Evolution of olfaction in non-avian theropod dinosaurs and birds

Little is known about the olfactory capabilities of extinct basal (non-neornithine) birds or the evolutionary changes in olfaction that occurred from non-avian theropods through modern birds. Although modern birds are known to have diverse olfactory capabilities, olfaction is generally considered to have declined during avian evolution as visual and vestibular sensory enhancements occurred in association with flight. To test the hypothesis that olfaction diminished through avian evolution, we assessed relative olfactory bulb size, here used as a neuroanatomical proxy for olfactory capabilities, in 157 species of non-avian theropods, fossil birds and living birds. We show that relative olfactory bulb size increased during non-avian maniraptoriform evolution, remained stable across the non-avian theropod/bird transition, and increased during basal bird and early neornithine evolution. From early neornithines through a major part of neornithine evolution, the relative size of the olfactory bulbs remained stable before decreasing in derived neoavian clades. Our results show that, rather than decreasing, the importance of olfaction actually increased during early bird evolution, representing a previously unrecognized sensory enhancement. The relatively larger olfactory bulbs of earliest neornithines, compared with those of basal birds, may have endowed neornithines with improved olfaction for more effective foraging or navigation skills, which in turn may have been a factor allowing them to survive the end-Cretaceous mass extinction.     

Ventilatory mechanics from maniraptoran theropods to extant birds

Shared behavioural, morphological and physiological characteristics are indicative of the evolution of extant birds from nonavian maniraptoran dinosaurs. One such shared character is the presence of uncinate processes and respiratory structures in extant birds. Recent research has suggested a respiratory role for these processes found in oviraptorid and dromaeosaurid dinosaurs. By measuring the geometry of fossil rib cage morphology, we demonstrate that the mechanical advantage, conferred by uncinate processes, for movements of the ribs in the oviraptorid theropod dinosaur, Citipati osmolskae, basal avialan species Zhongjianornis yangi, Confuciusornis sanctus and the more derived ornithurine Yixianornis grabaui, is of the same magnitude as found in extant birds. These skeletal characteristics provide further evidence of a flow-through respiratory system in nonavian theropod dinosaurs and basal avialans, and indicate that uncinate processes are a key adaptation facilitating the ventilation of a lung air sac system that diverged earlier than extant birds.     

Mesozoic birds of China-a synoptic review

A synoptic review of the discoveries and studies of Chinese Mesozoic birds is provided in this paper.40Ar/39Ar dating of several bird-bearing deposits in the Jehol Group has established a geochronological framework for the study of the early avian radiation.Chinese Mesozoic birds had lasted for at least 11 Ma during about 131 Ma and 120 Ma (Barremian to Aptian)of the middle and late Early Cretaceous,respectively.In order to further evaluate the change of the avian diversity in the Jehol Biota,six new orders and families are erected based on known genera and species,which brings the total number of orders of Chinese Mesozoic birds to 15 and highlights a remarkable radiation ever since the first appearante of birds in the Late Jurassic.Chinese Early Cretaceous birds had experienced a significant differentiation in morphology,flight,diet and habitat.Further examination of the foot of Jeholornis suggests this bird might not have possessed a fully reversed hallux.However,the attachment of metatarsal Ⅰ to the medial side of metatarsal Ⅱ does not preclude trunk climbing,a pre-adaptation for well developed perching life of early birds.Arboreality had proved to be a key adaptation in the origin and early evolution of bird flight,and the adaptation to lakeshore environment had played an equally important role in the origin of omithurine birds and their near-modern flight skill.Many Chinese Early Cretaceous birds had preserved the direct evidence of their diet,showing that the most primitive birds were probably mainly insectivorous and that specialized herbivorous or carnivorous (e.g.,piscivorous)dietary adaptation had appeared only in later advanced forms.The only known Early Cretaceous bird embryo fossil has shown that precocial birds had occurred prior to altricial birds in avian history,and the size of the embryo and other analysis indicate it probably had a short incubation period.Leg feathers probably have a wide range of distribution in early birds,further suggesting that leg feathers had played a key role in the beginning stage of the flight of birds.Finally,the Early Cretaceous avian radiation can be better understood against the background of their unique ecosystem.The advantage of birds in the competitions with other vertebrate groups such as pterosaurs had probably not only resulted in the rapid differentiation and radiation of birds but also the worldwide spreading of pterosaurs and other vertebrates from East Asia in the Early Cretaceous.     

Body size evolution in Mesozoic birds

The tendency for the mean body size of taxa within a clade to increase through evolution (Cope's Rule) has been demonstrated in a number of terrestrial vertebrate groups. However, because avian body size is strongly constrained by flight, any increase in size during the evolution of this lineage should be limited - there is a maximum size that can be attained by a bird for it to be able to get off the ground. Contrary to previous interpretations of early avian evolution, we demonstrate an overall increase in body size across Jurassic and Cretaceous flying birds: taxon body size increases from the earliest Jurassic through to the end of the Cretaceous, across a time span of 70 Myr. Although evidence is limited that this change is directional, it is certainly nonrandom. Relative size increase occurred presumably as the result of an increase in variance as the avian clade diversified after the origin of flight: a progression towards larger body size is seen clearly within the clades Pygostylia and Ornithothoraces. In contrast, a decrease in body size characterizes the most crownward lineage Ornithuromorpha, the clade that includes all extant taxa, and potentially may explain the survival of these birds across the Cretaceous-Palaeogene boundary. As in all other dinosaurs, counter selection for small size is seen in some clades, whereas body size is increasing overall.     

Foraging energetics of arctic cormorants and the evolution of diving birds

Efficient body insulation is assumed to have enabled birds and mammals to colonize polar aquatic ecosystems. We challenge this concept by comparing the bioenergetics of cormorants ( Phalacrocorax carbo ) living in temperate and arctic conditions. We show that although these birds have limited insulation, they maintain high body temperature (42.3 °C) when diving in cold water (1–10 °C). Their energy demand at these times is extremely high (up to 60 W kg⁻¹). Free-living cormorants wintering in Greenland (water temperature −1 °C) profoundly alter their foraging activity, thus minimizing time spent in water and the associated high thermoregulatory costs. They then meet their daily food demand within a single intense dive bout (lasting 9 min on average). Their substantial energy requirements are balanced by the highest predatory efficiency so far recorded for aquatic predators. We postulate that similar behavioural patterns allowed early diving birds (Cretaceous) to colonize cold coastal areas before they evolved efficient insulation.     

The scapulocoracoid of flightless birds: a primitive avian character similar to that of theropods

The scapula and coracoid of the first bird Archaeopteryx represents a highly specialized morphology approaching that of modern carinate birds, with the scapula meeting the coracoid at an angle of approximately 90^o. The primitive condition of the avian scapula and coracoid is exhibited by the flightless ratites, birds exhibiting many other primitive characters of the skeleton, either retained from primitive ancestors or derived through neoteny. The fact that the scapulocoracoid of ratites is the primitive condition is confirmed by its presence in the embryos of certain birds, and its presence through neoteny in other unrelated birds that have secondarily evolved Sightlessness. This morphology of the primitive avian scapulocoracoid closely approaches that of theropod dinosaurs, such as Deinonychits , and may well indicate relationship between the two groups.     

The quality of the fossil record of Mesozoic birds

The Mesozoic fossil record has proved critical for understanding the early evolution and subsequent radiation of birds. Little is known, however, about its relative completeness: just how 'good' is the fossil record of birds from the Mesozoic? This question has come to prominence recently in the debate over differences in estimated dates of origin of major clades of birds from molecular and palaeontological data. Using a dataset comprising all known fossil taxa, we present analyses that go some way towards answering this question. Whereas avian diversity remains poorly represented in the Mesozoic, many relatively complete bird specimens have been discovered. New taxa have been added to the phylogenetic tree of basal birds, but its overall shape remains constant, suggesting that the broad outlines of early avian evolution are consistently represented: no stage in the Mesozoic is characterized by an overabundance of scrappy fossils compared with more complete specimens. Examples of Neornithes (modern orders) are known from later stages in the Cretaceous, but their fossils are rarer and scrappier than those of basal bird groups, which we suggest is a biological, rather than a geological, signal.     

Foraging patterns of breeding birds in eucalypt forest and woodland of southeastern Australia

The foraging ecology of eucalypt forest and woodland birds was studied on three 10 ha plots in southeastern Australia. Quantitative data were obtained for 41 species of which 31 were insectivorous, eight were nectar-feeders, and two were parrots that fed primarily on eucalypt seeds. Birds-of-prey, large omnivores and frugivores were uncommon. Insectivorous birds differed in foraging behaviour, the substrates on which they found prey, and foraging height. Nectar- feeders exploited a variety of carbohydrates including nectar, honeydew, lerp, manna and sap. Nectarivorous birds were separated by foraging behaviour, substrate, height and by the extent to which they used the different types of carbohydrates. Carbohydrates were also an important food resource for some insectivores. By understanding how birds exploit food resources within forest and woodland environments, the features of the environment which need to be conserved or manipulated to manage forest avifaunas can be identified. For example, in addition to the substrates such as foliage and bark, usually associated with the foraging of forest birds, carbohydrates and loose bark were identified as important resources for birds in eucalypt forests and woodlands. The broad importance of these two resources to the avifauna had not been previously appreciated, yet both may be sensitive to environmental changes associated with logging and other forest management practices which alter the composition or age-class structure of forests.     

Optimal locomotion modes of foraging birds in trees

This paper compares the energy costs of various modes of locomotion of birds foraging in trees. For birds moving vertically in trees by climbing and hopping (but not by flying) the best choice of locomotion mode depends on the distance between visited trees in relation to the height h of the zone searched for food in trees.
When the distance between successively visisted trees is longer than about half the distance coverable in gliding flight with height loss h , then it is cheapest in energy to hop or climb upwards in a tree and fly downwards to the next tree. When the distance between successively visited trees is shorter than about half the distance coverable in gliding flight with height loss h , then it is cheapest in energy to move alternately downwards and upwards in trees (downwards in the first, upwards in the second, downwards in the third tree, etc.) and to fly level between trees.
Treecreepers and woodpeckers are adapted morphologically to the former mode, but more generalized tree foragers might use either mode depending on the spacing of trees.     

The craniofacial air sac system of Mesozoic birds (Aves)

Birds are characterized by pneumatization of their skeletons by epithelial diverticula from larger, air—filled cavities. The diverticula—or 'air sacs'—that invade the postcranium result from outgrowths of the lungs; postcranial pneumaticity has been very well studied. Much more poorly understood are the air sacs that pneumatize the skull. Study of craniofacial pneumaticity in modern birds (Neornithes) indicates the presence of two separate systems: nasal pneumaticity and tympanic pneumaticity. The lacrimal and maxillary bones are pneumatized by diverticula of the main paranasal cavity, the antorbital sinus. There are five tympanic diverticula in neornithines that pneumatic the quadrate, articulare and the bones of the braincase. The pneumatic features of the following six genera of Mesozoic birds are examined: Archaeopteryx, Enaliornis, Baptornis, Parahesperornis, Hesperornis and Ichthyornis. Despite the 'archaic' aspect of most of these birds, many of the pneumatic features of neornithines are found in Mesozoic birds and are considered primitive for Aves. The phylogenetic levels at which most of the avian pneumatic features arose within Archosauria are uncertain. Until the phylogenetic levels at which homologous pneumatic features arose are determined, it is unwise to use most pneumatic characters in the discussion of avian origins. Within avian phylogeny, Ornithurae and Neornithes are well–supported by pneumatic synapomorphies. There is a trend towards reduction of craniofacial pneumaticity within Hesperornithiformes. Within Neornithes, four derived pneumatic characters suggest that the Palaeognathae (ratites and tinamous) is monophyletic.     

Prevalence of different modes of parental care in birds

Estimates of the incidence of major classes of parental care by birds are drawn from classical studies that preceded both the publication of a massive secondary literature and the revolution driven by molecular approaches to avian phylogeny. Here, I review this literature in the light of new phylogenetic hypotheses and estimate the prevalence of six distinct modes of care: use of geothermal heat to incubate eggs, brood parasitism, male only care, female only care, biparental care and cooperative breeding. Female only care and cooperative breeding are more common than has previously been recognized, occurring in 8 and 9% of species, respectively. Biparental care by a pair-bonded male and female is the most common pattern of care but at 81% of species, the pattern is less common than once believed. I identify several problems with existing hypotheses for the evolution of parental care and highlight a number of poorly understood contrasts which, once resolved, should help elucidate avian social evolution.     

Foraging ecology of an assemblage of birds in lowland rainforest in northern Queensland

This paper presents an analysis of the foraging ecologies and hence means of partitioning resources amongst the twenty-eight most conspicuous species of birds in lowland tropical rainforest at Lacey's Creek, North Queensland. The height and site of foraging and the type of behaviour used by the species were compared. The height at which species foraged appeared to be more important for separating species than the site of foraging or the behaviour used. There appeared to be a greater proportion of species with generalized foraging behaviour at Lacey's Creek than in bird communities in the neotropics. Frugivorous species had very similar foraging ecologies but there were differences in the species of fruit eaten by some species. There were seasonal shifts in foraging by some species. It is postulated that the relative shortage of terrestrial species at Lacey's Creek compared with highland forest in North Queensland is due to the harsh dry season in the lowlands. The structure of this bird community appeared to be strongly affected by seasonality of climate.     

Body size evolution in Mesozoic birds: little evidence for Cope's rule

Cope's rule, the tendency towards evolutionary increases in body size, is a long-standing macroevolutionary generalization that has the potential to provide insights into directionality in evolution; however, both the definition and identification of Cope's rule are controversial and problematic. A recent study [J. Evol. Biol. 21 (2008) 618] examined body size evolution in Mesozoic birds, and claimed to have identified evidence of Cope's rule occurring as a result of among-lineage species sorting. We here reassess the results of this study, and additionally carry out novel analyses testing for within-lineage patterns in body size evolution in Mesozoic birds. We demonstrate that the nonphylogenetic methods used by this previous study cannot distinguish between among- and within-lineage processes, and that statistical support for their results and conclusions is extremely weak. Our ancestor-descendant within-lineage analyses explicitly incorporate recent phylogenetic hypotheses and find little compelling evidence for Cope's rule. Cope's rule is not supported in Mesozoic birds by the available data, and body size evolution currently provides no insights into avian survivorship through the Cretaceous-Paleogene mass extinction.     

Foraging guilds of aquatic birds on productive boreal lakes: environmental relations and concordance patterns

Previous studies have shown that when an ecosystem consists of many interacting components it becomes impossible to understand how it functions by focussing only on individual relationships. Alternatively, one can attempt to quantify system behaviour as a whole by developing ecological indicators that combine numerous environmental factors into a single value. One such holistic measure, called the system ‘ascendency’, arises from the analysis of networks of trophic exchanges. It deals with the joint quantification of overall system activity with the organisation of the component processes and can be used specifically to identify the occurrence of eutrophication. System ascendency analyses were applied to data over a gradient of eutrophication in a well documented small temperate intertidal estuary. Three areas were compared along the gradient, respectively, non eutrophic, intermediate eutrophic, and strongly eutrophic. Values of other measures related to the ascendency, such as the total system throughput, development capacity, and average mutual information, as well as the ascendency itself, were clearly higher in the non-eutrophic area. When the whole-system properties of the three areas were compared, however, the values associated with the intermediate eutrophic area turned out to be the lowest, which possibly could be attributed to the unstable nature of this area. The current study provided an example of how the measures arising out of␣network analysis might lead to an improved understanding of the system functioning and of the eutrophication process itself.     

Size scaling and stiffness of avian primary feathers: implications for the flight of Mesozoic birds

The primary feathers of birds are subject to cyclical forces in flight causing their shafts (rachises) to bend. The amount the feathers deflect during flight is dependent upon the flexural stiffness of the rachises. By quantifying scaling relationships between body mass and feather linear dimensions in a large data set of living birds, we show that both feather length and feather diameter scale much closer to predictions for geometric similarity than they do to elastic similarity. Scaling allometry also indicates that the primary feathers of larger birds are relatively shorter and their rachises relatively narrower, compared to those of smaller birds. Two-point bending tests indicated that larger birds have more flexible feathers than smaller species. Discriminant functional analyses (DFA) showed that body mass, primary feather length and rachis diameter can be used to differentiate between different magnitudes of feather bending stiffness, with primary feather length explaining 63% of variance in rachis stiffness. Adding fossil measurement data to our DFA showed that Archaeopteryx and Confuciusornis do not overlap with extant birds. This strongly suggests that the bending stiffness of their primary feathers was different to extant birds and provides further evidence for distinctive flight styles and likely limited flight ability in Archaeopteryx and Confuciusornis.     

Foraging niche dynamics and overlap in a guild of passerine birds in a south Swedish coniferous woodland

Summary Three foraging niche dimensions were measured at four seasons in six passerine bird species in a pine-dominated woodland. Total niche size increased in the order Certhia familiaris. In the first two species niche sizewas relatively constant at all seasons in spite of fluctuations in separatedimensions suggesting intraspecific complementarity between dimensionbreadths. Diffuse niche overlap was much lower in summer than at otherseasons in all species. Generally, this was due to movements of exploitationcurves along resource axes away from zones of interspecific overlap andto enlargement of total guild niche size, rather than to a reduction in totalniche size of the species. In some species a positive correlation was indicatedbetween breadth of niche dimension and diffuse overlap in the same dimension.Total niche size was correlated with total diffuse overlap up to acertain overlap value. Interspecific niche size complementarity was stronglysuggested by total guild niche being much less variable between seasons thanthe separate niche size of each species. The narrow niche of P. major, usuallyregarded a generalist, may be due to it being primarily adapted to broadleavedrather than coniferous woodland.     

Primary feather lengths may not be important for inferring the flight styles of Mesozoic birds

Chan, N.R., Dyke, G.J. & Benton, M.J. 2013: Primary feather lengths may not be important for inferring the flight styles of Mesozoic birds. Lethaia, Vol. 46, pp. 146–152. Although many Mesozoic fossil birds have been found with primary feathers preserved, these structures have rarely been included in morphometric analyses. This is surprising because the flight feathers of modern birds can contribute approximately 50% of the total wing length, and so it would be assumed that their inclusion or exclusion would modify functional interpretations. Here we show, contrary to earlier work, that this may not be the case. Using forelimb measurements and primary feather lengths from Mesozoic birds, we constructed morphospaces for different clades, which we then compared with morphospaces constructed for extant taxa classified according to flight mode. Consistent with older work, our results indicate that among extant birds some functional flight groups can be distinguished on the basis of their body sizes and that variation in the relative proportions of the wing elements is conservative. Mesozoic birds, on the other hand, show variable proportions of wing bones, with primary feather length contribution to the wing reduced in the earlier diverging groups. We show that the diverse Mesozoic avian clade Enantiornithes overlaps substantially with extant taxa in both size and limb element proportions, confirming previous morphometric results based on skeletal elements alone. However, these measurements cannot be used to distinguish flight modes in extant birds, and so cannot be used to infer flight mode in fossil forms. Our analyses suggest that more data from fossil birds, combined with accurate functional determination of the flight styles of living forms is required if we are to be able to predict the flight modes of extinct birds. □Birds, flight, morphospace, Mesozoic, wing.     

现生鸟类和孔子鸟(Confuciusornis)叉骨软骨的研究:初步分析以及对中生代鸟类飞行方式的启示（英文）

飞行的早期演化是古生物研究的热点问题之一。现生鸟类演化出多种不同飞行方式,然而对中生代鸟类飞行方式的研究亟需寻找具有指示意义的指标。骨骼与关节组织尤其是乌喙骨-叉骨关节是实现鸟类飞行功能的重要组成,因此其关节的组织形态可能有助于反映现生鸟类的飞行方式。鸟类膜质骨中的次级软骨受到表观遗传的重要影响,只能在关节受到肌肉运动刺激的情况下形成,因此能够反映鸟类关节的形成力学环境;对三种不同飞行方式的现生鸟类(珠颈斑鸠、树麻雀和普通楼燕),以及热河生物群中最为常见的古鸟类之一孔子鸟(Confuciusornis)的叉骨乌喙骨关节的组织学特征进行了分析,显示在所有三种现生鸟类中,叉骨与乌喙骨之间均存在可动关节,并且在叉骨上存在次级软骨。而孔子鸟的叉骨上也存在次级软骨,这是次级软骨组织在中生代鸟类叉骨化石中的首次报道。进一步分析发现,不同现生鸟类物种的次级软骨组织形态存在差异,还需更多数据建立相关形态功能关系以便用于帮助推断中生代鸟类的飞行方式。