The feeding apparatus of Rhea americana (Aves, Palaeognathae): Jaw myology and ontogenetic allometry.

In birds, the jaw musculature is a crucial adaptive feature involved in feeding. The morphological traits and postnatal growth patterns of jaw muscles constitute a useful proxy to interpret feeding function and ecology. This study aims to describe the jaw muscles of Rhea americana and explore their postnatal growth pattern. A total of 20 specimens of R. americana representing four ontogenetic stages were studied. Jaw muscles were described, weighed and their proportions with respect to body mass were calculated. Linear regression analysis was used to characterize ontogenetic scaling patterns. The morphological patterns of jaw muscles were characterized by their simplicity: bellies with few or no subdivisions and similar to those described for other flightless paleognathous birds. In all stages, the muscles pterygoideus lateralis, depressor mandibulae, and pseudotemporalis had the greatest mass values. The proportion of total jaw muscle mass decreased with age from 0.22% in 1-month-old chicks to 0.05% in adults. Linear regression analysis showed that all muscles scaled with negative allometry with respect to body mass. The progressive decrease of jaw muscle mass relative to body mass in adults could be related to the generation of less force, which is in accordance with the herbivorous diet of adults. In contrast, the diet of rhea chicks includes a large proportion of insects thus, this greater muscle proportion could be associated with the ability to generate more force, thus providing better abilities to grasp and hold more mobile prey.

Muscle architecture of the hindlimb of Tyto furcata (Aves Strigiformes): Highlights in owl morphology.

The American barn owl is a nocturnal bird of prey in which hind limb movements are a key factor in obtaining food; however, the architectural properties of its hind limb muscles have not been studied. This study sought to identify functional trends in the Tyto furcata hindlimb muscles by studying muscular architecture. The architectural parameters of the selected hip, knee, ankle, and digit muscles were studied in three specimens of the Tyto furcata and joint muscular proportions with an additional dataset were calculated. Previously published information on Asio otus was used for comparison. The flexor muscles of the digits had the highest muscle mass. Regarding architectural parameters, the main flexor of the digits (flexor digitorum longus) and the muscles that extend the knee and ankle joints (femorotibialis and gastrocnemius) showed a high physiological cross-sectional area (PCSA) and short fibers, allowing powerful digit flexion and knee and ankle extension. These mentioned features are in accordance with hunting behavior, in which prey capture is not only closely linked to the flexion of the digits but also to the movements of the ankle. During hunting, the distal hind limb is flexed and then fully extended at the moment of contact with the prey, whereas the digits are close to grasping the prey. The hip muscles showed a predominance of extensors over flexors, which were more massive, with parallel fibers and without tendons or short fibers. These features lead to a higher capacity to generate velocity to the detriment of forces, as indicated by the high values of the architectural index, their relatively low PCSA, and short or intermediate fiber length, which enhance the control of the joint positions and muscle length. Compared to Asio otus, Tyto furcata showed longer fibers; however, the relationship between fiber length and PCSA was similar for both species.

Wing and tail myology of Tyto furcata (Aves, Tytonidae).

Barn Owls (Tytonidae) are nocturnal raptors with the largest geographical distribution among Strigiformes. Several osteological, morphometrical, and biomechanical studies of this species were performed by previous authors. Nevertheless, the myology of forelimb and tail of the Barn Owls is virtually unknown. This study is the first detailed myological study performed on the wing and tail of the American Barn Owl (Tyto furcata). A total of 11 specimens were dissected and their morphology and muscle masses were described. Although T. furcata has the wing and tail myological pattern present in other species of Strigiformes, some peculiarities were observed including a difference in the attachment of m. pectoralis propatagialis due to the lack of the os prominence, and the presence of an osseous arch in the radius that seems to widen the anchorage area of the mm. pronator profundus, extensor longus alulae, and extensor longus digiti majoris. Furthermore, the m. biceps brachii has an unusual extra belly that flexes the forearm. The interosseous muscles have a small size and lacks ossified tendons. This feature may be indicative of a lower specialization in the elevation and flexion of the digiti majoris. Forelimb and tail muscle mass account for 10.66 and 0.24% of the total body mass, respectively. Forelimb muscle mass value is similar to the nocturnal (Strigiformes) and diurnal (Falconidae and Accipitridae) raptors, while the tail value is lower than in the diurnal raptors (Falconidae and Accipitridae). The myological differences with other birds of prey are here interpreted in association with their "parachuting" hunting style. This work complements our knowledge of the axial musculature of the American Barn owls, and provides important information for future studies related to functional morphology and ecomorphology.

Ontogenetic shape changes in the pelvis of the Greater Rhea (Aves, Palaeognathae) and their relationships with cursorial locomotion: a geometric morphometric approach.

Knowledge of the ontogenetic pattern of morphological features is essential to improve biological interpretations. The study of morphological features of the pelvic girdle and hind limb apparatus throughout growth is an excellent approach to understand how the skeletal morphology and muscles are interrelated during growth in a bird with a specialized mode of locomotion. The Greater Rhea (Rhea americana) is a large cursorial palaeognathous bird with long legs and powerful musculature. The postnatal shape changes of the pelvis of this bird were studied with geometric morphometric techniques, using landmarks and semilandmarks. In addition, regression analyses were used to explore the association between pelvic shape changes with muscle and body mass. The pelvises of 16 specimens of Rhea americana from 1 month old to adulthood were studied in dorsal and lateral views. Noticeable differences in pelvic shape were noted between ages, particularly in lateral view. In young birds, the pre- and post-acetabular ilium was subequal in length, whereas in adults the pre-acetabular ilium became shorter. In dorsal view, the main shape changes observed were the progressive thinning of both ilium portions and the elongation of the vertex craniolateralis ilii from chicks to adulthood. In this view, the only clear differentiation was between young and adult birds. Shape differences were influenced by body mass and pelvic muscles; the post-acetabular muscle mass explained the highest percentage of the variation. The specialized locomotion of Greater Rhea is reflected in their pelvic musculoskeletal system, in which the change to a longer post-acetabular ilium correlates with the growth of the powerful post-acetabular muscles. The actions of these muscles provide the necessary strength to support the body mass, minimize the body swinging movements and propel the body forward during locomotion. Bone morphology is affected by the forces produced by body mass and the muscle activity, demonstrating the presence of common growth mechanisms, which are primordial and gave rise to a functional and properly proportioned adult.

Hindlimb bone maturation during postnatal life in the Greater Rhea (Rhea americana, Aves, Palaeognathae): Implications for palaeobiological and zooarchaeological interpretations.

The objective of this study was to study the morphological pattern of bone maturation of the hindlimb bones of Rhea americana and find out how it can affect bone morphology after a taphonomic process. Juvenile specimens (n = 10) ranging from one month old to eight months old were studied. For comparison, bones from adults and juveniles from museum specimens (n = 4 and n = 6, respectively) were studied. In fresh bones, ossification centres were identified in the proximal and distal epiphyses of the tibiotarsi and in the proximal epiphysis of the tarsometatarsi, whereas the distal region of the femora and tarsometatarsi showed abundance of cartilage. The development and extension of the ossification centres of the tibiotarsi were different. In the proximal epiphysis, the centre presented less development with respect to the distal epiphysis. In the dry tibiotarsi, the proximal centre was absent and the distal one was well preserved. Both the fresh and dry juvenile tarsometatarsi presented unfused metatarsals and bulkier and wider aspect than the adult ones. The dry femora presented a noticeable excavation between condyles, whereas the dry tarsometatarsi showed the absence of the proximal epiphysis. The femora, tibiotarsi and tarsometatarsi possessed different traits of immaturity, which differentially affect the morphology of the preserved bones during a taphonomic process.

Hindlimb myology of the monk parakeet (Aves, Psittaciformes).

We studied the hindlimb myology of the monk parakeet (Myiopsitta monachus). Like all parrots, it has zygodactyl feet enabling perching, climbing, hanging, moving easily among trees, and handling food. Muscles were described and weighed, and physiological cross-sectional area (PCSA) of four flexors and one extensor was calculated. In comparison to other muscles, the M. tibialis cranialis and the M. fibularis brevis show increased development and high PCSA values, and therefore, large potential force production. Also, a large proportion of muscle mass was involved in flexing the digits. We hypothesize that these muscle traits are associated with the arboreal locomotion and food manipulation habits. In the monk parakeet, the M. extensor digitorum longus sends a branch to the hallux, and the connection between the M. flexor digitorum longus and the M. flexor hallucis longus is type I (Gadow's classification). We reaffirm the presence of the M. ambiens as a plesiomorphic condition that disappears in most members of the order. Among Psittaciformes, the M. fibularis brevis is stronger and the M. fibularis weaker in arboreal species than in basal terrestrial ones (e.g., Strigops).

Geometric morphometrics of the skull of Tinamidae (Aves, Palaeognathae).

The Tinamidae comprise exclusively Neotropical palaegnathous birds, with homogeneous body morphology and no sexual dimorphism. The goal of this work was to explore the variation in skull morphology between taxa and its possible correspondence with features such as diet or gender using geometric morphometric tools. Eleven landmarks were analyzed in 53 skulls of 4 genera that inhabit grasslands: Nothoprocta, Eudromia, Nothura and Rhynchotus. Intrageneric and intergeneric variability was analyzed. The genera studied here can be distinguished based on the geometric shape of their skull, with prenarial region length and neurocranium shape as the most outstanding features. In the genus Eudromia, males and females could be differentiated, while in the genus Nothoprocta, the species differentiated according to their trophic habits. This study allows establishing that genera and, in some cases, the gender of the Tinamidae can be differentiated based on cranial shape.

Virtual reconstructions of the endocranial cavity of Rhea americana (Aves, Palaeognathae): postnatal anatomical changes.

We examined the external anatomy of the endocast of the Greater Rhea (Rhea americana, Palaeognathae), during 3 main stages of its postnatal life, and compared it with information available on other palaeognathous birds. Series of scans with spiral computed tomographies were obtained from 3 skulls of different ages (chick, juvenile and adult) of R. americana; digital 3-dimensional reconstruction was performed and brain volumes were calculated from the models obtained. Qualitative assessment of the brain anatomy of R. americana indicates a conical and ventrally located bulbus olfactorius, laterally expanded hemispherium telencephali, well-developed eminentia sagittalis, and conspicuous cerebellum and tectum mesencephali. Anatomy of the chick brain was markedly different: less lateral expansion of the hemispherium telencephali, and lesser development of the eminentia sagittalis and auricula cerebelli. Little change between chicks and adults was observed in some brain regions such as the tectum mesencephali, while the eminentia sagittalis showed great increase in size. The large size of the eminentia sagittalis coupled with its increasing development during ontogeny could reflect its importance for visual processing functions and the way these improve during growth. Finally, the brain of R. americana is similar to that of Struthio and Dromaius, but differs from that of the Tinamidae and of Apteryx, allowing recognition of 3 distinct brain morphologies among the Palaeognathae.

Functional anatomy of the limbs of erethizontidae (Rodentia, Caviomorpha): Indicators of locomotor behavior in Miocene porcupines.

Functional analysis of the limb bones of the erethizontid Steiromys duplicatus, one of the most abundant Miocene porcupines from Patagonia, provides evidence to infer their locomotor behavior. Remains of the giant Neosteiromys pattoni (Late Miocene of Northeast Argentina) are also analyzed. Osteological and myological features of extant porcupines were evaluated and used as a model to interpret the functional significance of Miocene species' limbs. Several features in erethizontids are compatible with the ability to climb: the low humeral tuberosities indicate a mobile gleno-humeral joint; the prominent and distally extended deltopectoral crest indicates a powerful pectoral muscle, which is particularly active when climbing; the humero-ulnar and humero-radial joints are indicative of pronation-supination movements; the well-developed lateral epicondylar ridge and the medially protruding entepicondyle are in agreement with an important development of the brachioradialis, supinator, flexor digitorum profundus, and pronator teres muscles, acting in climbing and grasping functions; the mechanical advantage of the biceps brachii would be emphasized because of its distal attachment on the bicipital tuberosity. As with extant porcupines, in Miocene species, the large femoral head would have permitted a broad range of abduction of the femur, and the medially protruding lesser trochanter would have emphasized the abduction and outward rotation of the femur by the action of the ilio-psoas complex. In S. duplicatus, the shape of the hip, knee, and cruro-astragalar, calcaneo-astragalar, and astragalo-navicular joints would have allowed lateral and rotational movements, although probably to a lesser degree than in extant porcupines. Foot features of S. duplicatus (e.g., great medial sesamoid bone, medial astragalar head, complete hallux) indicate that this species would have had grasping ability, but would not have achieved the high degree of specialization of Coendou. Steiromys duplicatus would have been a semiarboreal dweller, resembling Erethizon dorsatum.