Running, jumping, hunting, and scavenging: Functional analysis of vertebral mobility and backbone properties in carnivorans.

Carnivorans are well-known for their exceptional backbone mobility, which enables them to excel in fast running and long jumping, leading to them being among the most successful predators amongst terrestrial mammals. This study presents the first large-scale analysis of mobility throughout the presacral region of the vertebral column in carnivorans. The study covers representatives of 6 families, 24 genera and 34 species. We utilized a previously developed osteometry-based method to calculate available range of motion, quantifying all three directions of intervertebral mobility: sagittal bending (SB), lateral bending (LB), and axial rotation (AR). We observed a strong phylogenetic signal in the structural basis of the vertebral column (vertebral and joint formulae, length proportions of the backbone modules) and an insignificant phylogenetic signal in most characteristics of intervertebral mobility. This indicates that within the existing structure (stabilization of which occurred rather early in different phylogenetic lineages), intervertebral mobility in carnivorans is quite flexible. Our findings reveal that hyenas and canids, which use their jaws to seize prey, are characterized by a noticeably elongated cervical region and significantly higher SB and LB mobility of the cervical joints compared to other carnivorans. In representatives of other carnivoran families, the cervical region is very short, but the flexibility of the neck (both SB and LB) is significantly higher than that of short-necked odd-toed and even-toed ungulates. The lumbar region of the backbone in carnivorans is dorsomobile in the sagittal plane, being on average ~23° more mobile than in artiodactyls and ~38° more mobile than in perissodactyls. However, despite the general dorsomobility, only some representatives of Canidae, Felidae, and Viverridae are superior in lumbar flexibility to the most dorsomobile ungulates. The most dorsomobile artiodactyls are equal or even superior to carnivorans in their ability to engage in dorsal extension during galloping. In contrast, carnivorans are far superior to ungulates in their ability to engage ventral flexion. The cumulative SB in the lumbar region in carnivorans largely depends on the mode of running and hunting. Thus, adaptation to prolonged and enduring pursuit of prey in hyenas is accompanied by markedly reduced SB flexibility in the lumbar region. A more dorsostable run is also a characteristic of the Ursidae, and the peculiar maned wolf. Representatives of Felidae and Canidae have significantly more available SB mobility in the lumbar region. However, they fully engage it only occasionally at key moments of the hunt associated with the direct capture of the prey or when running in a straight line at maximum speed.

Morphological peculiarities in the integument of enigmatic anomalurid gliders (Anomaluridae, Rodentia).

Scaly-tailed squirrels, the most poorly known group of gliding mammals, hold the record for variety of remarkable integument peculiarities. One of the most striking of these features is the scales on the tail, which apparently allow them to reduce energy costs when positioning themselves on a tree trunk. No less interesting is a peculiar spur that supports the flying membrane: the unciform element ('spur'). Despite the peculiarity of such elements, their nature has not yet been studied. Using anatomical, histological methods and scanning electron microscopy we studied the structure of the skin and its derivatives in five of the six species from both genera of extant gliding scaly-tailed squirrels (Anomaluridae, Rodentia): Idiurus macrotis, Idiurus zenkeri, Anomalurus beecrofti, Anomalurus pusillus and Anomalurus derbianus. In addition to the common mammalian skin structures, such as hair, vibrissae, sebaceous glands, meibomian glands of eyelids and eccrine sweat glands of the palmar and plantar pads, these animals have unique species-specific skin derivatives (the tail scaly organ and its specific glands, vibrissae of the withers, patagium and its hair brush) that play a significant role in their adaptation to gliding and to their environment in general. The structure of the elbow spur is also described and hypotheses on its evolutionary origin from the tendon of the triceps muscle are presented.

Anatomy of the forelimb musculature and ligaments of Psittacus erithacus (Aves: Psittaciformes).

Parrots (order Psittaciformes) are a rather homogeneous group of birds that can be easily distinguished by the notably modified morphology of the skull and hindlimb. Detailed description of the forelimb morphology in these birds has never been provided, though parrots are often used as model objects in flight studies. Parrots are also considered the closest living relatives of the perching birds (Passeriformes), and thus knowledge of the wing morphology in Psittaciformes is important for understanding the evolution of the locomotor apparatus on the way to the most speciose group of birds. Here we provide a comprehensive illustrated description of the wing morphology (musculature and ligaments) of the African grey parrot (Psittacus erithacus) and compare it with several closely related taxa of the high clade Eufalconimorphae and more distantly related outgroups (based on personal dissections and literature data). We note a general similarity of the wing musculature between P. erithacus and Falconidae. A number of features common with the outgroup Columbidae supports a generally plesiomorphic structure of the forelimb in parrots as compared with the Passeriformes. Nevertheless, the wing of the Psittaciformes displays a series of structural (likely autapomorphic) modifications, which can be explained in terms of adaptations for flight with vertical body. An analysis of the anatomical data for parrots (ratio of wing elevators and highly unusual development of the M. supracoracoideus), which is based on the current experiment-based knowledge of the flapping flight in birds, allows us to hypothesize that parrots are able to produce useful aerodynamic force during the upstroke, which is also known for pigeons and hummingbirds. This supposed ability of vertical flight and the zygodactyl foot together link the origin of parrots with the dense (likely tropical) forests.

Structural Diversity of the Extensor Digitorum Profundus Muscle Complex in Platyrrhini.

Separate extension of fingers in the hand of primates is performed by 3 muscles: m. extensor pollicis longus, m. extensor digiti secundi, and m. extensor digitorum lateralis. Here it is proposed to consider them as parts of the extensor digitorum profundus muscular complex. The diversity in structure of these muscles in primates is examined based both on original anatomical study of New World monkeys and analysis of extensive published data on primates from different taxonomic groups. It is shown that in these muscles there are 2 main types of structure variations - the division of the muscle belly into several heads which give rise to separate tendons, and the split of the single terminal tendon into several branches. The first type of modification ensures the possibility of a separate management of the fingers, and the second, on the contrary, ensures the coupled control of extension of fingers. A scheme of evolutionary transformations of muscles belonging to the complex of the deep extensors of fingers is proposed.

Are owls technically capable of making a full head turn?

The three-dimensional configuration of the neck that produces extreme head turn in owls was studied using the Joint Coordinate System. The limits of planar axial rotation (AR), lateral, and sagittal bending in each vertebral joint were measured. They are not extraordinary among birds, except probably for the extended ability for AR. The vertebral joint angles involved in the 360° head turn do not generally exceed the limits of planar mobility. Rotation in one plane does not expand the range of motion in the other, with one probable exception being extended dorsal bending in the middle of the neck. Therefore, the extreme 360° head turn can be presented as a simple combination of the three planar motions in the neck joints. Surprisingly, certain joints are always laterally bent or axially rotated to the opposite side than the head was turned. This allows keeping the anterior part of the neck parallel to the thoracic spine, which probably helps preserve the ability for peering head motions throughout the full head turn. The potential ability of one-joint muscles of the owl neck, the mm. intertransversarii, to ensure the 360° head turn was addressed. It was shown that the 360° head turn does not require these muscles to shorten beyond the known contraction limit of striated vertebrate muscles. Shortening by 50% or less is enough for the mm. intertransversarii in the middle neck region for the 360° head turn. This study has broad implications for further research on vertebral mobility and function in a variety of tetrapods, providing a new method for CT scan-based measurement of intervertebral angles.

Running of the feathertail glider (Acrobates pygmaeus) on level ground: Kinematics.

A complete kinematic analysis of the trunk and limbs was performed for the smallest gliding mammal, the feathertail glider (Acrobates pygmaeus). To compare the running technique of the glider with that of terrestrial mammals, only movement on horizontal substrate was studied. Kinematic analysis was performed separately for bound with extended suspension and bound with both suspensions. Terrestrial locomotion of the feathertail glider was classified as primitive ricochet, which is distinguished from true gallop by the absence of a delay in protraction of the hindlimbs after their take-off. The kinematic analysis showed significant impacts of gliding adaptation on the quadrupedal running, the most striking of which are the extremely sprawling position of the limbs, in particular, the wide stance of the forelimbs, and consequently, the absence of crossing of the fore- and hindlimbs in gathered stage of the cycle. The degree of limbs' sprawling in the feathertail glider is closer to the reptilian condition than to the mammalian parasagittal state. The sprawling condition of the feathertail glider is secondary and therefore is designated as "deparasagittalization."

Running of the feathertail glider (Acrobates pygmaeus) on level ground: Gaits.

Gliding is a crucial adaptation to arboreal habitats in several groups of mammals. Along with certain advantages it imposes limitations on the quadrupedal running since it affects the locomotor apparatus. To estimate the impact on quadrupedal running in gliders, the feathertail glider (Acrobates pygmaeus) was chosen considering that the small size allows minor morphological modifications for aerial locomotion. The gaits were studied on horizontal flat substrate which made it possible to compare running technique of the glider with that of terrestrial mammals. In all analyzed plots the footfall sequence was found to be asymmetrical; in most cases the bound was used, in contrast, the gallop occurred only occasionally. The half-bound with the fore lead, the most usual asymmetrical gait in quadrupedal marsupials, was much less common in A. pygmaeus than the bound; the rare among mammals half-bound with the hind lead was also found. The bound was not only the most common gait but also the steadiest one; therefore we can conclude that A. pygmaeus uses all other asymmetrical gaits as transitional forms associated with changes in speed, direction, etc. The bound with extended suspension is probably preferred by A. pygmaeus because it most closely resembles gliding by posture.

Gait mechanics of a blind echolocating rodent: Implications for the locomotion of small arboreal mammals and proto-bats.

Arboreal mammals have evolved a range of biomechanical adaptations that allow them to navigate trees effectively. One such feature that has received considerable attention is the importance of vision that helps arboreal animals assess gap distances, assure proper foot placement, and inspect potential risks. While there is considerable debate about the relative importance of the visual system specifics, there is little doubt that the ability to at least see the environment must confer some level of safety when navigating arboreal substrates. In this study, we explore spatiotemporal and kinematic patterns of arboreal locomotion in the Vietnamese pygmy dormouse (Typhlomys chapensis), a blind rodent that uses ultrasonic echolocation to navigate in tree canopies. We compare these data with five other species of arboreal rodents and primates. Spatiotemporal gait characteristics are largely similar between the Vietnamese pygmy dormouse and other small-bodied arboreal species analyzed. Most notable is the tendency for relatively high-speed asymmetrical gaits on large-diameter substrates and slower symmetrical lateral-sequence gaits on small-diameter substrates. Furthermore, for all species speed is primarily regulated by increasing stride frequency rather than length. Kinematics of the Vietnamese pygmy dormouse changed little in response substrate size and were primarily driven by speed. These findings suggest that the information gathered during ultrasonic scanning is sufficient to allow effective quadrupedal locomotion while moving on arboreal supports. The Vietnamese pygmy dormouse may serve as a model for the quadrupedal nocturnal ancestor of bats, which had started developing ultrasonic echolocation and reducing vision while likely occupying an arboreal niche.

A blind climber: The first evidence of ultrasonic echolocation in arboreal mammals.

The means of orientation is studied in the Vietnamese pygmy dormouse Typhlomys chapensis, a poorly known enigmatic semi-fossorial semi-arboreal rodent. Data on eye structure are presented, which prove that Typhlomys (translated as "the blind mouse") is incapable of object vision: the retina is folded and retains no more than 2500 ganglion cells in the focal plane, and the optic nerve is subject to gliosis. Hence, Typhlomys has no other means for rapid long-range orientation among tree branches other than echolocation. Ultrasonic vocalization recordings at the frequency range of 50-100 kHz support this hypothesis. The vocalizations are represented by bouts of up to 7 more or less evenly-spaced and uniform frequency-modulated sweep-like pulses in rapid succession. Structurally, these sweeps are similar to frequency-modulated ultrasonic echolocation calls of some bat species, but they are too faint to be revealed with a common bat detector. When recording video simultaneously with the ultrasonic audio, a significantly greater pulse rate during locomotion compared to that of resting animals has been demonstrated. Our findings of locomotion-associated ultrasonic vocalization in a fast-climbing but weakly-sighted small mammal ecotype add support to the "echolocation-first theory" of pre-flight origin of echolocation in bats.