Abdominal muscle function in ventilation and locomotion in new world opossums and basal eutherians: Breathing and running with and without epipubic bones

All tetrapods have the same four basic abdominal hypaxial muscle layers that wrap around the abdomen between the pelvis, ribcage, and spine. However, the marsupials and our immediate mammalian ancestors have epipubic bones extending anteriorly into the ventral hypaxial layers with two additional muscles connecting them to the ventral midline and femur. Studies of two marsupials have shown that all of the abdominal hypaxials play a part bilaterally in resting ventilation and during locomotion there is an asymmetrical pattern of activity as the hypaxial muscles form a cross‐couplet linkage that uses the epipubic bone as a lever to provide long‐axis support of the body between diagonal limb couplets during each step. The cross‐couplet epipubic lever system defines the earliest mammals and is lost in placental mammals. To expand our understanding of the evolution of mammalian abdominal muscle function and loco‐ventilatory integration we tested the generality of the cross‐couplet system in marsupials and conducted the first formal studies of hypaxial abdominal motor patterns in generalized placental mammals focusing on a representative rodent and insectivore. These new data reveal 1) that continuous abdominal muscle tonus during resting ventilation and a 1:1 breath to step cycle during locomotion appear to be the basal condition for mammals, 2) that the loss of epipubic bones in eutherians is associated with a shift from the cross‐couplet dominated motor pattern of marsupials to a shoulder‐to‐pelvis system with unilateral activation of abdominal muscles during locomotion and 3) that hypaxial function in generalized eutherians is more similar to marsupials than cursorial mammals. J. Morphol. 2009. © 2009 Wiley‐Liss, Inc.     

Abdominal muscle and epipubic bone function during locomotion in Australian possums: Insights to basal mammalian conditions and eutherian‐like tendencies in Trichosurus

Mammals have four hypaxial muscle layers that wrap around the abdomen between the pelvis, ribcage, and spine. However, the marsupials have epipubic bones extending anteriorly into the ventral hypaxial layers with two additional muscles extending to the ventral midline and femur. Comparisons of South American marsupials to basal eutherians have shown that all of the abdominal hypaxials are active bilaterally in resting ventilation. However, during locomotion marsupials employ an asymmetrical pattern of activity as the hypaxial muscles form a crosscouplet linkage that uses the epipubic bone as a lever to provide long‐axis support of the body between diagonal limb couplets during each step. In basal eutherians, this system shifts off the femur and epipubic bones (which are lost) resulting in a shoulder to pelvis linkage associated with shifts in both the positions and activity patterns of the pectineus and rectus abdominis muscles during locomotion. In this study, we present data on hypaxial function in two species (Pseudocheirus peregrinus and Trichosurus vulpecula) representing the two major radiations of possums in Australia: the Pseudocheiridae (within the Petauroidea) and the Phalangeridae. Patterns of gait, motor activity, and morphology in these two Australian species were compared with previous work to examine the generality of 1) the crosscouplet lever system as the basal condition for the Marsupialia and 2) several traits hypothesized to be common to all mammals (hypaxial tonus during resting ventilation, ventilation to step synchrony during locomotion, and bilateral transversus abdominis activity during locomotor expiration). Our results validate the presence of the crosscouplet pattern and basic epipubic bone lever system in Australian possums and confirm the generality of basal mammalian patterns. However, several novelties discovered in Trichosurus, reveal that it exhibits an evolutionary transition to intermediate eutherian‐like morphological and motor patterns paralleling many other unique features of this species. J. Morphol., 2010. © 2009 Wiley‐Liss, Inc.     

Molar scaling in strepsirrhine primates

We examined how maxillary molar dimensions change with body and skull size estimates among 54 species of living and subfossil strepsirrhine primates. Strepsirrhine maxillary molar areas tend to scale with negative allometry, or possibly isometry, relative to body mass. This observation supports several previous scaling analyses showing that primate molar areas scale at or slightly below geometric similarity relative to body mass. Strepsirrhine molar areas do not change relative to body mass(0.75), as predicted by the metabolic scaling hypothesis. Relative to basicranial length, maxillary molar areas tend to scale with positive allometry. Previous claims that primate molar areas scale with positive allometry relative to body mass appear to rest on the incorrect assumption that skull dimensions scale isometrically with body mass. We identified specific factors that help us to better understand these observed scaling patterns. Lorisiform and lemuriform maxillary molar scaling patterns did not differ significantly, suggesting that the two infraorders had little independent influence on strepsirrhine scaling patterns. Contrary to many previous studies of primate dental allometry, we found little evidence for significant differences in molar area scaling patterns among frugivorous, folivorous, and insectivorous groups. We were able to distinguish folivorous species from frugivorous and insectivorous taxa by comparing M1 lengths and widths. Folivores tend to have a mesiodistally elongated M1 for a given buccolingual M1 width when compared to the other two dietary groups. It has recently been shown that brain mass has a strong influence on primate dental eruption rates. We extended this comparison to relative maxillary molar sizes, but found that brain mass appears to have little influence on the size of strepsirrhine molars. Alternatively, we observed a strong correlation between the relative size of the facial skull and relative molar areas among strepsirrhines. We hypothesize that this association may be underlain by a partial sharing of the patterning of development between molar and facial skull elements.     

Selective breeding as a tool to probe skeletal response to high voluntary locomotor activity in mice

We present a novel mouse-model for the study of skeletal structureand evolution, based on selective breeding for high levels ofvoluntary wheel running. Whereas traditional models (originallyinbred strains, more recently knockouts and transgenics) relyon the study of mutant or laboratory-manipulated phenotypes,we have studied changes in skeletal morphometrics resultingfrom many generations of artificial selection for high activityin the form of wheel running, in which mice engage voluntarily.Mice from the four replicate High Runner (HR) lines run nearlythree times as many revolutions during days 5 and 6 of a 6-dayexposure to wheels (1.12 m circumference). We have found significantchanges in skeletal dimensions of the hind limbs, includingdecreased directional asymmetry, larger femoral heads, and widerdistal femora. The latter two have been hypothesized as evolutionaryadaptations for long-distance locomotion in hominids. Exercise-trainingstudies involving experimental groups with and without accessto wheels have shown increased diameters of both femora andtibiafibulae, and suggest genetic effects on trainability (genotype-by-environmentinteractions). Reanalysis of previously published data on bonemasses of hind limbs revealed novel patterns of change in bonemass associated with access to wheels for 2 months. Withoutaccess to wheels, HR mice have significantly heavier tibiafibulaeand foot bones, whereas with chronic access to wheels, a significantincrease in foot bone mass that was linearly related to increasesin daily wheel running was observed. Mice exhibiting a recentlydiscovered small-muscle phenotype ("mini-muscle," [MM] causedby a Mendelian recessive gene), in which the mass of the tricepssurae muscle complex is 50% lower than in normal individuals,have significantly longer and thinner bones in the hind limb.We present new data for the ontogenetic development of musclemass in Control, HR, and MM phenotypes in mice of 1–7weeks postnatal age. Statistical comparisons reveal highly significantdifferences both in triceps surae mass and mass-corrected tricepssurae mass between normal and MM mice at all but the postnatalage of 1 week. Based on previously observed differences in distributionsof myosin isoforms in adult MM mice, we hypothesize that a reductionof myosin heavy-chain type-IIb isoforms with accounts for ourobserved ontogenetic changes in muscle mass.     

Dynamic strain similarity in vertebrates; an alternative to allometric limb bone scaling

Galileo (1638) observed that "nature cannot grow a tree nor construct an animal beyond a certain size, while retaining the proportions which suffice in the case of a smaller structure". However, subsequent measurement has shown that limb bone dimensions are scaled geometrically with body size (Alexander et al., 1979a), and that the material properties of their constituent bone tissue are similar in animals over a wide range of body weight (Sedlin & Hirsch, 1966; Yamada, 1970; Burstein et al., 1972; Biewener, 1982). If, as suggested in previous scaling arguments (McMahon, 1973; Biewener, 1982), vigorous locomotion involved the same proportional forces over a wide range of animal size, this would create a paradox since large animals would be in far greater danger of skeletal failure than small ones. However, in vivo strain gauge implantations have shown that, during high speed running, axial force as a proportion of body weight (G) in the limb bones of animals decreases as a function of body size from 6.9 G in a 7 kg turkey to 2.8 G in a small (130 kg) horse. Estimates of axial force in larger animals suggest that this is further reduced to 0.8 G in a 2500 kg elephant. Nevertheless, it appears that, regardless of animal size or locomotory style, the peak stresses in the bones of these animals are remarkably similar. Therefore, throughout the range of animals considered (350 times differences in mass), we suggest that similar safety factors to failure are maintained, not by allometrically scaling bone dimensions, but rather by allometrically scaling the magnitude of the peak forces applied to them during vigorous locomotion.(ABSTRACT TRUNCATED AT 250 WORDS)     

Turning Semicircular Canal Function on Its Head: Dinosaurs and a Novel Vestibular Analysis

Previous investigations have correlated vestibular function to locomotion in vertebrates by scaling semicircular duct radius of curvature to body mass. However, this method fails to discriminate bipedal from quadrupedal non-avian dinosaurs. Because they exhibit a broad range of relative head sizes, we use dinosaurs to test the hypothesis that semicircular ducts scale more closely with head size. Comparing the area enclosed by each semicircular canal to estimated body mass and to two different measures of head size, skull length and estimated head mass, reveals significant patterns that corroborate a connection between physical parameters of the head and semicircular canal morphology. Head mass more strongly correlates with anterior semicircular canal size than does body mass and statistically separates bipedal from quadrupedal taxa, with bipeds exhibiting relatively larger canals. This morphologic dichotomy likely reflects adaptations of the vestibular system to stability demands associated with terrestrial locomotion on two, versus four, feet. This new method has implications for reinterpreting previous studies and informing future studies on the connection between locomotion type and vestibular function.     

Morphological and biomechanical analysis of a skeleton from Roman imperial necropolis of Casalecchio di Reno (Bologna, Italy, II-III c. A. D.). A possible case of crutch use

A Roman skeleton (T.130) from the roman necropolis of Casalecchio di Reno has been studied in order to understand if the hypothesis of crutch use, suggested by the severe articular degeneration at the hip joint that caused evident reduction of his locomotory possibilities, could be supported by the morphological alterations of other bones and joints. The pathological changes and muscular development of the upper limbs and shoulder girdle bones suggest that these parts were submitted to a great mechanical stress. The observations are consistent with the hypothesis of crutch use that would have involved a new weight-bearing function of the upper limbs in order to help locomotion, even though it is difficult to assess the number and type of the crutches. The comparison with other possible cases of crutch use reported in literature gives an additional support to the interpretation of the findings.     

Evolution of posture in amniotes–Diving into the trabecular architecture of the femoral head

Extant amniotes show remarkable postural diversity. Broadly speaking, limbs with erect (strongly adducted, more vertically oriented) posture are found in mammals that are particularly heavy (graviportal) or show good running skills (cursorial), while crouched (highly flexed) limbs are found in taxa with more generalized locomotion. In Reptilia, crocodylians have a “semi-erect” (somewhat adducted) posture, birds have more crouched limbs and lepidosaurs have sprawling (well-abducted) limbs. Both synapsids and reptiles underwent a postural transition from sprawling to more erect limbs during the Mesozoic Era. In Reptilia, this postural change is prominent among archosauriforms in the Triassic Period. However, limb posture in many key Triassic taxa remains poorly known. In Synapsida, the chronology of this transition is less clear, and competing hypotheses exist. On land, the limb bones are subject to various stresses related to body support that partly shape their external and internal morphology. Indeed, bone trabeculae (lattice-like bony struts that form the spongy bone tissue) tend to orient themselves along lines of force. Here, we study the link between femoral posture and the femoral trabecular architecture using phylogenetic generalized least squares. We show that microanatomical parameters measured on bone cubes extracted from the femoral head of a sample of amniote femora depend strongly on body mass, but not on femoral posture or lifestyle. We reconstruct ancestral states of femoral posture and various microanatomical parameters to study the “sprawling-to-erect” transition in reptiles and synapsids, and obtain conflicting results. We tentatively infer femoral posture in several hypothetical ancestors using phylogenetic flexible discriminant analysis from maximum likelihood estimates of the microanatomical parameters. In general, the trabecular network of the femoral head is not a good indicator of femoral posture. However, ancestral state reconstruction methods hold great promise for advancing our understanding of the evolution of posture in amniotes.     

Trabecular bone structure in the primate wrist

Trabecular (or cancellous) bone has been shown to respond to mechanical loading throughout ontogeny and thus can provide unique insight into skeletal function and locomotion in comparative studies of living and fossil mammalian morphology. Trabecular bone of the hand may be particularly functionally informative because the hand has more direct contact with the substrate compared with the remainder of the forelimb during locomotion in quadrupedal mammals. This study investigates the trabecular structure within the wrist across a sample of haplorhine primates that vary in locomotor behaviour (and thus hand use) and body size. High‐resolution microtomographic scans were collected of the lunate, scaphoid, and capitate in 41 individuals and eight genera (Homo, Gorilla, Pan, Papio, Pongo, Symphalangus, Hylobates, and Ateles). We predicted that particular trabecular parameters would 1) vary across suspensory, quadrupedal, and bipedal primates based on differences in hand use and load, and 2) scale with carpal size following similar allometric patterns found previously in other skeletal elements across a larger sample of mammals and primates. Analyses of variance (trabecular parameters analysed separately) and principal component analyses (trabecular parameters analysed together) revealed no clear functional signal in the trabecular structure of any of the three wrist bones. Instead, there was a large degree of variation within suspensory and quadrupedal locomotor groups, as well as high intrageneric variation within some taxa, particularly Pongo and Gorilla. However, as predicted, Homo sapiens, which rarely use their hands for locomotion and weight support, were unique in showing lower relative bone volume (BV/TV) compared with all other taxa. Furthermore, parameters used to quantify trabecular structure within the wrist scale with size generally following similar allometric patterns found in trabeculae of other mammalian skeletal elements. We discuss the challenges associated with quantifying and interpreting trabecular bone within the wrist. J. Morphol. 275:572–585, 2014. © 2013 Wiley Periodicals, Inc.     

Scaling in the animal kingdom

Several of the known scaling laws in the animal kingdom are based on a so-called allometric correlation in which some physical quantity is presumed to scale as some power of the mass of the animal. Such a simple correlation, when deduced purely as an empirical result, often hides the physical balances that fix the relevant scaling law. In particular, the emphasis on a simple allometric scaling has often masked the fundamental role played by time scales associated with the physical balances being struck. In this paper I have concentrated on three different attributes to which the use of dimensional analysis, scaling arguments and some judicious guesswork have led to new results and an understanding of some balances that occur in the animal kingdom. The running speed of animals is examined and a rationale deduced for the resolution of a conundrum first posed by A.V. Hill of why it is that many animals appear to have approximately the same maximum speed. A complete dimensional analysis for scaling the basal metabolic rate for a class of animals suggests that a detailed understanding of the physical balances that fix the metabolic rate could be quite subtle. However, the use of such an analysis has led to the discovery of a new correlation for mammals, relating the metabolic rate to the mass and the pulse rate of the animal. At the heart of many scaling laws for animal motion is the provision of an estimate of how the skeletal structure depends on the mass of the animal. It has been known for some time that the assumption of isometry between the builds of animals is too constrictive to describe the observed scaling laws. It is shown here how to relax the isometric assumption and deduce scaling laws in good agreement with observation. Thus, it appears that the skeletal dimensions of many animals with exoskeletons are fixed by the need to support static rather than dynamical loads. The scaling laws associated with endoskeletons are more complex, apparently, though the analysis does suggest that it is dynamical loading which is decisive for the skeletal design of land mammals.     

不同生境蚶形无齿蚌的形态观察

对不同生境下蚶形无齿蚌壳的形态、育儿囊的类型和结构以及钩介幼虫等进行了比较研究。结果表明,在不同生境下,蚶形无齿蚌个体大小有很大差异,壳具有高度的可塑性;育儿囊由两片外鳃构成,为外鳃类的同生型,钩介幼虫在育儿囊内呈散乱状态存在;在小同区域,钩介幼虫的大小不同,但其壳高与壳长的比例却是一致的,且不同生境钩介幼虫的超微结构相同,均为有钩型。     

Effects of metabolic level on the body size scaling of metabolic rate in birds and mammals

Metabolic rate is traditionally assumed to scale with body mass to the 3/4-power, but significant deviations from the '3/4-power law' have been observed for several different taxa of animals and plants, and for different physiological states. The recently proposed 'metabolic-level boundaries hypothesis' represents one of the attempts to explain this variation. It predicts that the power (log-log slope) of metabolic scaling relationships should vary between 2/3 and 1, in a systematic way with metabolic level. Here, this hypothesis is tested using data from birds and mammals. As predicted, in both of these independently evolved endothermic taxa, the scaling slope approaches 1 at the lowest and highest metabolic levels (as observed during torpor and strenuous exercise, respectively), whereas it is near 2/3 at intermediate resting and cold-induced metabolic levels. Remarkably, both taxa show similar, approximately U-shaped relationships between the scaling slope and the metabolic (activity) level. These predictable patterns strongly support the view that variation of the scaling slope is not merely noise obscuring the signal of a universal scaling law, but rather is the result of multiple physical constraints whose relative influence depends on the metabolic state of the organisms being analysed.     

Allometry of cetacean forelimb bones

This study examines the allometric scaling relationships of the cetacean humerus, radius, and ulna. Bone lengths and diameters were measured for 20 species of odontocete and three species of mysticete cetaceans, representing eight of the nine extant cetacean families. The scaling of individual bone proportions (bone length vs. cranio-caudal diameter, bone length vs. dorso-ventral diameter), and of individual bone dimensions against estimated body mass, are compared to models of geometric and elastic similarity. The geometric similarity model describes the scaling relationship of bone length vs. cranio-caudal diameter and body mass vs. cranio-caudal diameter for the humerus only; geometric similarity also describes the scaling relationship of body mass vs. bone length for all three bones. None of the scaling relationships fits the elastic similarity model. The scaling relationships of bone length vs. dorso-ventral diameter for all three bones, and bone length vs. cranio-caudal diameter for the radius and ulna, exhibit negative allometry, indicating that large bones are less robust than small bones. Negative allometry of structural support elements has not been previously described for terrestrial mammals or plants. The high relative swimming speeds of small delphinids may generate sufficient stresses to require more robust bones relative to those of larger whales. © 1994 Wiley-Liss, Inc.     

Comparative limb bone scaling in turtles: Phylogenetic transitions with changes in functional demands?

Several terrestrial vertebrate clades include lineages that have evolved nearly exclusive use of aquatic habitats. In many cases, such transitions are associated with the evolution of flattened limbs that are used to swim via dorsoventral flapping. Such changes in shape may have been facilitated by changes in limb bone loading in novel aquatic environments. Studies on limb bone loading in turtles found that torsion is high relative to bending loads on land, but reduced compared to bending during aquatic rowing. Release from torsion among rowers could have facilitated the evolution of hydrodynamically advantageous flattened limbs among aquatic species. Because rowing is regarded as an intermediate locomotor stage between walking and flapping, rowing species might show limb bone flattening intermediate between the tubular shapes of walkers and the flattened shapes of flappers. We collected measurements of humeri and femora from specimens representing four functionally divergent turtle clades: sea turtles (marine flappers), softshells (specialized freshwater rowers), emydids (generalist semiaquatic rowers), and tortoises (terrestrial walkers). Patterns of limb bone scaling with size were compared across lineages using phylogenetic comparative methods. Although rowing taxa did not show the intermediate scaling patterns we predicted, our data provide other functional insights. For example, flattening of sea turtle humeri was associated with positive allometry (relative to body mass) for the limb bone diameter perpendicular to the flexion-extension plane of the elbow. Moreover, softshell limb bones exhibit positive allometry of femoral diameters relative to body mass, potentially helping them maintain their typical benthic position in water by providing additional weight to compensate for shell reduction. Tortoise limb bones showed positive allometry of diameters, as well as long humeri, relative to body mass, potentially reflecting specializations for resisting loads associated with digging. Overall, scaling patterns of many turtle lineages appear to correlate with distinctive behaviors or locomotor habits.     

Fabricational morphology of oblique ribs in bivalves

Most analyses on allometry of long bones in terrestrial mammals have focused on dimensional allometry, relating external bone measurements either to each other or to body mass. In this article, an analysis of long bone mass to body mass in 64 different species of mammals, spanning three orders of magnitude in body mass, is presented. As previously reported from analyses on total skeletal mass to body mass in terrestrial vertebrates, the masses of most appendicular bones scale with significant positive allometry. These include the pectoral and pelvic girdles, humerus, radius+ulna, and forelimb. Total hindlimb mass and the masses of individual hindlimb bones (femur, tibia, and metatarsus) scale isometrically. Metapodial mass correlates more poorly with body mass than the girdles or any of the long bones. Metapodial mass probably reflects locomotor behavior to a greater extent than do the long bones. Long bone mass in small mammals (<50 kg) scales with significantly greater positive allometry than bone mass in large (>50 kg) mammals, probably because of the proportionally shorter long bones of large mammals as a means of preserving resistance to bending forces at large body sizes. The positive allometric scaling of the skeleton in terrestrial animals has implications for the maximal size attainable, and it is possible that the largest sauropod dinosaurs approached this limit.     

Long bone scaling in Captorhinidae: do limb bones scale according to elastic similarity in sprawling basal amniotes?

Captorhinids are a speciose clade of sauropsids that are crucial to understand several aspects of basal amniote general biology. Members of the Captorhinidae explored different diets and, amongst basal amniotes, were one of the first groups to demonstrate high‐fibre herbivory. Several papers have been published on the cranial anatomy of captorhinids, but there are relatively few studies which focus on the post‐cranium, especially on the appendicular skeleton and long bones. This contribution presents the first quantitative long bone scaling in Captorhinidae performed through morphometric analyses. From classical biomechanical research, it is well‐established that to accommodate an increase in size, gravity will result in elastic deformation of long bones. This outcome is especially significant in terrestrial tetrapods with a sprawling limb posture such as captorhinids, where great torsional stresses are applied to long bones, both during locomotion and in the resting phase. In this paper, we test whether the consistent evolutionary size increase in captorhinids led to major re‐patterning in long bone structure as theoretically expected, based on the theory of elastic similarity. Morphometric analysis shows that, apart from a small positive allometry in the humerus, captorhinid long bones scale geometrically as body size increases. Thus, the predicted elastic similarity to maintain similar levels in peak stress with an increase in dimensions does not seem not to apply to long bone evolution in captorhinids. We propose that, as already observed experimentally in larger‐bodied varanid lizards, large captorhinids could also mitigate size‐related increases in stress by reducing femur rotation and increasing the percentage of the stride cycle during which the right hindfoot was on the ground (i.e. the duty factor). In this way, large captorhinids could avoid reaching peak stress thresholds by sacrificing speed during locomotion and without a substantial long bone re‐patterning or postural change.     

Allometry and curvature in the long bones of quadrupedal mammals

The allometric relationships between basic structural proportions in long bones are examined in the humerus, radius, femur and tibia for a diverse group of 42 terrestrial quadrupedal mammals that span a size range from 0.02–6000 kg. Non-linear scaling is found for length vs. diameter in the tibia and radius, suggesting that the mechanical constraints on the skeleton differ within large and small body-size mammals. Curvature normalized to mid-shaft radius scales differently in the different long bones. Curvature is poorly related to size in the proximal limb bones (humerus and femur) while it decreases systematically with size in the tibia (mass exponent −0.13). The scaling of normalized curvature in the radius is unique among long bones. Variability of curvature in the radius is reduced at any size in comparison to that found in the other long bones. Normalized curvature is constant within the small body size group (0.02 to approximately 100 kg) while it decreases sharply with size within animals over 100 kg body mass. The unusual scaling found in the radius is probably the result of this bone's close alignment with the extrinsic forces which act on it during locomotion. The change in scaling within the radius for animals of different size may be indicative of more general size-dependent mechanical trade-offs which are masked by the complex loading circumstances of the other long bones.     

Brief communication: Locomotor limb preferences in captive chimpanzees (Pan troglodytes): implications for morphological asymmetries in limb bones

Understanding the evolutionary origins of hemispheric specialization remains a topic of considerable interest in a variety of scientific disciplines. Whether nonhuman primates exhibit population-level limb preferences continues to be a controversial topic. In this study, limb preferences for ascending and descending locomotion were assessed as a means of examining the hypothesis that asymmetries in forelimb bones might be attributed to asymmetries in posture. The results indicated that captive chimpanzees showed a population-level leftward asymmetry in descending locomotion but no group bias for ascending locomotion. The results are consistent with previous behavioral studies in captive chimpanzees as well as studies on skeletal asymmetries of the forelimbs of chimpanzees.     

What Lies Beneath: Sub-Articular Long Bone Shape Scaling in Eutherian Mammals and Saurischian Dinosaurs Suggests Different Locomotor Adaptations for Gigantism

Eutherian mammals and saurischian dinosaurs both evolved lineages of huge terrestrial herbivores. Although significantly more saurischian dinosaurs were giants than eutherians, the long bones of both taxa scale similarly and suggest that locomotion was dynamically similar. However, articular cartilage is thin in eutherian mammals but thick in saurischian dinosaurs, differences that could have contributed to, or limited, how frequently gigantism evolved. Therefore, we tested the hypothesis that sub-articular bone, which supports the articular cartilage, changes shape in different ways between terrestrial mammals and dinosaurs with increasing size. Our sample consisted of giant mammal and reptile taxa (i.e., elephants, rhinos, sauropods) plus erect and non-erect outgroups with thin and thick articular cartilage. Our results show that eutherian mammal sub-articular shape becomes narrow with well-defined surface features as size increases. In contrast, this region in saurischian dinosaurs expands and remains gently convex with increasing size. Similar trends were observed in non-erect outgroup taxa (monotremes, alligators), showing that the trends we report are posture-independent. These differences support our hypothesis that sub-articular shape scales differently between eutherian mammals and saurischian dinosaurs. Our results show that articular cartilage thickness and sub-articular shape are correlated. In mammals, joints become ever more congruent and thinner with increasing size, whereas archosaur joints remained both congruent and thick, especially in sauropods. We suggest that gigantism occurs less frequently in mammals, in part, because joints composed of thin articular cartilage can only become so congruent before stress cannot be effectively alleviated. In contrast, frequent gigantism in saurischian dinosaurs may be explained, in part, by joints with thick articular cartilage that can deform across large areas with increasing load.     

Sesamoids and Morphological Variation: a Hypothesis on the Origin of Rod-like Skeletal Elements in Aerial Mammals

Enigmatic rod-like skeletal structures that support compliant membranes (patagia) in aerial mammals have been often considered as neomorphic elements or as evolutionary novelties, and their origin has remained poorly understood. A potential source of skeletal plasticity and, probably, of morphofunctional innovations are sesamoids, which were recently demonstrated to have a common cellular origin with bone eminences. In this review, I compile information regarding anatomy, evolution, and development of rod-like skeletal elements in extant gliding and flying mammals and propose a working hypothesis on the origin of these structures. Rod-like skeletal elements, namely, the calcar in bats (Chiroptera), the unciform element in Anomaluridae (Rodentia), and the styliform cartilage in Pteromyini (Rodentia: Sciuridae), would derive from sesamoids, which, in turn, would have the same origin as eminences of long bones (or bones with a long-bone-like growth), i.e., calcaneus, ulna, and pisiform, respectively. Rod-like skeletal elements exhibit several features of sesamoids. However, further developmental data are needed to confirm this hypothesis, particularly whether these structures share a cellular origin and molecular developmental pathways with sesamoids and bone eminences. If this hypothesis were supported, a new role for sesamoids in generating morphofunctional innovations in mammals and, potentially, in other aerial amniotes, would be recognized. Rod-like skeletal elements, which are key in the evolution of aerial locomotion, might constitute an example of pre-existing traits that acquire novel functions through relatively little developmental plasticity.