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BackgroundOsteohistological examinations of fossil vertebrates have utilized a number of proxies, such as counts and spacing of lines of arrested growth (LAGs) and osteocyte lacunar densities (OLD), in order to make inferences related to skeletochronology and mass-specific growth rates. However, many of these studies rely on samplings of isolated bones from single individuals. These analyses do not take individual variation into account, and as a result may lead to misleading inferences of the physiology of extinct organisms. This study uses a multi-element, multi-individual sampling of ornithomimid dinosaurs to test the amount of individual variation in the aforementioned osteohistological indicators. Based on these results we also assess the conclusions of previous studies that tested paleohistological hypotheses using isolated elements.ResultsLAG number was found to be consistent within the hind limb bones of each individual, with the exception of the fibula, which preserves one additional LAG. Considerable differences in LAG spacing were found between elements of the sampled individuals, with larger variation found in elements of the foot compared with the femur, fibula, and tibia. Osteocyte lacunar density ranged between 29000 and 42000 osteocyte lacunae per mm3, and was found to vary more between hind limb bones of an individual and within bones, than between the average values of individuals.ConclusionsThe variation between hind limb elements in LAG number and LAG spacing suggests that direct comparisons of these elements may be misleading, and that LAG spacing is not a reliable proxy for mass-specific growth rates of an individual. Sampling of multiple bones should be performed as an internal check of model-based LAG retro-calculation and growth equations. The observation that osteocyte lacunar density varies more between individual bone elements than between average individual values suggests that the choice of sampled element can greatly influence the result, and care should be taken to not bias interpretations of the physiology of fossil tetrapods.Electronic supplementary materialThe online version of this article (doi:10.1186/s12862-014-0231-y) contains supplementary material, which is available to authorized users.

Electro-optic phase modulators are critical components in modern communication, microwave photonic, and quantum photonic systems. Important for these applications is to achieve modulators with low half-wave voltage at high frequencies.Here we demonstrate an integrated phase modulator, based on a thin-film lithium niobate platform, that simultaneously features small on-chip loss (∼ 1 dB) and low half-wave voltage over a large spectral range (3.5 -4.5 V at 5 -40 GHz). By driving the modulator with a strong 30-GHz microwave signal corresponding to around four half-wave voltages, we generate an optical frequency comb consisting of over 40 sidebands spanning 10 nm in the telecom L-band. The high electro-optic performance combined with the high RF power-handling ability (3.1 W) of our integrated phase modulator are crucial for future photonics and microwave systems.

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The independent evolution of gigantism among dinosaurs has been a topic of long-standing interest, but it remains unclear if gigantic theropods, the largest bipeds in the fossil record, all achieved massive sizes in the same manner, or through different strategies. We perform multi-element histological analyses on a phylogenetically broad dataset sampled from eight theropod families, with a focus on gigantic tyrannosaurids and carcharodontosaurids, to reconstruct the growth strategies of these lineages and test if particular bones consistently preserve the most complete growth record. We find that in skeletally mature gigantic theropods, weight-bearing bones consistently preserve extensive growth records, whereas non-weight-bearing bones are remodelled and less useful for growth reconstruction, contrary to the pattern observed in smaller theropods and some other dinosaur clades. We find a heterochronic pattern of growth fitting an acceleration model in tyrannosaurids, with allosauroid carcharodontosaurids better fitting a model of hypermorphosis. These divergent growth patterns appear phylogenetically constrained, representing extreme versions of the growth patterns present in smaller coelurosaurs and allosauroids, respectively. This provides the first evidence of a lack of strong mechanistic or physiological constraints on size evolution in the largest bipeds in the fossil record and evidence of one of the longest-living individual dinosaurs ever documented.

Bonebeds can provide a wealth of anatomical, taphonomic, and ontogenetic information about the specimens preserved within them, and can provide evidence for inferred behavior. The material described here represents the first known bonebed of ornithomimids in North America, and the fourth record of an ornithomimosaur bonebed in the world. Partial skeletons representing three individuals are preserved in this assemblage, each comprising primarily portions of the posterior postcrania (pelvis, hind limbs and tail). All three individuals are morphologically similar, although one is larger in overall size. Given the stratigraphic position of the site, and the morphology of the postcrania, the preserved material represents a taxon from the clade containing Ornithomimus and Struthiomimus. Pedal ungual morphology is examined and found to be too variable to be useful in distinguishing these species taxonomically. This site provides additional evidence of gregarious behavior in ornithomimids and the first probable record of that behavior in North American forms.

BackgroundThe Belly River Group of southern Alberta is one of the best-sampled Late Cretaceous terrestrial faunal assemblages in the world. This system provides a high-resolution biostratigraphic record of terrestrial vertebrate diversity and faunal turnover, and it has considerable potential to be a model system for testing hypotheses of dinosaur palaeoecological dynamics, including important aspects of palaeoecommunity structure, trophic interactions, and responses to environmental change. Vertebrate fossil microsites (assemblages of small bones and teeth concentrated together over a relatively short time and thought to be representative of community composition) offer an unparalleled dataset to better test these hypotheses by ameliorating problems of sample size, geography, and chronostratigraphic control that hamper other palaeoecological analyses. Here, we assembled a comprehensive relative abundance dataset of microsites sampled from the entire Belly River Group and performed a series of analyses to test the influence of environmental factors on site and taxon clustering, and assess the stability of faunal assemblages both temporally and spatially. We also test the long-held idea that populations of large dinosaur taxa were particularly sensitive to small-scale environmental gradients, such as the paralic (coastal) to alluvial (inland) regimes present within the time-equivalent depositional basin of the upper Oldman and lower Dinosaur Park Formations.ResultsPalaeoenvironment (i.e. reconstructed environmental conditions, related to relative amount of alluvial, fluvial, and coastal influence in associated sedimentary strata) was found to be strongly associated with clustering of sites by relative-abundance faunal assemblages, particularly in relation to changes in faunal assemblage composition and marine-terrestrial environmental transitions. Palaeogeography/palaeolandscape were moderately associated to site relative abundance assemblage clustering, with depositional setting and time (i.e. vertical position within stratigraphic unit) more weakly associated. Interestingly, while vertebrate relative abundance assemblages as a whole were strongly correlated with these marine-terrestrial transitions, the dinosaur fauna does not appear to be particularly sensitive to them.ConclusionsThis analysis confirms that depositional setting (i.e. the sediment type/sorting and associated characteristics) has little effect on faunal assemblage composition, in contrast to the effect of changes in the broader palaeoenvironment (e.g. upper vs. lower coastal plain, etc.), with marine-terrestrial transitions driving temporal faunal dynamics within the Belly River Group. The similarity of the dinosaur faunal assemblages between the time-equivalent portions of the Dinosaur Park Formation and Oldman Formation suggests that either these palaeoenvironments are more similar than characterized in the literature, or that the dinosaurs are less sensitive to variation in palaeoenvironment than has often been suggested. A lack of sensitivity...

Sexual selection is one of the earliest areas of interest in evolutionary biology. And yet, the evolutionary history of sexually dimorphic traits remains poorly characterized for most vertebrate lineages. Here, we report on evidence for the early evolution of dimorphism within a model mammal group, the pinnipeds. Pinnipeds show a range of sexual dimorphism and mating systems that span the extremes of modern mammals, from monomorphic taxa with isolated and dispersed mating to extreme size dimorphism with highly ordered polygynous harem systems. In addition, the degree of dimorphism in pinnipeds is closely tied to mating system, with strongly dimorphic taxa always exhibiting a polygynous system, and more monomorphic taxa possessing weakly polygynous systems. We perform a comparative morphological description, and provide evidence of extreme sexual dimorphism (similar to sea lions), in the Miocene-aged basal pinniped taxon Enaliarctos emlongi. Using a geometric morphometric approach and combining both modern and fossil taxa we show a close correlation between mating system and sex-related cranial dimorphism, and also reconstruct the ancestral mating system of extant pinnipeds as highly polygynous. The results suggest that sexual dimorphism and extreme polygyny in pinnipeds arose by 27 Ma, in association with changing climatic conditions.