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Conceived and designed the experiments: Therizinosauria is one of the most enigmatic and peculiar clades among theropod dinosaurs, exhibiting an unusual suite of characters, such as lanceolate teeth, a rostral rhamphotheca, long manual claws, and a wide, opisthopubic pelvis. This specialized anatomy has been associated with a shift in dietary preferences and an adaptation to herbivory. Despite a large number of just click for source in recent years, the fossil record for Therizinosauria is still relatively poor, and cranial remains are particularly rare.

Based on computed tomographic CT scanning of the nearly complete and articulated skull of Erlikosaurus andrewsias well as partial braincases of two other therizinosaurian taxa, the endocranial anatomy is reconstructed and described.

The wider phylogenetic range of the described specimens permits the evaluation of sensory and cognitive capabilities of Therizinosauria in an evolutionary context. The endocranial anatomy reveals a mosaic of plesiomorphic and derived characters in therizinosaurians. The anatomy of the olfactory apparatus and the endosseous labyrinth suggests that olfaction, hearing, and equilibrium were well-developed in therizinosaurians and might have affected or benefited from an enlarged telencephalon.

This study presents the first appraisal of the evolution of endocranial anatomy and sensory adaptations in Therizinosauria.

Despite their phylogenetically basal position among maniraptoran dinosaurs, therizinosaurians had developed the neural pathways for a well developed sensory repertoire. Therizinosauria comprises an enigmatic clade of Cretaceous Barremian — Maastrichtian maniraptoran dinosaurs found in Asia and North American [1].

They are distinguished from other theropods by an unusual suite of morphological characters, which include lanceolate, tightly packed teeth, a rostral rhamphotheca, an elongate neck, enlarged manual claws, and a wide, opisthopubic pelvis. Many of these characters have been associated with an herbivorous diet and specializations in foraging behavior [2].

Recent phylogenetic analyses [3][4] have recovered Therizinosauria close to the base of Maniraptora. Nested between basal coelurosaurs and derived maniraptorans, this makes them of special importance not only in terms of the evolution of anatomical and developmental patterns, but also for the radiation of Athens Georgia Dating Free Artwork Of Rhinosinusitis Vs Sinusitis among theropods [5].

Their unique morphology has resulted in a plethora of hypotheses Athens Georgia Dating Free Artwork Of Rhinosinusitis Vs Sinusitis the evolution, dietary preferences, and behavior of therizinosaurians — most of which have been inferred, based on skeletal traits and anatomical characters [6] — [9].

However, the endocranial http://malishka.info/lit/why-do-guys-have-big-balls.php has rarely been considered, and little is known about the sensory and cognitive abilities in therizinosaurians and whether any of these reflect adaptations to an herbivorous lifestyle.

As the neurocranium and the brain are closely associated during growth, the endocranial cavity constitutes a representation of the gross surficial structure of the brain itself. A cast of the endocranial cavity, thus can provide the means to reconstruct the brain anatomy and associated structures [10][11].

The neural centers for the senses of vision, olfaction, hearing and balance either reside in the brain or peripherally in the inner ear, allowing us to evaluate the relative sensory capabilities using a series of qualitative and quantitative measures.

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A variety of cranial endocasts both digital and physical of dinosaurs in general have been described in recent years [12] — [15]. For theropods, these include mainly basal and non-maniraptoriform taxa [16] — [23]probably in part because braincase elements are more likely to be preserved in larger animals.

In contrast, data on brain anatomy and endocasts are rare in Maniraptoriformes and restricted to some scattered specimens among the individual lineages [24] — [27]. Given the paucity of available material, it may not be surprising that only a handful of studies focus on a broader evolutionary context of brain anatomy and sensory adaptations [10][14][21][28][29][30].

Although cranial remains are still rare, numerous discoveries have expanded our knowledge of therizinosaurians in the last 20 years. Based on the endocasts of these four specimens, we present an extensive description of the endocranial anatomy of therizinosaurians Figure 1.

The wide phylogenetic range of these samples, spanning both basal and derived taxa and thus a considerable part of the known therizinosaurian diversity, allows inferences to be drawn about sensory and cognitive capabilities.

It further provides the foundation for the study of endocranial and sensorineural evolution in Therizinosauria. This is potentially relevant to the discussion of whether therizinosaurian sensory and cognitive abilities were specially adapted to herbivory.

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The resulting rotational projections were processed with custom-built software provided by X-Tek Systems Ltd. The respective scan data are part of an ongoing doctoral dissertation of the first author and will be deposited in an electronic archive and with the respective museum collections after completion of the dissertation.

The final data files were subsequently imported into Avizo Version 6. Anatomical structures of interest endocasts, endosseous labyrinths, cranial nerves, and blood vessels were highlighted and separately labeled using Avizo's segmentation editor.

Additionally, all measurements presented in this article and used for various calculations were derived from the digital models using Avizo see Table 1. A 3d pdf document with interactive figures can be found in supplementary information. Due to some extensive click to the caudal left braincase wall basisphenoid, laterosphenoid, prootic, exoccipital of Erlikosaurusit was not possible to completely reconstruct this part of the endocast.

To obtain a complete endocast, the individual slices of the data set were mirrored and the missing part of the endocast was created and registered for the left side separately. To automate these steps two tcl tool check this out language scripts were created for Avizo.

Their unique morphology has resulted in a plethora of hypotheses on the evolution, dietary preferences, and behavior of therizinosaurians — most of which have been inferred, based on skeletal traits and anatomical characters [6] — [9]. Based on the findings for Tyrannosaurus rex Lawrence County and Northern New York. Alexander RM All-time giants: Orange Regional Medical Center.

The endocast description is therefore mostly based on the original right side, unless specified otherwise. This method was also applied to acquire and supplement data for some parts of the endosseous labyrinths of Falcarius and Nothronychus.

Athens Georgia Dating Free Artwork Of Rhinosinusitis Vs Sinusitis

Following Sampson and Witmer [20]we will use the term cranial endocast in reference to the space within the braincase, housing the actual brain, the meninges and dural venous sinuses, the cerebral blood vessels, and the cranial nerve roots. To facilitate the discussion of these elements, we will further refer to the digital cast of the structures as if they were the structures themselves. We focus our description on Erlikosaurus andrewsias its endocast is the most complete and shows the majority of anatomical features.

The phylogenetic and taxonomic classifications follow Zanno [4]by using the definitions for Therizinosauria and correspondingly therizinosaurian, to describe a clade including the basal taxon Falcarius utahensiswhereas the terms Therizinosauroidea and therizinosauroid, and Therizinosauridae and therizinosaurid define more exclusive clades within Therizinosauria.

No Athens Georgia Dating Free Artwork Of Rhinosinusitis Vs Sinusitis were required for the described study, which complied with all relevant regulations. Relative and correlated measures of sensory capabilities are typically scaled to body mass. In the case of therizinosaurians, the general lack of preserved fossil remains usable for go here calculations poses a considerable problem. Moreover, the exceptional body morphology and proportions, which differ from those of other theropods, make comparisons to better-known theropod groups difficult.

Of the three therizinosaurian taxa discussed here, the only published mass estimate was given by Russell and Dong [37] for Erlikosaurus andrewsiwho estimated its body mass at approximately kg.

This value was obtained by calculating a mass estimate of kg for the more complete Alxasaurus elesitaiensis using Anderson et al. Here we use theropod-specific equations relating body mass to femoral length FL [40] to produce mass estimates for ErlikosaurusNothronychus mckinleyiand Falcarius Table 2. The largest femur from the holotype locality of Falcarius utahensis UMNH measures mm, yielding an estimated mass of kg.

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However, femora are unknown for Erlikosaurus and Nothronychus mckinleyi. In order to estimate femoral length for these taxa, we performed bivariate regression analyses on log-transformed data to describe the relationship between femoral length FL and humeral Things To Do In Eddyville Ky HL in six therizinosauriod taxa Nothronychus graffamiAlxasaurus elesitaiensisErliansaurus bellamanusNeimongosaurus yangiand two undescribed taxa, Y.

Although not a weight-bearing element, we chose the humerus because, among the fragmentary materials known for Erlikosaurus [41]it has the largest representation across Therizinosauria. Despite being small in number, the sampled taxa bracket the humeral lengths of Erlikosaurus and Nothronychus mckinleyi allowing interpolated prediction of FL and minimizing potential error. Applying the maximum humeral length of mm for Erlikosaurus given by Perle [41]yields an estimated FL of A humeral length of mm for Nothronychus mckinleyiresults in a FL of mm and a mass value of kg.

Given the large uncertainty inherent in our approach and in body-mass estimates in general, we assume a range of — kg for Erlikosaurus and — for Nothronychus mckinleyi see also Table 2. The cranial endocasts of all four examined specimens are very similar in their general morphology, at least as far as can be determined from those parts that are preserved.

The full extent of the endocast could only be reconstructed for Erlikosaurus andrewsi. Its endocast is relatively elongate and straight.

More info individual parts of the brain fore- mid- and hindbrain are arranged in the same horizontal plane and do not show Athens Georgia Dating Free Artwork Of Rhinosinusitis Vs Sinusitis obvious angulation, indicating only slight cephalic and pontine flexures. This is in contrast to more basal theropods, where the angulation of the midbrain can be up to 45—60 degrees [20] — [22]but also in contrast to the endocast of the holotype of Falcarius utahensis UMNH VP Although incomplete, its morphology suggests a more pronounced angulation between the hindbrain and the midbrain.

The excavation of the braincase floor and the corresponding angulation in the endocast are less pronounced in the referred specimen UMNH VP of Falcarius.

Nothronychus shows a more moderate flexure similar to Erlikosaurus Figure 2B. However, as these reconstructions are based on partial braincases only, such morphological differences should be considered with caution. In Erlikosaurus Figure 3A—Cthe forebrain region—most notably the olfactory apparatus and the cerebral hemispheres—is well differentiated and clearly demarcated.

The olfactory tracts and bulbs are preserved only in Erlikosaurus. However, due to the lack of preserved or ossified sphenethmoid and mesethmoid elements [42] compare Figure 4the ventral extent of the olfactory apparatus is uncertain and has been reconstructed only tentatively as far as possible, tracing the ventral margins of frontal bones. The olfactory tracts are elongate and extend relatively far from the cerebral hemispheres and the actual brain. They are slightly downturned, following the shape of the skull roof.

The olfactory bulbs are only moderately indicated by shallow impressions on the ventral surface of the frontals. They are of moderate size, as in other Maniraptoriformes Struthiomimus altus Athens Georgia Dating Free Artwork Of Rhinosinusitis Vs Sinusitis, Deinonychus antirrhopusfor which endocranial casts are available [21]. In Falcariustwo incomplete frontals UMNH VP; [43] show preserved, although only partial, impressions of olfactory tracts.

Olfactory bulb size and tract length cannot be ascertained, but a considerable expansion of the olfactory apparatus is evident. The cerebral hemispheres in Erlikosaurus are relatively large and are broader transversely than the caudal part of the endocast. Their contours are clearly reproduced by the endocranial cavity for the entire hemispheres, indicating a close association between the brain and the endocranial wall and the presence of only a thin dural envelope between the two.

Complex vascular grooves on the endocranial surface, typically found in birds and mammals [44]but also in some ornithischians [45]tyrannosaurs [21]respective figure 7oviraptorids [46] and troodontids [47]are not observed in Erlikosaurus. A pair of semi-articulated frontals referred to Falcarius utahensis UMNH VP; [43] shows pronounced endocranial impressions. Although only incompletely preserved, their size indicates equally enlarged cerebral hemispheres as in Erlikosaurus.

In the latter, the cerebral hemispheres are separated dorsally by an interhemispherical fissure Figure 3C. Optic lobes or tecta are not visible in the endocast of Erlikosaurus. The enlarged cerebral hemispheres would suggest that the optic lobes are situated in a ventrolateral position caudal to the cerebral hemispheres and rostral to the transverse sinus [21]. However, Athens Georgia Dating Free Artwork Of Rhinosinusitis Vs Sinusitis structures in this region are not clear enough to distinctly demarcate the optic lobes.

Pronounced and ventrolaterally displaced optic lobes are considered to reflect a more derived organization of the brain. Within coelurosaurs, and especially Maniraptoriformes, there is a clear trend for the expression of this feature [10][21][29]. Given that more basal coelurosaurs such as ornithomimosaurs also display the derived condition, it is likely that therizinosaurians also had ventrolaterally displaced optic lobes that were concealed within the dura, but we cannot rule out that they secondarily acquired the primitive, more dorsal position or were simply too small to make their position apparent.

Endo H, Frey R, editors. Share this article Share. I have created a range of material resources to support clients and others. BoxAtlantic Beach, New York The quest of Peaceful Minds is to offer effective treatment to those who have served our country and are now struggling to integrate back into the life they knew before. Nested between basal coelurosaurs and derived maniraptorans, this makes them of special importance not only in terms of the evolution of anatomical and developmental patterns, but also for the radiation of herbivory among theropods [5].

It is situated in the space between the endosseous labyrinths. The height, however, cannot be exactly determined, as a dural venous sinus covered the dorsal part of the endocast, obscuring cerebellar details. A moderately developed dural peak is present in all the therizinosaurian endocasts. Ventrally, the outline of the pons is indicated by an expanded area, which merges with the medulla oblongata. The latter is considerably enlarged and as wide as the cerebellum at its widest expansion, where it exits the braincase through the foramen magnum.

Athens Georgia Dating Free Artwork Of Rhinosinusitis Vs Sinusitis

The floccular lobes cerebellar auricles in all the therizinosaurian endocasts project caudolaterally from the cerebellar region and enter the space between the semicircular canals of the endosseous labyrinth. The floccular lobes are enlarged and have a round to oval cross-section in Erlikosauruswhereas they are transversely flattened in Falcarius and Nothronychus.

In Falcariusthe flocculi barely extend caudally beyond the crus communis Figure 6A. They extend further in Erlikosaurus and Nothronychusbut do not pass beyond the lateral and caudal semicircular canals, filling the space between the canals to a large extent.

Due to the lack of preservation of bony correlates, the pituitary fossa could not be faithfully reconstructed for Erlikosaurusand only partially click Falcarius and Nothronychus.