Forelimb Kinematics of Rats Using XROMM,with Implications for Small Eutherians and Their Fossil Relatives

The earliest eutherian mammals were small-bodied locomotor generalists with a forelimb morphology that strongly resembles that of extant rats. Understanding the kinematics of the humerus, radius, and ulna of extant rats can inform and constrain hypotheses concerning typical posture and mobility in early eutherian forelimbs. The locomotion of Rattus norvegicus has been extensively studied, but the three-dimensional kinematics of the bones themselves remains under-explored. Here, for the first time, we use markerless XROMM (Scientific Rotoscoping) to explore the three-dimensional long bone movements in Rattus norvegicus during a normal, symmetrical gait (walking). Our data show a basic kinematic profile that agrees with previous studies on rats and other small therians: rats maintain a crouched forelimb posture throughout the step cycle, and the ulna is confined to flexion/extension in a parasagittal plane. However, our three-dimensional data illuminate long-axis rotation (LAR) movements for both the humerus and the radius for the first time. Medial LAR of the humerus throughout stance maintains an adducted elbow with a caudally-facing olecranon process, which in turn maintains a cranially-directed manus orientation (pronation). The radius also shows significant LAR correlated with manus pronation and supination. Moreover, we report that elbow flexion and manus orientation are correlated in R. norvegicus: as the elbow angle becomes more acute, manus supination increases. Our data also suggest that manus pronation and orientation in R. norvegicus rely on a divided system of labor between the ulna and radius. Given that the radius follows the flexion and extension trajectory of the ulna, it must rotate at the elbow (on the capitulum) so that during the stance phase its distal end lies medial to ulna, ensuring that the manus remains pronated while the forelimb is supporting the body. We suggest that forelimb posture and kinematics in Juramaia, Eomaia, and other basal eutherians were grossly similar to those of rats, and that humerus and radius LAR may have always played a significant role in forelimb and manus posture in small eutherian mammals.     

Growth of the pectoral girdle of the Leopard frog, Rana pipiens (Anura: Ranidae)

This article describes the growth of the anuran pectoral girdle of Rana pipiens and compares skeletal development of the shoulder to that of long bones. The pectoral girdle chondrifies as two halves, each adjacent to a developing humerus. In each, the scapula and coracoid form as single foci of condensed chondrocytes that fuse, creating a cartilaginous glenoid bridge articulating with the humerus. Based on histological sections, both the dermal clavicle and cleithrum begin to ossify at approximately the same time as the periosteum forms around the endochondral bones. The dermal and endochondral bones of the girdle form immobile joints with neighboring girdle elements; however, the cellular organization and growth pattern of the scapula and coracoid closely resemble those of a long bone. Similar to a long bone epiphysis, distal margins of both endochondral elements have zones of hyaline, stratified, and hypertrophic cartilages. As a result, fused elements of the girdle can grow without altering the glenoid articulation with the humerus. Comparisons of anuran long bone and pectoral girdle growth suggest that different bones can have similar histology and development regardless of adult morphology.     

A functional-morphometric analysis of forelimbs in bipedal and quadrupedal heteromyid rodents

The rodent family Heteromyidae contains bipedal hoppers and quadrupedal runners. The possibility that bipedalism is associated with forelimb specialization for nonlocomotory functions, such as burrowing and seed-gathering, motivated a static functional-morphometric and interspecific allometric analysis of 18 metric characters of the forelimb skeleton. A principal-components analysis, across 28 species in six genera, showed that lengths of proximal (scapula, humerus) and distal (ulna, radius, metacarpal) elements were negatively allometric, and widths were positively allometric. Quadrupedal and bipedal species groups showed qualitatively similar allometric patterns, except that scapula width anterior to the spine was positively allometric in quadrupeds and negatively allometric in bipeds; scapula width posterior to the spine was positively allometric in bipeds and isometric in quadrupeds; and olecranon length was isometric in bipeds and positively allometric in quadrupeds. Most morphometric characters varied significantly among species within genera, even when effects of size variation were reduced by reconstructing all species to a common general size (as indicated by their score on the first principal component). These shape differences caused species to vary in the mechanical advantage of the forelimb, of possible importance for digging and seed-harvesting performance. Relative to quadrupeds, bipedal species tended to have greater mechanical advantage for proximal forelimb elements and smaller mechanical advantage for distal forelimb elements, but only the distal pattern remained in reconstructed forms, and no functional character was significantly different when tested over variation among genera nested within locomotion type. Cluster analysis confirmed that forelimb characters related to digging or seed-harvest are not coincident with mode of locomotion. Forelimb characters were, however, associated with digging or seed-harvest performance. Mechanical advantage of the proximal forelimb was positively related to an index of the compaction of soils with which 26 desert-dwelling species are associated, and also to relative use of heavy vs. light soils by nine species in the laboratory. Across 10 species, deviations in seed-harvest rate from expected allometric values were negatively correlated with mechanical advantage of the distal forelimb.     

Bony‐toothed birds (Aves: Pelagornithidae) from the Middle Eocene of Belgium

Abstract: We describe well‐preserved remains of the Pelagornithidae (bony‐toothed birds) from the middle Eocene of Belgium, including a sternum, pectoral girdle bones and humeri of a single individual. The specimens are tentatively assigned to Macrodontopteryx oweni Harrison and Walker, 1976 , which has so far only been known from the holotype skull and a referred proximal ulna. Another species, about two times larger, is represented by an incomplete humerus and tentatively identified as Dasornis emuinus ( Bowerbank, 1854 ). The fossils provide critical new data on the osteology of the pectoral girdle of bony‐toothed birds. For the first time, the sternum of one of the smaller species is preserved, and this bone exhibits a more plesiomorphic morphology than the recently described sternum of the giant Miocene taxon Pelagornis. The coracoid resembles that of the Diomedeidae (albatrosses) in overall morphology, but because bony‐toothed birds lack apomorphies of the Procellariiformes, the similarities are almost certainly owing to convergence. Bony‐toothed birds were often compared with the ‘Pelecaniformes’ by previous authors, who especially made comparisons with the Sulidae (gannets and boobies). However, the coracoid distinctly differs from that of extant ‘pelecaniform’ birds, and the plesiomorphic presence of a foramen nervi supracoracoidei as well as the absence of a well‐delimited articulation facet for the furcula supports a position outside the Suloidea, the clade to which the Sulidae belong.     

BILATERAL DIRECTIONAL ASYMMETRY OF THE APPENDICULAR SKELETON OF THE HARBOR PORPOISE (PHOCOENA PHOCOENA)

Directional asymmetry ( DA ) of the lengths, diameters, and masses of the scapula, humerus, radius, and ulna were analyzed on a sample of 213 harbor porpoises from Denmark and West Greenland. The levels of DA were consistent across yearlings and older animals, mature and immature animals, sexes, and populations. All investigated variables showed significant DA favoring the right side. For the pooled sample, DA of lengths ranged from 0.2% SE 0.1 (ulna) to 1.2% SE 0.1 (scapula). DA s of diameters were 1.5% SE 0.2 for the humerus, 0.6% SE 0.1 for the radius, and 1.3% SE 0.2 for the ulna. DA of mass ranged from 2.8% SE 0.5 (humerus) to 4.3% SE 0.7 (ulna). The humerus and ulna had significantly larger mean diameter/ length ratios on the right side than the left, making them more robust. The large DA of scapula length indicates larger muscle mass associated with the right flipper, while the generally more robust right humerii and ulnae may be designed for higher levels of mechanical stress. These DA s and the examples of lateralized behavior recorded in cetaceans, point to the existence of lateralized use of the flippers at the population level in harbor porpoises and possibly other cetacean species.     

陕甘宁盆地二马营组的假鳄类

在陕甘宁盆地东北缘二马营组发现的两种假鳄类,经鉴定均属Euparkeridae。其中之一与Euparkeria较相近;根据肩胛骨、肱骨和骨质背板等的显著不同,定名为侨家梁哈拉寨鳄(Halazhaisuchus qiaoensis),新属、新种。依据骨髂形态,另一标本归入世鲁番鳄属(Turfanosuchus),取名沙圪堵吐鲁番鳄(Turfanosuchus shageduensis)、新种。     

Age estimation of adolescent and young adult male and female skeletons II, epiphyseal union at the upper limb and scapular girdle in a modern Portuguese skeletal sample

This study completes previously reported ages for timing of epiphyseal union in the postcranial skeleton in a recent sample, with data from the scapula, clavicle, humerus, radius, and ulna. A sample of 121 individuals between the ages of 9 and 29 (females = 65, males = 56) was derived from the Lisbon documented skeletal collection. Epiphyseal union was scored at 16 anatomical locations, using a three-stage scheme: 1) no union; 2) partial union; and 3) completed union, all traces of fusion having disappeared. In the upper limb, the epiphyses of the elbow are the first to fuse at around 11 to 15 years of age, followed by those of the shoulder and wrist. In the scapular girdle, the coracoid area is the first to fuse, followed by the glenoid surface and remaining epiphyses, with the medial clavicle fusing last, by the age of 25-27. There is a sex difference in maturation, with females showing an advance relative to males of about 2 years in the upper limb. Sex differences in maturation are less noticeable in the scapular girdle, but data suggest that females are also ahead of males. Results suggest overall similar age ranges for stages of union as previous dry bone observations, but some studies show significant divergences which seem to derive from methodological issues. Although some radiographic reference standards provide comparable age ranges, they should probably be avoided when aging skeletal remains.     

Morphological variation in the appendicular skeleton of atlantic forest sigmodontine rodents

Rodents of the subfamily Sigmodontinae comprise a highly diversified group in the Atlantic Forest, with semifossorial, terrestrial, semiaquatic, scansorial, and arboreal forms. In this study, we analyzed morphometric variation in humerus, scapula, ulna, radius, femur, tibia, and pelvis to investigate its possible relationship with the different types of locomotion recorded in the literature. Skeletal characters were measured in 321 specimens belonging to 29 species and 19 genera either restricted to or recorded in this ecoregion. Multivariate morphometric analyses (principal component and canonical variate analyses) arranged individuals of different genera in groups congruent with the different types of locomotion. This arrangement was more clearly defined when analyses included only forelimb measurements, indicating that most of the variation in appendicular traits associated with the different locomotor modes occurs in the forelimb skeleton. Semifossorial forms exhibited the most distinct appendicular morphology, as well as the greatest frequency of endemism among analyzed species. These results suggest that this mode of locomotion led to greater differentiation in semifossorial Atlantic forest sigmodontines than in terrestrial and arboreal forms, which were found to have more subtle differentiation and fewer endemics. Scansorial species could not be set apart from terrestrial ones in terms of appendicular morphology, suggesting that these two modes of locomotion are the most similar and generalized for the group, as they occur in most lineages in the subfamily. The results of this study corroborate previous observations on the relevance of appendicular characters in the differentiation of species and genera in the subfamily Sigmodontinae. J. Morphol. 2013. © 2013 Wiley Periodicals, Inc.     

The role of the forelimb in prey capture in the Late Triassic reptile Megalancosaurus (Diapsida,Drepanosauromorpha)

The pectoral girdle and forelimb of the Late Triassic drepanosauromorph reptile Megalancosaurus are redescribed and their function reinterpreted. The whole skeleton of this diapsid is highly specialised for arboreal life, and also the peculiarities of the shoulder girdle and forelimb were interpreted as adaptations for a limb-based locomotion using gap-bridging to move from one support to another, as in chameleons. Re-examination of the pectoral girdle and forelimb revealed the presence of clavicles fused into a furcula-like structure, a saddle-shaped glenoid and a tight connection between the radius and ulna that strengthened the forearm but hindered pronation and supination movements at that joint. The new information plus a reconstruction of the pectoral and forelimb musculature suggests that the forelimb was also specialised for grasping and raking in addition to climbing and thus prey capture may have been an important function for the forelimb. The new functional interpretation fits well with the overall body architecture of Megalancosaurus’ skeleton, suggesting that this reptile was an ambush predator that may have assumed a stable tripodal position, secured by the hooked tail and hind limbs, freeing its forelimbs to catch prey by sudden extension of the arm and firm grasping with the pincer-like digits.     

Perforation of the coronoid-olecranon septum. Humero-ulnar relationships in Netherlands and African populations

The mechanical interaction of the humerus and ulna during articulation has been examined in skeletal material from African and European populations. It was found that the presence or absence of perforation of the humeral septum and the thickness of the septum when unperforated can be predicted with considerable accuracy when two factors are known: the length of the coronoid and olecranon processes of the ulna relative to the minimum diameter of the trochlea humeri, and the total angle of movement at the elbow from full extension to full flexion. The influences which control either or both of these factors also determine the thickness of the septum of the humerus. These influences are not well known and there is opportunity for both genetic and environmental effects. The coefficients of correlation between septum thickness, flexion angle, humeral robusticity, minimum circumference of the humerus, trochlea diameter and brachial index are given. Septum thickness is correlated more highly with trochlea diameter than with robusticity or minimum circumference of the humerus. The higher frequency of perforation of the septum in the African sample (47%) compared with the European (6%) is associated with a greater range of movement at the elbow in the Africans. Bilateral, sexual and population differences in the frequency of perforation of the humeral septum may be associated with differences in the extent of movement at the elbow joint. Variation in this feature, in turn, may be associated with muscularity and somatotype.     

New genera of baleen whales (Cetacea,Mammalia) from the Miocene of the northern Caucasus and Ciscaucasia: 3. Zygiocetus gen. nov. (Middle Sarmatian,Adygea)

Based on an almost complete skeleton (skull, scapula fragment, humerus, ulna, radius, 7 cervical, 12 thoracic, 17 lumbar and caudal vertebrae, sternum, and hyoid), a new cetotheriid, Zygiocetus nartorum gen. et sp. nov., is described. It comes from the Middle Sarmatian beds (Upper Miocene) of the Krasnooktyabr’sk Formation of the Polevoe 1 locality (Republic of Adygea).     

Reidentification of Avian Embryonic Remains from the Cretaceous of Mongolia

Embryonic remains within a small (4.75 by 2.23 cm) egg from the Late Cretaceous, Mongolia are here re-described. High-resolution X-ray computed tomography (HRCT) was used to digitally prepare and describe the enclosed embryonic bones. The egg, IGM (Mongolian Institute for Geology, Ulaanbaatar) 100/2010, with a three-part shell microstructure, was originally assigned to Neoceratopsia implying extensive homoplasy among eggshell characters across Dinosauria. Re-examination finds the forelimb significantly longer than the hindlimbs, proportions suggesting an avian identification. Additional, postcranial apomorphies (strut-like coracoid, cranially located humeral condyles, olecranon fossa, slender radius relative to the ulna, trochanteric crest on the femur, and ulna longer than the humerus) identify the embryo as avian. Presence of a dorsal coracoid fossa and a craniocaudally compressed distal humerus with a strongly angled distal margin support a diagnosis of IGM 100/2010 as an enantiornithine. Re-identification eliminates the implied homoplasy of this tri-laminate eggshell structure, and instead associates enantiornithine birds with eggshell microstructure composed of a mammillary, squamatic, and external zones. Posture of the embryo follows that of other theropods with fore- and hindlimbs folded parallel to the vertebral column and the elbow pointing caudally just dorsal to the knees. The size of the egg and embryo of IGM 100/2010 is similar to the two other Mongolian enantiornithine eggs. Well-ossified skeletons, as in this specimen, characterize all known enantiornithine embryos suggesting precocial hatchlings, comparing closely to late stage embryos of modern precocial birds that are both flight- and run-capable upon hatching. Extensive ossification in enantiornithine embryos may contribute to their relatively abundant representation in the fossil record. Neoceratopsian eggs remain unrecognized in the fossil record.     

Skeletal indicators of ecological specialization in pika (Mammalia,Ochotonidae)

Pika species generally fall into two ecotypes, meadow‐dwelling (burrowing) or talus‐dwelling, a classification that distinguishes a suite of different ecological, behavioral, and life history traits. Despite these differences, little morphological variation has previously been documented to distinguish among ecotypes. The aim of this study was to test whether postcranial features related to burrowing are present in meadow‐dwelling species and whether talus‐dwelling species exhibit postcranial modifications related to frequent leaping between rocks. To test this, the scapula, humerus, ulna, radius, innominate, femur, tibia, and calcaneus of 15 species were studied and measured. Twenty‐three measurements were taken on 199 skeletons, and 19 indices were constructed from these measurements. Indices were compared between the two ecotypes using Student's t‐test. Comparisons among ecotypes, species, and subgenera were made using one‐way ANOVA with the Tukey honest significant difference post hoc test. Multivariate results were generated using principal components analyses. Thirteen forelimb and hind limb indices proved significant in distinguishing the meadow‐dwelling, talus‐dwelling, and intermediate forms. A number of these indices are associated with burrowing or leaping in other mammals, providing some support for the hypothesis that postcranial modifications in pika are related to locomotor differences. This evidence of morphological responses to ecological specialization will be useful for reconstructing the paleobiology of extinct taxa, assessing the behavioral variability of extant species, and improving our understanding of the evolutionary history of pikas. J. Morphol., 2013. © 2013 Wiley Periodicals, Inc.     

Scaling of long bones in ruminants with respect to the scapula

The significance of the scapula for locomotion is becoming more and more established. Studies of locomotion in small and medium‐sized mammals show a considerable amplitude of the scapula and a large contribution to step length. Taking this into account, long bone studies of forelimb movement restricted to the ‘arm’ miss one important segment. A regression model (reduced major axis) was used for analysis of a sample of 77 species of ruminants. This sample was divided according to (1) phylogenetic relationships and (2) habitat. The proximal elements of the limbs, scapula and humerus in the anterior extremity, femur in the hindlimb, show a similar scaling in the different analyses. The changes to limb proportions in the different subsamples are caused by the variability of the distal segments. The anterior extremity scales with a higher coefficient than the hindlimb in all analyses. Concepts like elastic or geometric similarity are inadequate for long bone scaling when the full range of body size in the sample is used. Taking all analyses into account, the differences in limb proportions are due more to phylogenetic relationships than to habitat.     

Integration of the mammalian shoulder girdle within populations and over evolutionary time

Morphological integration has the potential to link morphological variation within populations with morphological evolution among species. This study begins to investigate this link by comparing integration among shoulder girdle elements (e.g. scapular blade, glenoid, coracoid, etc.) during the origin and evolution of therian mammals, and within modern bat, opossum and mouse populations. In this study, correlations among skeletal elements and patterns of allometry are used as proxies for integration. Results suggest that shoulder girdle elements tended to vary and evolve independently during the origin of mammals and subsequent radiation of placentals, consistent with the elements’ distinct developmental and evolutionary origins. This finding suggests that skeletal element correlations, and therefore integration, can be conserved over large taxonomic and temporal scales. However, marsupials display a different pattern in which shoulder girdle elements tend to be more integrated, with the exception of the coracoid. This finding is consistent with a shift in the pattern of skeletal element integration coincident with the appearance of the marsupial mode of reproduction. This finding provides further evidence that development can play a significant role in the establishment of patterns of skeletal element correlation and that patterns of skeletal element correlation can themselves evolve when faced with sufficient selective pressures.     

The developmental reduction of the marsupial coracoid: A case study in Monodelphis domestica

During their embryogenesis, marsupials develop a unique structure, the shoulder arch, which provides the structural and muscle‐attachment support necessary for the newborn's crawl to the teat. One of the most pronounced and important aspects of the shoulder arch is an enlarged coracoid. After marsupial newborns reach the teat, the shoulder arch is remodeled and the coracoid is reduced to a small process on the scapula. Although an understanding of marsupial coracoid reduction has the potential to provide insights into both, marsupial evolution and the origin of mammals, little is known about the morphological and cellular processes controlling this process. To remedy this situation, this study examined the morphological and cellular mechanisms behind coracoid reduction in the gray short‐tailed opossum, Monodelphis domestica. A quantitative, morphometric study of shoulder girdle development revealed that the coracoid is reduced in size relative to other aspects of the shoulder girdle by growing at a slower rate. Using a series of molecular assays for cell death, no evidence was found for programmed cell death playing a role in the reduction of coracoid size in marsupials (in contrast to hypotheses of previous researchers). Although it is likely the case that coracoid growth is reduced through a relatively lower rate of cellular proliferation, differences in proliferative rates in the coracoid and scapula were not great enough to be quantified using standard molecular assays. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.     

Appendicular skeleton in Bachia bicolor (Squamata: Gymnophthalmidae): osteology,limb reduction and postnatal skeletal ontogeny

The osteology of the appendicular skeleton and its postnatal development are described in Bachia bicolor, a serpentiform lizard with reduced limbs. The pectoral girdle is well developed and the forelimb consists of a humerus, ulna, radius, five carpal elements (ulnare, radiale, distal carpals 4–3, centrale), four metacarpals (II, III, IV, V) and phalanges (phalangeal formula X‐2‐2‐2‐2). In the hindlimb, the femur is small and slender, and articulates distally with a series of ossified amorphous and extremely reduced elements that correspond to a fibula, tibia and proximal and distal tarsals 4 and 3. The pelvic girdle consists of ischium, pubis and ilium, but its two halves are widely separated; the ilium is the least reduced element. We describe the ossification and development during postnatal skeletal ontogeny, especially of epiphyseal secondary centres, ossifications of carpal elements, apophyseal ossifications and sesamoids. Compared to other squamates, B. bicolor shows an overall reduction in limb size, an absence of skeletal elements, a fusion of carpal elements, an early differentiation of apophyseal centres, and a low number of sesamoids and apophyseal centres. These observations suggest that the reductions are produced by heterochronic changes during postnatal development and probably during embryonic development; therefore the appendicular skeleton exhibits a pattern of paedomorphic features.     

四川自贡大山铺中侏罗世肉食龙一新种

自贡四川龙(Szechuanosourus zigongensis sp.nov.)为自贡市大山铺蜀龙动物群中发现的又一肉食龙新材料,本文对它进行了记述。     

Shoulder movements during quadrupedal locomotion in arboreal primates

The kinematics of scapula and shoulder joint movements were analyzed in three species of arboreal quadrupedal primates using cineradiography. Our findings indicate that scapular movement is highly important for forelimb movement in primates with this ancestral mode of locomotion. Retroversion of the scapula (syn. caudal rotation or extension) during the stance phase contributes more than 40% to the stride length of the forelimb. Lateral forelimb excursions, a general feature for arboreal primates, are based on complex three-dimensional scapular movements guided by the clavicle. Humeral abduction is achieved by scapular abduction and transversal rotation of the scapula about its longitudinal axis, and is therefore strikingly different from humeral abduction in humans. At the same time, the movements of the shoulder joint are limited to flexion and extension only.