Reinterpretation of a previously described Jehol bird clarifies early trophic evolution in the Ornithuromorpha.
ABSTRACT: STM35-3 from the Yixian Formation is the only Early Cretaceous ornithuromorph preserving direct evidence of granivory. The crop contains numerous seeds and the preservation of gastroliths presumably within the ventriculus indicates this diet was paired with the presence of a gastric mill as in living granivorous birds. STM35-3 was originally referred to Hongshanornis longicresta, member of a diverse clade of small, basal ornithuromorphs with elongate hindlimbs known as the Hongshanornithidae. Hindlimb proportions suggest that hongshanornithids were wading birds, an ecological inference somewhat in conflict with direct evidence suggesting Hongshanornis fed on seeds. However, close inspection of STM35-3 reveals that the specimen represents a new species not closely related to hongshanornithids, distinguished by large forelimbs that exceed the length of the hindlimbs, robust and narrow coracoids, and a delicate edentulous rostrum. By contrast, all hongshanornithids have hindlimbs that far exceed the length of the forelimbs, coracoids with wide sternal margins, and small teeth throughout the upper and lower jaws. Reinterpretation of this new taxon, Eogranivora edentulata gen. et sp. nov, helps to clarify trophic driven patterns of tooth loss within the Ornithuromorpha. Apparent loss of the hallux may represent the first such occurrence in a Mesozoic bird and suggests a highly terrestrial lifestyle.
Project description:The midline pattern of sternal ossification characteristic of the Cretaceous enantiornithine birds is unique among the Ornithodira, the group containing birds, nonavian dinosaurs and pterosaurs. This has been suggested to indicate that Enantiornithes is not the sister group of Ornithuromorpha, the clade that includes living birds and their close relatives, which would imply rampant convergence in many nonsternal features between enantiornithines and ornithuromorphs. However, detailed comparisons reveal greater similarity between neornithine (i.e. crown group bird) and enantiornithine modes of sternal ossification than previously recognized. Furthermore, a new subadult enantiornithine specimen demonstrates that sternal ossification followed a more typically ornithodiran pattern in basal members of the clade. This new specimen, referable to the Pengornithidae, indicates that the unique ossification pattern observed in other juvenile enantiornithines is derived within Enantiornithes. A similar but clearly distinct pattern appears to have evolved in parallel in the ornithuromorph lineage. The atypical mode of sternal ossification in some derived enantiornithines should be regarded as an autapomorphic condition rather than an indication that enantiornithines are not close relatives of ornithuromorphs. Based on what is known about molecular mechanisms for morphogenesis and the possible selective advantages, the parallel shifts to midline ossification that took place in derived enantiornithines and living neognathous birds appear to have been related to the development of a large ventral keel, which is only present in ornithuromorphs and enantiornithines. Midline ossification can serve to medially reinforce the sternum at a relatively early ontogenetic stage, which would have been especially beneficial during the protracted development of the superprecocial Cretaceous enantiornithines.
Project description:Here we report on three new species of ornithuromorph birds from the Lower Cretaceous Xiagou Formation in the Changma Basin of Gansu Province, northwestern China: Yumenornis huangi gen. et sp. nov., Changmaornis houi gen. et sp. nov., and Jiuquanornis niui gen. et sp. nov.. The last of these is based on a previously published but unnamed specimen: GSGM-05-CM-021. Although incomplete, the specimens can be clearly distinguished from each other and from Gansus yumenensis Hou and Liu, 1984. Phylogenetic analysis resolves the three new taxa as basal ornithuromorphs. This study reveals previously unrecognized ornithuromorph diversity in the Changma avifauna, which is largely dominated by Gansus but with at least three other ornithuromorphs. Body mass estimates demonstrate that enantiornithines were much smaller than ornithuromorphs in the Changma avifauna. In addition, Changma enantiornithines preserve long and recurved pedal unguals, suggesting an arboreal lifestyle; in contrast, Changma ornithuromorphs tend to show terrestrial or even aquatic adaptions. Similar differences in body mass and ecology are also observed in the Jehol avifauna in northeastern China, suggesting niche partitioning between these two clades developed early in their evolutionary history.
Project description:The Lower Cretaceous Huajiying Formation of the Sichakou Basin in northern Hebei Province, northern China contains key vertebrate taxa of the early Jehol Biota, e.g., Protopteryx fengningensis, Archaeornithura meemannae, Peipiaosteus fengningensis, and Eoconfuciusornis zhengi This formation arguably documents the second-oldest bird-bearing horizon, producing the oldest fossil records of the two major Mesozoic avian groups Enantiornithes and Ornithuromorpha. Hence, precisely determining the depositional ages of the Huajiying Formation would advance our understanding of the evolutionary history of the Jehol Biota. Here we present secondary ion mass spectrometry (SIMS) U-Pb zircon analysis results of eight interbedded tuff/tuffaceous sandstone samples from the Huajiying Formation. Our findings, combined with previous radiometric dates, suggest that the oldest enantiornithine and ornithuromorph birds in the Jehol Biota are ?129-131 Ma, and that the Jehol Biota most likely first appeared at ?135 Ma. This expands the biota's temporal distribution from late Valanginian to middle Aptian with a time span of about 15 My.
Project description:Ornithuromorpha is the most inclusive clade containing extant birds but not the Mesozoic Enantiornithes. The early evolutionary history of this avian clade has been advanced with recent discoveries from Cretaceous deposits, indicating that Ornithuromorpha and Enantiornithes are the two major avian groups in Mesozoic. Here we report on a new ornithuromorph bird, Archaeornithura meemannae gen. et sp. nov., from the second oldest avian-bearing deposits (130.7?Ma) in the world. The new taxon is referable to the Hongshanornithidae and constitutes the oldest record of the Ornithuromorpha. However, A. meemannae shows few primitive features relative to younger hongshanornithids and is deeply nested within the Hongshanornithidae, suggesting that this clade is already well established. The new discovery extends the record of Ornithuromorpha by five to six million years, which in turn pushes back the divergence times of early avian lingeages into the Early Cretaceous.
Project description:Can lower limb motor function be improved after a spinal cord lesion by re-engaging functional activity of the upper limbs? We addressed this issue by training the forelimbs in conjunction with the hindlimbs after a thoracic spinal cord hemisection in adult rats. The spinal circuitries were more excitable, and behavioural and electrophysiological analyses showed improved hindlimb function when the forelimbs were engaged simultaneously with the hindlimbs during treadmill step-training as opposed to training only the hindlimbs. Neuronal retrograde labelling demonstrated a greater number of propriospinal labelled neurons above and below the thoracic lesion site in quadrupedally versus bipedally trained rats. The results provide strong evidence that actively engaging the forelimbs improves hindlimb function and that one likely mechanism underlying these effects is the reorganization and re-engagement of rostrocaudal spinal interneuronal networks. For the first time, we provide evidence that the spinal interneuronal networks linking the forelimbs and hindlimbs are amenable to a rehabilitation training paradigm. Identification of this phenomenon provides a strong rationale for proceeding toward preclinical studies for determining whether training paradigms involving upper arm training in concert with lower extremity training can enhance locomotor recovery after neurological damage.
Project description:Background:Kinetic motion analysis has been used in canines and equines as a fundamental objective evaluation measurement. Cats are very capable jumpers, and this ability has biomimetic applications. It is essential to understand movement patterns and physical adaptations of this species, as cats are popular pets for humans. Further to this, motion analysis of a cat's movement patterns may provide potentially valuable information in relation to limb disease and injury. Therefore, the aim of this study was to investigate kinetic differences in cats when landing from varying preselected controlled heights. Methods:The peak vertical force (PVF) and paw contact area (CA) of both the forelimbs and hindlimbs were collected from seven healthy Chinese domesticated cats while landing from heights of 30 cm, 50 cm, 70 cm and 90 cm respectively. The falling motivation for the cats was facilitated with the use of a flip board. This device provided the basis for the cats to land passively. Results:The results indicated that the PVF of all examined limbs (fore right, fore left, hind right, hind left) significantly increased as the height increased. When the PVF from the hindlimbs and forelimbs were compared, the forelimbs recorded significantly greater values for all heights examined (P < 0.001). The PVF of the hindlimbs was symmetrical at all heights, but forelimb symmetry only occurred at the lower heights. The hindlimbs demonstrated larger CA than the forelimbs measured from all heights on landing (P < 0.001). Moreover, the paw CA on the left and right limbs were symmetrical. Discussion:The paw CA of cats may be an effective parameter to evaluate abnormalities or diseases in the limbs of cats. Additionally, these findings highlight how cats land from varying heights, which may also provide reference values for the bionic design of artificial limbs for felines and treatment for limb diseases in this species.
Project description:Some rats spinalized P1/P2 achieve autonomous weight-supported locomotion and quiet stance as adults. We used force platforms and robot-applied perturbations to test such spinalized rats (n = 6) that exhibited both weight-supporting locomotion and stance, and also normal rats (n = 8). Ground reaction forces in individual limbs and the animals' center of pressure were examined. In normal rats, both forelimbs and hindlimbs participated actively to control horizontal components of ground reaction forces. Rostral perturbations increased forelimb ground reaction forces and caudal perturbations increased hindlimb ground reaction forces. Operate rats carried 60% body weight on the forelimbs and had a more rostral center of pressure placement. The pattern in normal rats was to carry significantly more weight on the hindlimbs in quiet stance (roughly 60%). The strategy of operate rats to compensate for perturbations was entirely in forelimbs; as a result, the hindlimbs were largely isolated from the perturbation. Stiffness magnitude of the whole body was measured: its magnitude was hourglass shaped, with the principal axis oriented rostrocaudally. Operate rats were significantly less stiff--only 60-75% of normal rats' stiffness. The injured rats adopt a stance strategy that isolates the hindlimbs from perturbation and may thus prevent hindlimb loadings. Such loadings could initiate reflex stepping, which we observed. This might activate lumbar pattern generators used in their locomotion. Adult spinalized rats never achieve independent hindlimb weight-supported stance. The stance strategy of the P1 spinalized rats differed strongly from the behavior of intact rats and may be difficult for rats spinalized as adults to master.
Project description:The rich repertoire of locomotor behaviors in quadrupedal animals requires flexible inter-limb and inter-segmental coordination. Here we studied the kinematic coordination of different gaits (walk, trot, gallop, and swim) of six dogs (Canis lupus familiaris) and, in particular, the planar covariation of limb segment elevation angles. The results showed significant variations in the relative duration of rearward limb movement, amplitude of angular motion, and inter-limb coordination, with gait patterns ranging from a lateral sequence of footfalls during walking to a diagonal sequence in swimming. Despite these differences, the planar law of inter-segmental coordination was maintained across different gaits in both forelimbs and hindlimbs. Notably, phase relationships and orientation of the covariation plane were highly limb specific, consistent with the functional differences in their neural control. Factor analysis of published muscle activity data also demonstrated differences in the characteristic timing of basic activation patterns of the forelimbs and hindlimbs. Overall, the results demonstrate that the planar covariation of inter-segmental coordination has emerged for both fore- and hindlimbs and all gaits, although in a limb-specific manner.
Project description:Limb skeletal elements originate from the limb progenitor cells, which undergo expansion and patterning to develop each skeletal element. Posterior-distal skeletal elements, such as the ulna/fibula and posterior digits develop in a Sonic hedgehog (Shh)-dependent manner. However, it is poorly understood how anterior-proximal elements, such as the humerus/femur, the radius/tibia and the anterior digits, are developed. Here we show that the zinc finger factors Sall4 and Gli3 cooperate for proper development of the anterior-proximal skeletal elements and also function upstream of Shh-dependent posterior skeletal element development. Conditional inactivation of Sall4 in the mesoderm before limb outgrowth caused severe defects in the anterior-proximal skeletal elements in the hindlimb. We found that Gli3 expression is reduced in Sall4 mutant hindlimbs, but not in forelimbs. This reduction caused posteriorization of nascent hindlimb buds, which is correlated with a loss of anterior digits. In proximal development, Sall4 integrates Gli3 and the Plzf-Hox system, in addition to proliferative expansion of cells in the mesenchymal core of nascent hindlimb buds. Whereas forelimbs developed normally in Sall4 mutants, further genetic analysis identified that the Sall4-Gli3 system is a common regulator of the early limb progenitor cells in both forelimbs and hindlimbs. The Sall4-Gli3 system also functions upstream of the Shh-expressing ZPA and the Fgf8-expressing AER in fore- and hindlimbs. Therefore, our study identified a critical role of the Sall4-Gli3 system at the early steps of limb development for proper development of the appendicular skeletal elements.
Project description:Scincine lizards in Madagascar form an endemic clade of about 60 species exhibiting a variety of ecomorphological adaptations. Several subclades have adapted to burrowing and convergently regressed their limbs and eyes, resulting in a variety of partial and completely limbless morphologies among extant taxa. However, patterns of limb regression in these taxa have not been studied in detail. Here we fill this gap in knowledge by providing a phylogenetic analysis of DNA sequences of three mitochondrial and four nuclear gene fragments in an extended sampling of Malagasy skinks, and microtomographic analyses of osteology of various burrowing taxa adapted to sand substrate. Based on our data we propose to (i) consider Sirenoscincus Sakata & Hikida, 2003, as junior synonym of Voeltzkowia Boettger, 1893; (ii) resurrect the genus name Grandidierina Mocquard, 1894, for four species previously included in Voeltzkowia; and (iii) consider Androngo Brygoo, 1982, as junior synonym of Pygomeles Grandidier, 1867. By supporting the clade consisting of the limbless Voeltzkowia mira and the forelimb-only taxa V. mobydick and V. yamagishii, our data indicate that full regression of limbs and eyes occurred in parallel twice in the genus Voeltzkowia (as hitherto defined) that we consider as a sand-swimming ecomorph: in the Voeltzkowia clade sensu stricto the regression first affected the hindlimbs and subsequently the forelimbs, whereas the Grandidierina clade first regressed the forelimbs and subsequently the hindlimbs following the pattern prevalent in squamates. Timetree reconstructions for the Malagasy Scincidae contain a substantial amount of uncertainty due to the absence of suitable primary fossil calibrations. However, our preliminary reconstructions suggest rapid limb regression in Malagasy scincids with an estimated maximal duration of 6 MYr for a complete regression in Paracontias, and 4 and 8 MYr respectively for complete regression of forelimbs in Grandidierina and hindlimbs in Voeltzkowia.