa removal caused some reorganization of pairwise interactions but did not prompt large changes in specialization, despite the large magnitude of our manipulation (on average, >100 inflorescences removed in exclusion areas of >1 ha). Although some individual hummingbirds sampled through time showed modest increases in niche breadth following Heliconia removal (relative to birds that did not experience resource loss), these changes were not reflected in species- and network-level specialization metrics. Our results suggest that, at least over short time-scales, animals may not necessarily shift to alternative resources after losing an abundant food resource-even in species thought to be highly opportunistic foragers, such as hummingbirds. Given that rewiring contributes to theoretical predictions of network stability, future studies should investigate why pollinators might not expand their diets after a local resource extinction.

Bene"fit" Assessment in Pollination Coevolution: Mechanistic Perspectives on Hummingbird Bill-Flower Matching.

One of the reasons why flowering plants became the most diverse group of land plants is their association with animals to reproduce. The earliest examples of this mutualism involved insects foraging for food from plants and, in the process, pollinating them. Vertebrates are latecomers to these mutualisms, but birds, in particular, present a wide variety of nectar-feeding clades that have adapted to solve similar challenges. Such challenges include surviving on small caloric rewards widely scattered across the landscape, matching their foraging strategy to nectar replenishment rate, and efficiently collecting this liquid food from well-protected chambers deep inside flowers. One particular set of convergent traits among plants and their bird pollinators has been especially well studied: the match between the shape and size of bird bills and ornithophilous flowers. Focusing on a highly specialized group, hummingbirds, we examine the expected benefits from bill-flower matching, with a strong focus on the benefits to the hummingbird and how to quantify them. Explanations for the coevolution of bill-flower matching include (1) that the evolution of traits by bird-pollinated plants, such as long and thin corollas, prevents less efficient pollinators (e.g., insects) from accessing the nectar and (2) that increased matching, as a result of reciprocal adaptation, benefits both the bird (nectar extraction efficiency) and the plant (pollen transfer). In addition to nectar-feeding, we discuss how interference and exploitative competition also play a significant role in the evolution and maintenance of trait matching. We present hummingbird-plant interactions as a model system to understand how trait matching evolves and how pollinator behavior can modify expectations based solely on morphological matching, and discuss the implications of this behavioral modulation for the maintenance of specialization. While this perspective piece directly concerns hummingbird-plant interactions, the implications are much broader. Functional trait matching is likely common in coevolutionary interactions (e.g., in predator-prey interactions), yet the physical mechanisms underlying trait matching are understudied and rarely quantified. We summarize existing methods and present novel approaches that can be used to quantify key benefits to interacting partners in a variety of ecological systems.

Vertebral pneumaticity is correlated with serial variation in vertebral shape in storks.

Birds and their ornithodiran ancestors are unique among vertebrates in exhibiting air-filled sinuses in their postcranial bones, a phenomenon called postcranial skeletal pneumaticity. The factors that account for serial and interspecific variation in postcranial skeletal pneumaticity are poorly understood, although body size, ecology, and bone biomechanics have all been implicated as influencing the extent to which pneumatizing epithelia invade the skeleton and induce bone resorption. Here, I use high-resolution computed-tomography to holistically quantify vertebral pneumaticity in members of the neognath family Ciconiidae (storks), with pneumaticity measured as the relative volume of internal air space. These data are used to describe serial variation in extent of pneumaticity and to assess whether and how pneumaticity varies with the size and shape of a vertebra. Pneumaticity increases dramatically from the middle of the neck onwards, contrary to previous predictions that cervical pneumaticity should decrease toward the thorax to maintain structural integrity as the mass and bending moments of the neck increase. Although the largest vertebrae sampled are also the most pneumatic, vertebral size cannot on its own account for serial or interspecific variation in extent of pneumaticity. Vertebral shape, as quantified by three-dimensional geometric morphometrics, is found to be significantly correlated with extent of pneumaticity, with elongate vertebrae being less pneumatic than craniocaudally short and dorsoventrally tall vertebrae. Considered together, the results of this study are consistent with the hypothesis that shape- and position-specific biomechanics influence the amount of bone loss that can be safely tolerated. These results have potentially important implications for the evolution of vertebral morphology in birds and their extinct relatives.

Inter-amphibian predation in the Early Cretaceous of China.

For most fossil taxa, dietary inference relies primarily on indirect evidence from jaw morphology and the dentition. In rare cases, however, preserved gut contents provide direct evidence of feeding strategy and species interaction. This is important in the reconstruction of food webs and energy flow through ancient ecosystems. The Early Cretaceous Chinese Jehol Biota has yielded several such examples, with lizards, birds, small dinosaurs, and mammals as both predator and prey. Here we describe an Early Cretaceous fossil frog specimen, genus Genibatrachus, that contains an adult salamander within its body cavity. The salamander is attributed to the hynobiid-like genus Nuominerpeton. The salamander skeleton is complete and articulated, suggesting it was caught and swallowed shortly before the frog itself died and was buried.

Archaeorhynchus preserving significant soft tissue including probable fossilized lungs.

We describe a specimen of the basal ornithuromorph Archaeorhynchus spathula from the Lower Cretaceous Jiufotang Formation with extensive soft tissue preservation. Although it is the fifth specimen to be described, unlike the others it preserves significant traces of the plumage, revealing a pintail morphology previously unrecognized among Mesozoic birds, but common in extant neornithines. In addition, this specimen preserves the probable remnants of the paired lungs, an identification supported by topographical and macro- and microscopic anatomical observations. The preserved morphology reveals a lung very similar to that of living birds. It indicates that pulmonary specializations such as exceedingly subdivided parenchyma that allow birds to achieve the oxygen acquisition capacity necessary to support powered flight were present in ornithuromorph birds 120 Mya. Among extant air breathing vertebrates, birds have structurally the most complex and functionally the most efficient respiratory system, which facilitates their highly energetically demanding form of locomotion, even in extremely oxygen-poor environments. Archaeorhynchus is commonly resolved as the most basal known ornithuromorph bird, capturing a stage of avian evolution in which skeletal indicators of respiration remain primitive yet the lung microstructure appears modern. This adds to growing evidence that many physiological modifications of soft tissue systems (e.g., digestive system and respiratory system) that characterize living birds and are key to their current success may have preceded the evolution of obvious skeletal adaptations traditionally tracked through the fossil record.

A bizarre Early Cretaceous enantiornithine bird with unique crural feathers and an ornithuromorph plough-shaped pygostyle.

Enantiornithes are the most successful clade of Mesozoic birds. Here, we describe a new enantiornithine bird, Cruralispennia multidonta gen. et sp. nov., from the Protopteryx-horizon of the Early Cretaceous Huajiying Formation of China. Despite being among the oldest known enantiornithines, Cruralispennia displays derived morphologies that are unexpected at such an early stage in the evolution of this clade. A plough-shaped pygostyle, like that of the Ornithuromorpha, evolved convergently in the Cruralispennia lineage, highlighting the homoplastic nature of early avian evolution. The extremely slender coracoid morphology was previously unknown among Early Cretaceous enantiornithines but is common in Late Cretaceous taxa, indicating that by 131 million years ago this clade had already experienced considerable morphological differentiation. Cruralispennia preserves unusual crural feathers that are proximally wire-like with filamentous distal tips, a new morphotype previously unknown among fossil or modern feathers, further increasing the known diversity of primitive feather morphologies.

Archaeopteryx feathers and bone chemistry fully revealed via synchrotron imaging.

Evolution of flight in maniraptoran dinosaurs is marked by the acquisition of distinct avian characters, such as feathers, as seen in Archaeopteryx from the Solnhofen limestone. These rare fossils were pivotal in confirming the dinosauria-avian lineage. One of the key derived avian characters is the possession of feathers, details of which were remarkably preserved in the Lagerstätte environment. These structures were previously simply assumed to be impressions; however, a detailed chemical analysis has, until now, never been completed on any Archaeopteryx specimen. Here we present chemical imaging via synchrotron rapid scanning X-ray fluorescence (SRS-XRF) of the Thermopolis Archaeopteryx, which shows that portions of the feathers are not impressions but are in fact remnant body fossil structures, maintaining elemental compositions that are completely different from the embedding geological matrix. Our results indicate phosphorous and sulfur retention in soft tissue as well as trace metal (Zn and Cu) retention in bone. Other previously unknown chemical details of Archaeopteryx are also revealed in this study including: bone chemistry, taphonomy (fossilization process), and curation artifacts. SRS-XRF represents a major advancement in the study of the life chemistry and fossilization processes of Archaeopteryx and other extinct organisms because it is now practical to image the chemistry of large specimens rapidly at concentration levels of parts per million. This technique has wider application to the archaeological, forensic, and biological sciences, enabling the mapping of "unseen" compounds critical to understanding biological structures, modes of preservation, and environmental context.

New flutes document the earliest musical tradition in southwestern Germany.

Considerable debate surrounds claims for early evidence of music in the archaeological record. Researchers universally accept the existence of complex musical instruments as an indication of fully modern behaviour and advanced symbolic communication but, owing to the scarcity of finds, the archaeological record of the evolution and spread of music remains incomplete. Although arguments have been made for Neanderthal musical traditions and the presence of musical instruments in Middle Palaeolithic assemblages, concrete evidence to support these claims is lacking. Here we report the discovery of bone and ivory flutes from the early Aurignacian period of southwestern Germany. These finds demonstrate the presence of a well-established musical tradition at the time when modern humans colonized Europe, more than 35,000 calendar years ago. Other than the caves of the Swabian Jura, the earliest secure archaeological evidence for music comes from sites in France and Austria and post-date 30,000 years ago.

A gigantic bird-like dinosaur from the Late Cretaceous of China.

An evolutionary trend of decreasing size is present along the line to birds in coelurosaurian theropod evolution, but size increases are seen in many coelurosaurian subgroups, in which large forms are less bird-like. Here we report on a new non-avian dinosaur, Gigantoraptor erlianensis, gen. et sp. nov., from the Late Cretaceous Iren Dabasu Formation of Nei Mongol, China. Although it has a body mass of about 1,400 kg, a phylogenetic analysis positions this new taxon within the Oviraptorosauria, a group of small, feathered theropods rarely exceeding 40 kg in body mass. A histological analysis suggests that Gigantoraptor gained this size by a growth rate considerably faster than large North American tyrannosaurs such as Albertosaurus and Gorgosaurus. Gigantoraptor possesses several salient features previously unknown in any other dinosaur and its hind limb bone scaling and proportions are significantly different from those of other coelurosaurs, thus increasing the morphological diversity among dinosaurs. Most significantly, the gigantic Gigantoraptor shows many bird-like features absent in its smaller oviraptorosaurian relatives, unlike the evolutionary trend seen in many other coelurosaurian subgroups.

Adaptive trade-off in floral morphology mediates specialization for flowers pollinated by bats and hummingbirds.

Evolution toward increased specificity in pollination systems is thought to have played a central role in the diversification of angiosperms. Theory predicts that the presence of trade-offs in adapting to different pollinator types will favor specialization, yet few studies have attempted to characterize such interactions in nature. I conducted flight cage experiments with bats, hummingbirds, and artificial flowers to examine effects of corolla width on pollination. I videotaped visits to analyze pollinator behavior and counted pollen grains transferred to stigmas. Results demonstrated that flower-pollinator fit is critical to effective pollination; wide corollas guided bat snouts better, and narrow corollas guided hummingbird bills better. Poor fit resulted in variable entry angles and decreased pollen transfer. A model using these results predicts that wide corollas will be selected for when bats make more than 44% of the visits and narrow corollas when they make fewer. Intermediate corollas are never favored (i.e., generalization is always suboptimal). This is the first study to clearly document a pollinator-mediated fitness trade-off in floral morphology.

Origin of flight: Could 'four-winged' dinosaurs fly?

Our understanding of the origin of birds, feathers and flight has been greatly advanced by new discoveries of feathered non-avian dinosaurs, but functional analyses have not kept pace with taxonomic descriptions. Zhang and Zhou describe feathers on the tibiotarsus of a new basal enantiornithine bird from the Early Cretaceous of China. They infer, as did Xu and colleagues from similar feathers on the small non-avian theropod Microraptor found in similar deposits, that these leg feathers had aerodynamic properties and so might have been used in some kind of flight.

Thalamotelencephalic organization in birds.

Investigation of thalamo-telencephalic connections reveals correspondences between the avian and mammalian thalamic subdivisions (which may or may not mean true homologies). Based mainly on hodological comparisons, the avian thalamus possesses the principal anatomical and functional subdivisions characteristic for mammals. The current review is focused on a comparative analysis of intralaminar, midline and mediodorsal nuclei. There is evidence for matching subdivisions in the case of midline thalamic and mediodorsal nuclei within the avian dorsal thalamic zone, whereas such correspondence is evident, if less complete, in the case of the intralaminar nuclei. Thalamic connections are also relevant to the debated issue of the avian 'prefrontal' cortex. From the current study it is suggested that the prefrontal analogue regions of the bird may spread across the rostrocaudal extent of telencephalon, the rostral nidopallial/mesopallial region (formerly known as medial neostriatum/hyperstriatum) being one subdivision, receiving direct input from the paraventricular thalamic nucleus homologue of midline thalamic region (the medial juxtaventricular region of the nucleus dorsomedialis posterior). Hodological evidence from the current study and other reports argues for the possibility that the area corticoidea dorsolateralis might be hodologically comparable to the cingulate cortex, receiving input from a mediodorsal thalamic-relevant subdivision (lateral subdivision of nucleus dorsomedialis anterior, and medial aspect of nucleus dorsolateralis pars medialis), which also projects on the caudal nidopallium close to (but not coextensive with) the nidopallium caudolaterale, another potential analogue of avian prefrontal cortex. The rostral dorsolateral aspect of nucleus dorsomedialis anterior thalami and the dorsal aspect of nucleus dorsolateralis pars medialis are partially comparable to the mammalian intralaminar nuclei, sharing connections to non-limbic 'corticoid' areas (the Wulst), and the reticular thalamic nuclei.

Feeding mechanisms: Hummingbird jaw bends to aid insect capture.

The upper jaws of birds, unlike those in many tetrapods, move relative to the skull and are often flexible along their length, whereas the lower jaw (mandible) is usually a rigid structure formed by the fusion of several bones, flexing only where it meets the skull. Here we describe a previously unnoticed mandibular bending movement in hummingbirds, in which the distal half of the mandible is actively flexed downwards and the gape widens to catch flying insects. The hummingbird is thought to have developed a long narrow bill as it specialized in feeding on floral nectar, but the bird's need to supplement its diet with insects must have contributed to the surprising flexibility of its jaw.

New Early Cretaceous fossil from China documents a novel trophic specialization for Mesozoic birds.

We report on a new Mesozoic bird, Longirostravis hani, from the Early Cretaceous Jehol Biota of northeastern China. The new taxon has a long, slender rostrum and mandible, and a small number of rostralmost teeth. Postcranial characters such as a furcular ramus wider ventrally than dorsally, a centrally concave proximal margin of the humeral head, and a minor metacarpal that projects distally more than the major metacarpal, support the placement of Longirostravis within euenantiornithine Enantiornithes, the most diverse clade of Mesozoic birds. The morphology of the skull, however, suggests that Longirostravis had a probing feeding behavior, a specialization previously unknown for Enantiornithes. Indeed, this discovery provides the first evidence in support of the existence of such a foraging behavior among basal lineages of Mesozoic birds.

Stimulatory effects of pituitary adenylate cyclase-activating polypeptide on inositol phosphates accumulation in avian cerebral cortex and hypothalamus.

This study has demonstrated that the short and long form of the pituitary adenylate cyclase-activating polypeptide (PACAP), i.e. PACAP(27) and PACAP(38), moderately but significantly, and in a concentration (0.5-5 microM)-dependent manner, stimulated inositol phosphates (IPs) accumulation in myo-[(3)H]inositol-prelabeled cerebral cortical and hypothalamal slices of chick and duck, and in slices of rat cerebral cortex; both peptides had no effect on IPs formation in rat hypothalamus. Vasoactive intestinal peptide (VIP; 0.5-5 microM) weakly enhanced IPs accumulation in chick hypothalamus, had no significant action in chick cerebral cortex (in fact there was a tendency to attenuate the IPs response in this tissue), and slightly, but significantly, inhibited the IPs accumulation in rat cerebral cortex. VIP showed no activity in rat hypothalamus. It is concluded that the stimulatory action of PACAP on phosphoinositide metabolism in avian cerebral cortex, similar to rat cerebral cortex, is mediated via phospholipase C-linked PAC(1) type receptors. In chick hypothalamus, however, there may be a component of VPAC type receptors stimulating IPs formation.

Domesticated birds as a model for the genetics of speciation by sexual selection.

In theory, even populations occupying identical environments can diverge in sexually selected traits, as a consequence of different mutational input. I evaluate the potential of this process by comparing the genetics of breeds of domesticated birds to what is known about the genetics of differences among species. Within domesticated species there is a strong correlation of time since domestication with the number of breeds. Descendants of the rock dove, Columba livia (the oldest domesticate) show differences in courtship, vocalizations, body shape, feather ornaments (crests and tails) and colors and color patterns. When nine other domesticated species are included there is a striking hierarchy, with more recent domesticates having a nested subset of these traits: the youngest domesticated species have breeds distinguished only by color. This suggests that selection of new, visible, mutations is driving the process of breed diversification, with mutations that appeal to the breeder happening the most frequently in color. In crosses among related species, color, feather ornaments and many vocalizations and displays show both intermediate dominance and pure dominance. Although the number of loci affecting each of these traits is typically unknown, limited evidence of the genetics of species' differences suggests that some differences are due to the substitution of single genes of major effect. While neither the genetics of breeds nor the genetics of species provide a perfect model for the genetics of speciation, similarities between the two are sufficiently striking to infer that major, visible, mutations can provide the impetus underlying new directions of sexual selection.