Gareth Dyke

Sustained miniaturization and anatomical innovation in the dinosaurian ancestors of birds

Turning large dinosaurs into small birds Most paleontologists agree that birds are descended from dinosaurs. How did such large terrestrial or aquatic animals evolve into small feathered fliers? Lee et al. used two large databases of theropod morphology to explore possible evolutionary patterns that may have driven this dramatic transformation (see the Perspective by Benton). They found no clear pattern of miniaturization across the entire clade of Theropoda. However, several lines of evidence suggested that the lineage leading to birds underwent sustained miniaturization. Within that lineage, body sizes decreased and species evolved faster. They also developed ecological and morphological innovations linked to smaller body sizes. Science, this issue p. 562; see also p. 508 Birds evolved from their dinosaurian ancestors through a period characterized by sustained and repeated decreases in size. [Also see Perspective by Benton] Recent discoveries have highlighted the dramatic evolutionary transformation of massive, ground-dwelling theropod dinosaurs into light, volant birds. Here, we apply Bayesian approaches (originally developed for inferring geographic spread and rates of molecular evolution in viruses) in a different context: to infer size changes and rates of anatomical innovation (across up to 1549 skeletal characters) in fossils. These approaches identify two drivers underlying the dinosaur-bird transition. The theropod lineage directly ancestral to birds undergoes sustained miniaturization across 50 million years and at least 12 consecutive branches (internodes) and evolves skeletal adaptations four times faster than other dinosaurs. The distinct, prolonged phase of miniaturization along the bird stem would have facilitated the evolution of many novelties associated with small body size, such as reorientation of body mass, increased aerial ability, and paedomorphic skulls with reduced snouts but enlarged eyes and brains.

Morphological Clocks in Paleontology, and a Mid-Cretaceous Origin of Crown Aves

Birds are among the most diverse and intensivelystudied vertebrate groups, but many aspects of theirhigher-level phylogeny and evolution still remaincontroversial. One contentious issue concerns theantiquity of modern birds (=crown Aves): the ageof the most recent common ancestor of all livingbirds (Gauthier 1986). Very few Mesozoic fossilsare attributable to modern birds (e.g., Clarke et al.2005; Dyke and Kaiser 2011; Brocklehurst et al. 2012;Ksepka and Boyd 2012) suggesting that they diversifiedlargely or entirely in the early Paleogene, perhaps in theecologicalvacuumcreatedbytheextinctionofnon-aviandinosaurs, pterosaurs, and many archaic (stem) birds(e.g.,Longrichetal.2011).Incontrast,molecularstudiesindicate that modern birds commenced radiating deepwithin the Mesozoic, for example ∼130Ma(Cooperand Penny 1997; Haddrath and Baker 2012)or∼113 Ma(Jetz et al. 2012), with ratites, galliforms, anseriforms,shorebirds, and even passerines surviving acrossthe KPg boundary (∼66 Ma). The oldest moleculardates further imply an extraordinarily rapid earlybird evolution, with the modern birds appearingonly 20 myr after

Reduced plumage and flight ability of a new Jurassic paravian theropod from China.

Feathered theropods were diverse in the Early Cretaceous Jehol Group of western Liaoning Province, China. Recently, anatomically distinct feathered taxa have been discovered in the older Middle-Late Jurassic Tiaojishan Formation in the same region. Phylogenetic hypotheses including these specimens have challenged the pivotal position of Archaeopteryx in bird phylogeny. Here we report a basal troodontid from the Tiaojishan Formation that resembles Anchiornis, also from Jianchang County (regarded as sister-taxa). The feathers of Eosinopteryx are less extensive on the limbs and tail than Anchiornis and other deinonychosaurians. With reduced plumage and short uncurved pedal claws, Eosinopteryx would have been able to run unimpeded (with large foot remiges cursorial locomotion was likely problematic for Anchiornis). Eosinopteryx increases the known diversity of small-bodied dinosaurs in the Jurassic, shows that taxa with similar body plans could occupy different niches in the same ecosystem and suggests a more complex picture for the origin of flight.

A reversal of fortunes: climate change ‘winners’ and ‘losers’ in Antarctic Peninsula penguins

Climate change is a major threat to global biodiversity. Antarctic ecosystems are no exception. Investigating past species responses to climatic events can distinguish natural from anthropogenic impacts. Climate change produces ‘winners’, species that benefit from these events and ‘losers’, species that decline or become extinct. Using molecular techniques, we assess the demographic history and population structure of Pygoscelis penguins in the Scotia Arc related to climate warming after the Last Glacial Maximum (LGM). All three pygoscelid penguins responded positively to post-LGM warming by expanding from glacial refugia, with those breeding at higher latitudes expanding most. Northern (Pygoscelis papua papua) and Southern (Pygoscelis papua ellsworthii) gentoo sub-species likely diverged during the LGM. Comparing historical responses with the literature on current trends, we see Southern gentoo penguins are responding to current warming as they did during post-LGM warming, expanding their range southwards. Conversely, Adélie and chinstrap penguins are experiencing a ‘reversal of fortunes’ as they are now declining in the Antarctic Peninsula, the opposite of their response to post-LGM warming. This suggests current climate warming has decoupled historic population responses in the Antarctic Peninsula, favoring generalist gentoo penguins as climate change ‘winners’, while Adélie and chinstrap penguins have become climate change ‘losers’.

Aerodynamic performance of the feathered dinosaur Microraptor and the evolution of feathered flight

Understanding the aerodynamic performance of feathered, non-avialan dinosaurs is critical to reconstructing the evolution of bird flight. Here we show that the Early Cretaceous five-winged paravian Microraptor is most stable when gliding at high-lift coefficients (low lift/drag ratios). Wind tunnel experiments and flight simulations show that sustaining a high-lift coefficient at the expense of high drag would have been the most efficient strategy for Microraptor when gliding from, and between, low elevations. Analyses also demonstrate that anatomically plausible changes in wing configuration and leg position would have made little difference to aerodynamic performance. Significant to the evolution of flight, we show that Microraptor did not require a sophisticated, modern wing morphology to undertake effective glides. This is congruent with the fossil record and also with the hypothesis that symmetric flight feathers first evolved in dinosaurs for non-aerodynamic functions, later being adapted to form lifting surfaces.

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 Sebeş area of Transylvania, Romania. A lens of calcareous mudstone with minimum dimensions of 80xa0cm length, 50xa0cm width and 20xa0cm depth contains thousands of tightly packed, morphologically homogenous eggshell fragments, seven near-complete eggs and neonatal and adult avialan skeletal elements. Eggshell forms 70–80xa0% 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.

Limb disparity and wing shape in pterosaurs

The limb proportions of the extinct flying pterosaurs were clearly distinct from their living counterparts, birds and bats. Within pterosaurs, however, we show that further differences in limb proportions exist between the two main groups: the clade of short‐tailed Pterodactyloidea and the paraphyletic clades of long‐tailed rhamphorhynchoids. The hindlimb to forelimb ratios of rhamphorhynchoid pterosaurs are similar to that seen in bats, whereas those of pterodactyloids are much higher. Such a clear difference in limb ratios indicates that the extent of the wing membrane in rhamphorhynchoids and pterodactyloids may also have differed; this is borne out by simple ternary analyses. Further, analyses also indicate that the limbs of Sordes pilosus, a well‐preserved small taxon used as key evidence for inferring the extent and shape of the wing membrane in all pterosaurs, are not typical even of its closest relatives, other rhamphorhynchoids. Thus, a bat‐like extensive hindlimb flight membrane, integrated with the feet and tail may be applicable only to a small subset of pterosaur diversity. The range of flight morphologies seen in these extinct reptiles may prove much broader than previously thought.

A New Azhdarchid Pterosaur from the Late Cretaceous of the Transylvanian Basin, Romania: Implications for Azhdarchid Diversity and Distribution

We describe a new taxon of medium-sized (wing span ca. 3 m) azhdarchid pterosaur from the Upper Cretaceous Transylvanian Basin (Sebeş Formation) of Romania. This specimen is the most complete European azhdarchid yet reported, comprising a partially articulated series of vertebrae and associated forelimb bones. The new taxon is most similar to the Central Asian Azhdarcho lancicollis Nessov but possesses a suite of autapomorphies in its vertebrae that include the relative proportions of cervicals three and four and the presence of elongated prezygapophyseal pedicles. The new taxon is interesting in that it lived contemporaneously with gigantic forms, comparable in size to the famous Romanian Hatzegopteryx thambema. The presence of two distinct azhdarchid size classes in a continental depositional environment further strengthens suggestions that these pterosaurs were strongly linked to terrestrial floodplain and wooded environments. To support this discussion, we outline the geological context and taphonomy of our new specimen and place it in context with other known records for this widespread and important Late Cretaceous pterosaurian lineage.

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.

A New Small-Bodied Azhdarchoid Pterosaur from the Lower Cretaceous of England and Its Implications for Pterosaur Anatomy, Diversity and Phylogeny

Background Pterosaurs have been known from the Cretaceous sediments of the Isle of Wight (southern England, United Kingdom) since 1870. We describe the three-dimensional pelvic girdle and associated vertebrae of a small near-adult pterodactyloid from the Atherfield Clay Formation (lower Aptian, Lower Cretaceous). Despite acknowledged variation in the pterosaur pelvis, previous studies have not adequately sampled or incorporated pelvic characters into phylogenetic analyses. Methodology/Principal Findings The new specimen represents the new taxon Vectidraco daisymorrisae gen. et sp. nov., diagnosed by the presence of a concavity posterodorsal to the acetabulum and the form of its postacetabular process on the ilium. Several characters suggest that Vectidraco belongs to Azhdarchoidea. We constructed a pelvis-only phylogenetic analysis to test whether the pterosaur pelvis carries a useful phylogenetic signal. Resolution in recovered trees was poor, but they approximately matched trees recovered from analyses of total evidence. We also added Vectidraco and our pelvic characters to an existing total-evidence matrix for pterosaurs. Both analyses recovered Vectidraco within Azhdarchoidea. Conclusions/Significance The Lower Cretaceous strata of western Europe have yielded members of several pterosaur lineages, but Aptian pterosaurs from western Europe are rare. With a pelvis length of 40 mm, the new animal would have had a total length of c. 350 mm, and a wingspan of c. 750 mm. Barremian and Aptian pterodactyloids from western Europe show that small-bodied azhdarchoids lived alongside ornithocheirids and istiodactylids. This assemblage is similar in terms of which lineages are represented to the coeval beds of Liaoning, China; however, the number of species and specimens present at Liaoning is much higher. While the general phylogenetic composition of western European and Chinese communities appear to have been approximately similar, the differences may be due to different palaeoenvironmental and depositional settings. The western Europe pterodactyloid record may therefore be artificially low in diversity due to preservational factors.