Low morphological disparity and decelerated rate of limb size evolution close to the origin of birds.

The origin of birds from theropod dinosaurs involves many changes in musculoskeletal anatomy and epidermal structures, including multiple instances of convergence and homology-related traits that contribute to the refinement of flight capability. Changes in limb sizes and proportions are important for locomotion (for example, the forelimb for bird flight); thus, understanding these patterns is central to investigating the transition from terrestrial to volant theropods. Here we analyse the patterns of morphological disparity and the evolutionary rate of appendicular limbs along avialan stem lineages using phylogenetic comparative approaches. Contrary to the traditional wisdom that an evolutionary innovation like flight would promote and accelerate evolvability, our results show a shift to low disparity and decelerated rate near the origin of avialans that is largely ascribed to the evolutionarily constrained forelimb. These results suggest that natural selection shaped patterns of limb evolution close to the origin of avialans in a way that may reflect the winged forelimb 'blueprint' associated with powered flight.

Magnetic resonance imaging, computed tomography, and gross anatomy of forelimb joints in New Zealand rabbit (Oryctolagus cuniculus L.).

The present study aimed to define the anatomical structures by comparing the transversal computed tomography (CT) and magnetic resonance (MR) images of the forelimb joints of the rabbits with the cross-sectional plastinated images. A total of 14 (seven females, seven males) one-year-old adult New Zealand rabbits were used in the study. After the CT and MR imaging procedures were completed, the forelimbs were removed from the body. The forelimbs were plastinated using the silicone plastination method and sectioned transversal. Cross-sectional plastinates were evaluated and correlated anatomically with conjugate CT and MR images. Joint and surrounding anatomical structures were defined in sections. Cross-sectional plastinated samples were highly correlated with CT and MR images in terms of bone and soft tissue, respectively. It is thought that the anatomical and radiological data obtained from the forelimb joints of rabbits will provide a basis for scientists who are involved in both experimental surgical interventions and clinical anatomy education.

[Anatomical observation and CT three-dimensional reconstruction of Five-shu acupoints of Shaoyin Heart Meridian in rabbit's forelimb].

OBJECTIVE: To investigate the location and anatomical structure of "Shaochong"(HT9), "Shaofu"(HT8), "Shenmen"(HT7), "Lingdao"(HT4) and "Shaohai"(HT3) in the rabbit's forelimb. METHODS: Sixteen rabbits (half male and half female) were used in the present study. By referring to the national standards on the location of acupoints in the human body and the literature about the location of acupoints in the rabbit, and by using the method of comparative anatomy, the location and needling operation of the Five-shu acupoints of Shaoyin Heart Meridian on the rabbit's forelimb were defined, and these acupoints were needled and CT three-dimensional reconstruction were conducted. Then, the rabbits were killed, and intravascular perfusion was performed, followed by inserting acupuncture needles into these five acupoints for observing the anatomical relationship between the inserted acupuncture needle and the structure of surrounding tissues. RESULTS: HT9 is located at the medial side of the little finger of forelimb, about 1 mm beside the nail root, and is adjacent to the superficial flexor tendon of the finger, the dorsal branches of the proper palmar digital artery and vein, and the endings of dorsal branch of palmar digital proper nerve of the ulnar nerve on the fifth finger side. HT8 is located at the palm side of the forelimb, horizontally parallel to the proximal end of the 5th metacarpophalangeal joint and between the 4th and 5th metacarpal bones, and is adjacent to the lumbricalis, the 4th and 5th interossei, and common palmar digital artery and vein and the palmar digital proper nerve of the ulnar nerve. HT7 is located at the medial margin of the extensor carpal tendon on the ulnar side, between the distal end of the ulna and the ulnar carpal bone, and is adjacent to the tendons of flexor carpi ulnaris and extensor carpi ulnaris, ulnar artery, ulnar vein and ulnar nerve. HT4 is located at the medial border of the ulnar flexor tendon, about 1.5 cun superior to HT7, and is adjacent to extensor carpi ulnaris, flexor carpi ulnaris, flexor digitorum superficialis, flexor digitorum profundus, ulnar artery, vein and ulnar nerve. HT3 is located at the depression, medial to the condyle of humerus when the elbow is bent at 90°, its neighbor structure is composed of pronator teres, biceps brachii, brachial artery and vein, radial collateral artery, radial collateral vein, medial antebrachial cutaneous nerve and median nerve. CONCLUSION: In the rabbit, there is a close relationship between HT9, HT8, HT7, HT4 and HT3 regions and brachial vascular and its branches, cephalic vein and its branches, medial antebrachial cutaneous nerve, median nerve and ulnar nerve, which is the morphological basis of the Five-shu acupoints of Shaoyin Heart Meridian for treating some related clinical disorders.

Fossorial mammals emphasise the forelimb muscle moment arms used for digging: New indices for reconstruction of the digging ability and behaviours in extinct taxa.

Among fossorial mammals, forelimbs are major digging apparatuses for dwelling, sheltering and foraging underground. Forelimb-diggers have independently evolved in many lineages of mammals; thus, the method of digging with forelimbs varies by taxon. Therefore, the reconstruction of digging behaviours in extinct animals leads us to understand the evolutionary process of fossorial adaptation in each lineage. However, no morphological index was found to reconstruct if, or how, extinct taxa dug with forelimbs. In this study, we used the shoulder and elbow muscle moment arms in relation to the out-force lever on the manus as indices of the efficiency of motions. The mechanical advantage of two shoulder motions (medial rotation and retraction) and three elbow motions (extension, flexion and adduction) was measured in 381 extant mammal specimens representing 332 species, 279 genera, 103 families and 24 orders. Assuming that both forelimb-digging and -paddling in water require relatively high-output moment arm efficiency, the studied taxa were categorised into four groups based on the presence or absence of forelimb-digging and -paddling abilities. We found that the efficiencies of all five muscle moment arms in the forelimb-diggers and -paddlers were higher than those of the non-diggers and non-paddlers. Furthermore, among the forelimb-diggers, the taxa that dig compact substrates or frequently burrow tend to emphasise the muscle moment arms compared to the taxa that dig loose substrates or dig less frequently. The comparison among the 53 extant forelimb-diggers revealed that the efficiency marked among the five muscle moment arms reflects the difference in digging strategy: humeral rotation diggers emphasise the shoulder medial rotator and elbow adductor, hook-and-pull diggers emphasise the shoulder retractor and elbow flexor and scratch diggers emphasise the shoulder retractor and elbow extensor. We propose that these indices will be powerful tools for reconstructing the fossorial behaviours of extinct mammals. Applying these indices to extinct taxa, Ceratogaulus, Ernanodon, Metacheiromys and Prozaedyus are capable of more efficient forelimb-digging, and each may have adopted different digging strategies.

Functional space analyses reveal the function and evolution of the most bizarre theropod manual unguals.

Maniraptoran dinosaurs include the ancestors of birds, and most used their hands for grasping and in flight, but early-branching maniraptorans had extraordinary claws of mysterious function. Alvarezsauroids had short, strong arms and hands with a stout, rock-pick-like, single functional finger. Therizinosaurians had elongate fingers with slender and sickle-like unguals, sometimes over one metre long. Here we develop a comprehensive methodological framework to investigate what the functions of these most bizarre bony claws are and how they formed. Our analysis includes finite element analysis and a newly established functional-space analysis and also involves shape and size effects in an assessment of function and evolution. We find a distinct functional divergence among manual unguals of early-branching maniraptorans, and we identify a complex relationship between their structural strength, morphological specialisations, and size changes. Our analysis reveals that efficient digging capabilities only emerged in late-branching alvarezsauroid forelimbs, rejecting the hypothesis of functional vestigial structures like T. rex. Our results also support the statement that most therizinosaurians were herbivores. However, the bizarre, huge Therizinosaurus had sickle-like unguals of such length that no mechanical function has been identified; we suggest they were decorative and lengthened by peramorphic growth linked to increased body size.

Reconstruction of the pectoral girdle and forelimb musculature of Megaraptora (Dinosauria: Theropoda).

Megaraptora is a group of enigmatic, carnivorous non-avian theropod dinosaurs from the Cretaceous of Asia, Australia, and especially South America. Perhaps the most striking aspect of megaraptoran morphology is the large, robustly constructed forelimb that, in derived members of the clade, terminates in a greatly enlarged manus with hypertrophied, raptorial unguals on the medialmost two digits and a substantially smaller ungual on digit III. The unique forelimb anatomy of megaraptorans was presumably associated with distinctive functional specializations; nevertheless, its paleobiological significance has not been extensively explored. Here we draw from observations of the pectoral girdle and forelimb skeletons of Megaraptora and myological assessments of other archosaurian taxa to provide a comprehensive reconstruction of the musculature of this anatomical region in these singular theropods. Many muscle attachment sites on megaraptoran forelimb bones are remarkably well developed, which in turn suggests that the muscles themselves were functionally significant and important to the paleobiology of these theropods. Furthermore, many of these attachments became increasingly pronounced through megaraptoran evolutionary history, being substantially better developed in derived taxa such as Australovenator wintonensis and especially Megaraptor namunhuaiquii than in early branching forms such as Fukuiraptor kitadaniensis. When considered alongside previous range of motion hypotheses for Australovenator, our results indicate that megaraptorans possessed a morphologically and functionally specialized forelimb that was capable of complex movements. Notable among these were extensive extension and flexion, particularly in the highly derived manus, as well as enhanced humeral protraction, attributes that very probably aided in prey capture.

Elbow-joint morphology in the North American 'cheetah-like' cat Miracinonyx trumani.

The North American cheetah-like cat Miracinonyx trumani is an extinct species that roamed the Pleistocene prairies 13 000 years ago. Although M. trumani is more closely related to the cougar (Puma concolor) than to the living cheetah (Acinonyx jubatus), it is believed that both A. jubatus and M. trumani possess a highly specialized skeleton for fast-running, including limbs adapted for speed at the expense of restricting the ability of prey grappling. However, forelimb dexterity of M. trumani has not been yet investigated. Here, we quantify the 3D-shape of the humerus distal epiphysis as a proxy for elbow-joint morphology in a sample of living cats to determine whether the extinct M. trumani was specialized to kill open-country prey using predatory behaviour based on fast running across the prairies and steppe terrains of the North American Pleistocene. We show that M. trumani had an elbow morphology intermediate to that of P. concolor and A. jubatus, suggesting that M. trumani had a less specialized pursuit predatory behaviour than A. jubatus. We propose that M. trumani probably deployed a unique predatory behaviour without modern analogues. Our results bring into question the degree of ecomorphological convergence between M. trumani and its Old World vicar A. jubatus.

Early diversification of avian limb morphology and the role of modularity in the locomotor evolution of crown birds.

High disparity among avian forelimb and hind limb segments in crown birds relative to non-avialan theropod dinosaurs, potentially driven by the origin of separate forelimb and hind limb locomotor modules, has been linked to the evolution of diverse avian locomotor behaviors. However, this hypothesized relationship has rarely been quantitatively investigated in a phylogenetic framework. We assessed the relationship between the evolution of limb morphology and locomotor behavior by comparing a numerical proxy for locomotor disparity to morphospace sizes derived from a dataset of 1,241 extant species. We then estimated how limb disparity accumulated during the crown avian radiation. Lastly, we tested whether limb segments evolved independently between each limb module using phylogenetically informed regressions. Hind limb disparity increased significantly with locomotor disparity after accounting for clade age and species richness. We found that forelimb disparity accumulated rapidly early in avian evolution, whereas hind limb disparity accumulated later, in more recent divergences. We recovered little support for strong correlations between forelimb and hind limb morphology. We posit that these findings support independent evolution of locomotor modules that enabled the striking morphological and behavioral disparity of extant birds.

Muscle architecture and muscle fibre type composition in the forelimb of two African mole-rat species, Bathyergus suillus and Heterocephalus glaber.

The scratch-digging Cape dune mole-rat (Bathyergus suillus), and the chisel-toothed digging naked mole-rat (Heterocephalus glaber) are African mole-rats that differ in their digging strategy. The aim of this study was to determine if these behavioural differences are reflected in the muscle architecture and fibre-type composition of the forelimb muscles. Muscle architecture parameters of 39 forelimb muscles in both species were compared. Furthermore, muscle fibre type composition of 21 forelimb muscles were analysed using multiple staining protocols. In B. suillus, muscles involved with the power stroke of digging (limb retractors and scapula elevators), showed higher muscle mass percentage, force output and shortening capacity compared to those in H. glaber. Additionally, significantly higher percentages of glycolytic fibres were observed in the scapular elevators and digital flexors of B. suillus compared to H. glaber, suggesting that the forelimb muscles involved in digging in B. suillus provide fast, powerful motions for effective burrowing. In contrast, the m. sternohyoideus a head and neck flexor, had significantly more oxidative fibres in H. glaber compared to B. suillus. In addition, significantly greater physiological cross-sectional area and fascicle length values were seen in the neck flexor, m. sternocleidomastoideus, in H. glaber compared to B. suillus, which indicates a possible adaptation for chisel-tooth digging. While functional demands may play a significant role in muscle morphology, the phylogenetic differences between the two species may play an additional role which needs further study.

Reversibility of digit loss revisited: Limb diversification in Bachia lizards (gymnophthalmidae).

Strict interpretations of the Dollo's Law lead to postulation that trait loss is irreversible and organisms never recover ancestral phenotypes. Dollo, however, admitted the possibility of reversals in trait loss when predicted differences between reversed (derived) and ancestral forms. Phenotypic signatures from reversals are expected, as the historical context of a reversal in trait loss differs from the initial setting where the trait originally evolved. This article combines morphological and molecular information for Bachia scolecoides to discuss phenotypic and genetic patterns established during processes that reversed digit loss in Gymnophthalmidae (also termed microteiid lizards). Results suggest that pathways leading to the derived tetradactyl state of B. scolecoides comprise particularities in their origin and associated processes. Autopodial bones of B. scolecoides lack digit identity, and muscle anatomy is very similar between manus and pes. Gymnophthalmidae sequence patterns in the limb-specific sonic hedgehog enhancer (ZRS) suggest that regulation of shh expression did not degenerate in Bachia, given the prediction of similar motifs despite mutations specific to Bachia. Persistence of developmental mechanisms might explain intermittent character expression leading to reversals of digit loss, as ZRS signaling pathways remain active during the development of at least one pair of appendices in Bachia, especially if some precursors persisted at early stages. Patterns of ZRS sequences suggest that irreversibility of trait loss might be lineage-specific (restricted to Gymnophthalmini) and contingent to the type of signature established. These results provide insights regarding possible mechanisms that may allow reactivation of developmental programs in specific regions of the embryo.

Morphological and radiological study of the shoulder and elbow joints of the giant anteater (Myrmecophaga tridactyla).

The giant anteater (Myrmecophaga tridactyla) has specialized thoracic limbs to forage by breaking the walls of anthills and termite mounds. They also play critical roles in defense posture and locomotion. This study aimed to provide a morphological and radiographic description of the shoulder joint and elbow joint of the giant anteater. Both joints of 13 tamanduas were assessed by morphological dissections and histological evaluation and radiographies without and with positive contrast. The radiographic projections selected to this study were the mediolateral and craniocaudal projections. The radiographic and anatomical findings were compared with the following results: the shoulder joint had a continuous joint capsule with the tendon sheaths of the short head and long head of the biceps brachii muscle, which could be visualized with an injection of 3 ml of intra-articular contrast. The shoulder joint arthrography was performed with the needle positioned cranially to the joint for contrast injection. The elbow joint presented three articular compartments, and the insertion of the joint capsule was proximal to the radial fossa and distal to the radial tuberosity in the radial notch of the ulna, which were possible to identify with 2 ml of intra-articular contrast. The elbow joint arthrography was performed with the needle positioned laterally to the joint for contrast injection. Moreover, the joint capsule presented a caudomedial distention and fat pads. The powerful muscles of the forelimb play a fundamental role in maintaining the shoulder joint and elbow joint stability due to bony adaptations and the absence of usual ligaments. The morphological and radiological study provided relevant information on the soft-tissue characteristics of shoulder and elbow joints, which may aid clinical-surgical and diagnostic imaging procedures.

Osteology and radiographic anatomy of the thoracic limb of the greater cane rat (Thryonomys swinderianus).

The greater cane rat (Thryonomys swinderianus) is mainly distributed in the humid and sub-humid regions of the sub-Saharan Africa. It is believed to be the second largest African rodent. The aim of this study was to describe the normal osteology and radiographic anatomy of the thoracic limb of the greater cane rat as a reference for teaching, clinical use, anatomical studies and biomedical research. Five intact dead greater cane rats were used for radiographic examination. Radiographic findings were correlated with bone specimens. The clavicle was seen in all animals. The scapula was wide with well-developed acromion and coracoid process. The major and minor tubercles did not extend higher than the head of the humerus. The deltoid tuberosity was very prominent. The medial epicondyle was larger than its lateral counterpart, and the supratrochlear foramen was present. The radial tuberosity was further distally located. The ulnar head was well-developed, and the olecranon tuber was square-shaped. The distal physes for the radius and ulna appeared transverse. The carpus had eight bones. The manus presented with five widely spread digits. The first digit was reduced in size and placed such that the dorsal and palmar surfaces face almost medial and lateral, respectively. Additionally, two rudimentary digits were seen. Variations exist in the normal osteology and radiographic anatomy of the thoracic limb in different species. Knowledge of the normal osteology and radiographic anatomy of the thoracic limb of individual species is important for species identification and diagnosis of musculoskeletal diseases involving the thoracic limb.

Ultrasonographic Evaluation of the Suspensory Ligament in Quarter Horses Used for Cutting.

Ultrasound remains a mainstay in proximal suspensory ligament (PSL) evaluation. Despite recent improvements facilitating earlier diagnosis/treatment, needs exist for (1) characterization of normal ultrasonographic PSL cross-sectional area (CSA) in specific breeds/disciplines, (2) improved detection of early/subtle changes in the PSL, and (3) an understanding of suspensory ligament (SL) morphology change from work-related stress. The objectives of this study were to establish normal PSL CSA in Quarter Horses (QH) used for cutting via angle contrast ultrasonography, ultrasonographically monitor SL morphology as horses progress through training, and assess the impact of PSL disease on performance (Lifetime Earnings - $LTE). One hundred ten 2-year-old cutting horses had PSL ultrasound examination on all limbs, and examinations were repeated 2 years later on 51 remaining horses. PSL CSA at select locations was compared between time points and entire examinations subjectively graded by an equine radiologist. $LTE was obtained from the National Cutting Horse Association for each horse and analyzed by abnormality number, location, and development over time. A total of 267 PSL ultrasound examinations across both times points were measured and graded. Normal CSA was established by zone for forelimbs and hindlimbs. The prevalence of abnormal limbs increased between the 2-year-old and 4-year-old years, forelimb abnormalities were statistically more prevalent than hindlimb abnormalities at both time points, and osseous attachment abnormalities were statistically more prevalent than ligament abnormalities at both times points. No significant differences in PSL CSA change over time were seen in any group and there was no significant difference in $LTE among or between groups.

The forelimbs of Alvarezsauroidea (Dinosauria: Theropoda): Insight from evolutionary teratology.

Alvarezsauroidea (Tetanurae) are nonavian theropod dinosaurs whose forelimb evolution is characterised by the overdevelopment of digit I, at the expense of the other two digits, complemented by a drastic forelimb shortening in derived species (Parvicursorinae). These variations are recognised as evolutionary developmental anomalies. Evolutionary teratology hence leads to a double diagnosis with (1) macrodactyly of digit I and microdactyly of digits II and III, plus (2) anterior micromelia. The teratological macrodactyly/microdactyly coupling evolved first. Developmental mechanisms disturbing limb proportion are thought to be convergent with those of other Tetanurae (Tyrannosauridae, Carcharodontosauridae). As for the manual anomalies, both are specific to Alvarezsauroidea (macrodactyly/microdactyly) and inherited (digit loss/reduction). While considering the frame-shift theory, posterior digits develop before the most anterior ones. There would therefore be a decrease in the area devoted to digits II (condensation 3) and III (condensation 4), in connection with the Shh signalling pathway, interacting with other molecular players such as the GLI3 protein and the Hox system. Developmental independence of digit I (condensation 2) would contribute to generating a particular morphology. Macrodactyly would be linked to a variation in Hoxd-13, impacting Gli3 activity, and increasing cell proliferation. The loss/reduction of digital ray/phalanges (digits II and III), would be associated with Shh activity, a mechanism inherited from the theropodan ancestry. The macrodactyly/microdactyly coupling, and then anterior micromelia, fundamentally changed the forelimb mechanical function, compared to the 'classical' grasping structure of basal representatives and other theropods. The distal ossification of the macrodactylian digit has been identified as physiological, implying the use of the structure. However, the debate on a particular 'adaptive' use is pointless as the ecology of an organism is interactively complex, being both at the scale of the individual and dependent on circumstances. Other anatomical features also allow for compensation and different predation (cursorial hindlimbs).

Low effective mechanical advantage of giraffes' limbs during walking reveals trade-off between limb length and locomotor performance.

Giraffes (Giraffa camelopardalis) possess specialized locomotor morphology, namely elongate and gracile distal limbs. While this contributes to their overall height and enhances feeding behavior, we propose that the combination of long limb segments and modest muscle lever arms results in low effective mechanical advantage (EMA, the ratio of in-lever to out-lever moment arms), when compared with other cursorial mammals. To test this, we used a combination of experimentally measured kinematics and ground reaction forces (GRFs), musculoskeletal modeling, and inverse dynamics to calculate giraffe forelimb EMA during walking. Giraffes walk with an EMA of 0.34 (±0.05 SD), with no evident association with speed within their walking gait. Giraffe EMA was about four times lower than expectations extrapolated from other mammals, ranging from 0.03 to 297 kg, and this provides further evidence that EMA plateaus or even diminishes in mammals exceeding horse size. We further tested the idea that limb segment length is a factor which determines EMA, by modeling the GRF and muscle moment arms in the extinct giraffid Sivatherium giganteum and the other extant giraffid, Okapia johnstoni. Giraffa and Okapia shared similar EMA, despite a four to sixfold difference in body mass (Okapia EMA = 0.38). In contrast, Sivatherium, sharing a similar body mass with Giraffa, had greater EMA (0.59), which we propose reflects behavioral differences, such as a somewhat increased capability for athletic performance. Our modeling approach suggests that limb length is a determinant of GRF moment arm magnitude and that unless muscle moment arms scale isometrically with limb length, tall mammals are prone to low EMA.

Local mechanical stimuli correlate with tissue growth in axolotl salamander joint morphogenesis.

Movement-induced forces are critical to correct joint formation, but it is unclear how cells sense and respond to these mechanical cues. To study the role of mechanical stimuli in the shaping of the joint, we combined experiments on regenerating axolotl (Ambystoma mexicanum) forelimbs with a poroelastic model of bone rudiment growth. Animals either regrew forelimbs normally (control) or were injected with a transient receptor potential vanilloid 4 (TRPV4) agonist during joint morphogenesis. We quantified growth and shape in regrown humeri from whole-mount light sheet fluorescence images of the regenerated limbs. Results revealed significant differences in morphology and cell proliferation between groups, indicating that TRPV4 desensitization has an effect on joint shape. To link TRPV4 desensitization with impaired mechanosensitivity, we developed a finite element model of a regenerating humerus. Local tissue growth was the sum of a biological contribution proportional to chondrocyte density, which was constant, and a mechanical contribution proportional to fluid pressure. Computational predictions of growth agreed with experimental outcomes of joint shape, suggesting that interstitial pressure driven from cyclic mechanical stimuli promotes local tissue growth. Predictive computational models informed by experimental findings allow us to explore potential physical mechanisms involved in tissue growth to advance our understanding of the mechanobiology of joint morphogenesis.

How does bone microanatomy and musculature covary? An investigation in the forelimb of two species of martens (Martes foina, Martes martes).

The long bones and associated musculature play a prominent role in the support and movement of the body and are expected to reflect the associated mechanical demands. But in addition to the functional response to adaptive changes, the conjoined effects of phylogenetic, structural and developmental constraints also shape the animal's body. In order to minimise the effect of the aforementioned constraints and to reveal the biomechanical adaptations in the musculoskeletal system to locomotor mode, we here study the forelimb of two closely related martens: the arboreal pine marten (Martes martes) and the more terrestrial stone marten (Martes foina), focusing on their forelimb muscle anatomy and long bone microanatomy; and, especially, on their covariation. To do so, we quantified muscle data and bone microanatomical parameters and created 3D and 2D maps of the cortical thickness distribution for the three long bones of the forelimb. We then analysed the covariation of muscle and bone data, both qualitatively and quantitatively. Our results reveal that species-specific muscular adaptations are not clearly reflected in the microanatomy of the bones. Yet, we observe a global thickening of the bone cortex in the radius and ulna of the more arboreal pine marten, as well a stronger flexor muscle inserting on its elbow. We attribute these differences to variation in their locomotor modes. Analyses of our 2D maps revealed a shift of cortical thickness distribution pattern linked to ontogeny, rather than species-specific patterns. We found that although intraspecific variation is not negligible, species distinction was possible when taking muscular and bone microanatomical data into consideration. Results of our covariation analyses suggest that the muscle-bone correlation is linked to ontogeny rather than to muscular strength at zones of insertion. Indeed, if we find a correlation between cortical thickness distribution and the strength of some muscles in the humerus, that is not the case for the others and in the radius and ulna. Cortical thickness distribution appears rather linked to bone contact zones and ligament insertions in the radius and ulna, and to some extent in the humerus. We conclude that inference on muscle from bone microanatomy is possible only for certain muscles in the humerus.

Embryonic muscle splitting patterns reveal homologies of amniote forelimb muscles.

Limb muscles are remarkably complex and evolutionarily labile. Although their anatomy is of great interest for studies of the evolution of form and function, their homologies among major amniote clades have remained obscure. Studies of adult musculature are inconclusive owing to the highly derived morphology of modern amniote limbs but correspondences become increasingly evident earlier in ontogeny. We followed the embryonic development of forelimb musculature in representatives of six major amniote clades and found, contrary to current consensus, that these early splitting patterns are highly conserved across Amniota. Muscle mass cleavage patterns and topology are highly conserved in reptiles including birds, irrespective of their skeletal modifications: the avian flight apparatus results from slight early topological modifications that are exaggerated during ontogeny. Therian mammals, while conservative in their cleavage patterns, depart drastically from the ancestral amniote musculoskeletal organization in terms of topology. These topological changes occur through extension, translocation and displacement of muscle groups later in development. Overall, the simplicity underlying the apparent complexity of forelimb muscle development allows us to resolve conflicting hypotheses about homology and to trace the history of each individual forelimb muscle throughout the amniote radiations.

Muscular anatomy of the forelimb of tiger (Panthera tigris).

Dissection reports of large cats (family Felidae) have been published since the late 19th century. These reports generally describe the findings in words, show drawings of the dissection, and usually include some masses of muscles, but often neglect to provide muscle maps showing the precise location of bony origins and insertions. Although these early reports can be highly useful, the absence of visual depictions of muscle attachment sites makes it difficult to compare muscle origins and insertions in living taxa and especially to reconstruct muscle attachments in fossil taxa. Recently, more muscle maps have been published in the primary literature, but those for large cats are still limited. Here, we describe the muscular anatomy of the forelimb of the tiger (Panthera tigris), and compare muscle origins, insertions, and relative muscle masses to other felids to identify differences that may reflect functional adaptations. Our results reiterate the conservative nature of felid anatomy across body sizes and behavioral categories. We find that pantherines have relatively smaller shoulder muscle masses, and relatively larger muscles of the caudal brachium, pronators, and supinators than felines. The muscular anatomy of the tiger shows several modifications that may reflect an adaptation to terrestrial locomotion and a preference for large prey. These include in general a relatively large m. supraspinatus (shoulder flexion), an expanded origin for m. triceps brachii caput longum, and relatively large m. triceps brachii caput laterale (elbow extension), as well as relatively large mm. brachioradialis, abductor digiti I longus, and abductor digiti V. Muscle groups that are well developed in scansorial taxa are not well developed in the tiger, including muscles of the cranial compartment of the brachium and antebrachium, and m. anconeus. Overall, the musculature of the tiger strongly resembles that of the lion (Panthera leo), another large-bodied terrestrial large-prey specialist.

Comparative development of limb musculature in phylogenetically and ecologically divergent lizards.

BACKGROUND: Squamate reptiles (lizards, snakes, and amphisbaenians) exhibit incredible diversity in their locomotion, behavior, morphology, and ecological breadth. Although they often are used as models of locomotor diversity, surprisingly little attention has been given to muscle development in squamate reptiles. In fact, the most detailed examination was conducted almost 80 years ago and solely focused on the proximal limb regions. Herein, we present forelimb and hindlimb muscle morphogenesis data for three lizard species with different locomotion and feeding strategies: the desert grassland whiptail lizard, the central bearded dragon, and the veiled chameleon. This study fills critical gaps in our understanding of muscle morphogenesis in squamate reptiles and presents a comparative and temporospatial analysis of muscle development. RESULTS: Our results reveal a conserved pattern of early muscle development among lizards with different adult morphologies and ecologies. The variations that exist are concentrated in distal regions, particularly the specialized autopodia of chameleons, where differentiation of muscles associated with the digits is delayed. CONCLUSIONS: The chameleon autopod provides an example of major evolutionary modifications to the skeleton with only minor disruption of the conserved order and pattern of limb muscle development. This robustness of muscle patterning facilitates the evolution of extreme yet functional phenotypes.