Limb bone strength and habits in large glyptodonts

Fariña, R.A. 1995 11 30 Limb bone strength and habits in large glyptodonts.
The masses of some large Pleistocene species of the fossil family Glyptodontidae (Mammalia; Xenarthra) were estimated from the volumes of models. Their centres of mass were also estimated. Dimensions of limb bones and limb muscles were used to assess the athleticism of these species, using an approach previously applied to dinosaurs. The femora show higher athletic indicators (even when supporting the whole weight of the animal) than humeri in the quadrupedal stance. It is therefore proposed that performing strenuous locomotor activities bipedally was not only possible but even advantageous for minimizing risk of bone failure. The muscular dimensions analysed are consistent with this conclusion. The possible biological meaning of these mechanical results is considered. Since the smaller and older (early Miocene) glyptodont Propalaehoplophorus does not share this condition, it is suggested that it was developed later in the history of the group, perhaps as a feature related to the acquisition of large size. Glyptodonts, fossil, Xenarthra, biomechanics, locomotion, extinct megafauna, palaeobiology, evolution .     

Human body proportions explained on the basis of biomechanical principles

On the basis of theoretical biomechanics and of experiments, we investigated the mechanical requirements to which the body of a bipedally walking primate is subject, and the possibilities to meet these requirements with a minimum amount of energy. The least energy-consuming adaptation is clearly a body shape favourable for the preferred locomotion. Some characteristics of human body shape, in particular its proportions, could be identified as advantageous for fulfilling obvious biological roles or mechanical necessities. The characteristic length and the extended position of human hindlimbs make walking faster without additional input of energy. Mass distribution on the hindlimbs reduces the energy necessary for accelerating the swing limb after liftoff and for decelerating the swing limb before the heelstrike. Length and mass distribution in the forelimb gives it a pendulum length comparable to that of the hindlimb, so that both extremities swing at the same frequency. This swinging of the forelimbs counters in part the movements exerted by the moved hindlimbs on the trunk. The elongate and slim shape of the trunk provides great mass moments of inertia and that means stability against being flexed ventrally and dorsally by the forward and rearward movements of the heavy and long hindlimbs. Shoulder breadth in combination with the shallow shape of the thorax yield higher mass moments of inertia against the rotation of the trunk about a vertical axis than a cylindrical trunk shape. Further elongation of the hindlimbs is limited by the energy necessary for acceleration and deceleration, as well as for lifting them during the swing phase. In addition, the reaction forces exerted by the hindlimbs would expose the trunk to undue excursions if the proportions trunk length/limb length or trunk mass/limb mass would decrease. The above-noted kinetic requirements are partly in line, partly in conflict with the requirements of statics.     

Morphological conservation of limb natural pendular period in the domestic dog (Canis familiaris): Implications for locomotor energetics

For better understanding of the links between limb morphology and the metabolic cost of locomotion, we have characterized the relationships between limb length and shape and other functionally important variables in the straightened forelimbs and hindlimbs of a sample of 12 domestic dogs (Canis familiaris). Intra-animal comparisons show that forelimbs and hindlimbs are very similar (not significantly different) in natural pendular period (NPP), center-of-mass, and radius of gyration, even though they differ distinctly in mass, length, moment-of-inertia, and other limb proportions. The conservation of limb NPP, despite pronounced dissimilarity in other limb characteristics, appears to be the result of systematic differences in shape, forelimbs tending to be cylindrical and hindlimbs conical. Estimating limb NPP for other species from data in the literature on segment inertia and total limb length, we present evidence that the similarity between forelimbs and hindlimbs in NPP is generally true for mammals across a large size range. Limbs swinging with or near their natural pendular periods will maximize within-limb pendular exchange of potential and kinetic energy. As all four limbs of moderate- and large-size animals swing with the same period during walking, maximal advantage can be derived from the pendular exchange of energy only if forelimbs and hindlimbs are very similar in NPP. We hypothesize that an important constraint in the evolution of limb length and shape is the locomotor economy derived from forelimbs and hindlimbs of similar natural pendular period. J. Morphol. 234:183–196, 1997. © 1997 Wiley-Liss, Inc.     

Body mass estimation in xenarthra: A predictive equation suitable for all quadrupedal terrestrial placentals?

The Magnorder Xenarthra includes strange extinct groups, like glyptodonts, similar to large armadillos, and ground sloths, terrestrial relatives of the extant tree sloths. They have created considerable paleobiological interest in the last decades; however, the ecology of most of these species is still controversial or unknown. The body mass estimation of extinct species has great importance for paleobiological reconstructions. The commonest way to estimate body mass from fossils is through linear regression. However, if the studied species does not have similar extant relatives, the allometric pattern described by the regression could differ from those shown by the extinct group. That is the case for glyptodonts and ground sloths. Thus, stepwise multiple regression were developed including extant xenarthrans (their taxonomic relatives) and ungulates (their size and ecological relatives). Cases were weighted to maximize the taxonomic evenness. Twenty‐eight equations were obtained. The distribution of the percent of prediction error (%PE) was analyzed between taxonomic groups (Perissodactyla, Artiodactyla, and Xenarthra) and size groups (0–20 kg, 20–300 kg, and more than 300 kg). To assess the predictive power of the functions, equations were applied to species not included in the regression development [test set cross validation, (TSCV)]. Only five equations had a homogeneous %PE between the aforementioned groups. These were applied to five extinct species. A mean body mass of 80 kg was estimated for Propalaehoplophorus australis (Cingulata: Glyptodontidae), 594 kg for Scelidotherium leptocephalum (Phyllophaga: Mylodontidae), and 3,550.7 kg for Lestodon armatus (Phyllophaga: Mylodontidae). The high scatter of the body mass estimations obtained for Catonyx tarijensis (Phyllophaga: Mylodontidae) and Thalassocnus natans (Phyllophaga: Megatheriidae), probably due to different specializations, prevented us from predicting its body mass. Surprisingly, although obtained from ungulates and xenarthrans, these five selected equations were also able to predict the body mass of species from groups as different as rodents, carnivores, hyracoideans, or tubulidentates. This result suggests the presence of a complex common allometric pattern for all quadrupedal placentals. J. Morphol., 2008. © 2008 Wiley‐Liss, Inc.     

Morphological changes leading to hominid bipedalism

The adoption of habitual bipedal locomotion required a backward shift of the centre of gravity of the body, to a level relative to the supporting surface area of the body and pivotal axis of the hips at which walking with extended knees became practicable. In the morphology of the immediate ancestors to hominids, there were relatively few features whose change could have effectively affected the position of the gravity vertical of the body. Weight could be distributed posteriorly mainly by increasing the mass of the hindlimbs and the kyphotic curve of the vertebral column, by developing buttocks. and by flattening the thorax and abdomen. Estimates indicate that the adoption of a suitable curvature of the vertebral column alone was not sufficient for shifting the centre of gravity behind the 'threshold of bipedalism'. A considerable increase in the mass of the hindlimbs was also required, and the addition of the mass of the buttocks may have represented the decisive factor for crossing the threshold. A possible physical enviroment in which the change to bipedalism could have taken place was a mountainous terrain with long, steep slopes, transected by gorges and precipices. In such a terrain the increase in the power and mass of both the hindlimbs and the gluteus maximus proprius muscle could have been favoured by selection, leading ultimately to a condition in which the combined effect of heavy hindlimbs. a suitable curvature of the spine and the weight of the buttocks shifted the centre of gravity of the body backwards to a level at which habitual walking with extended knees became practicable. □ Hominid evolution. bipedalism, evolutionary thresholds.     

On birds: scale effects in the neognath hindlimb and differences in the gross morphology of wings and hindlimbs

Scale effects on whole limb morphology (i.e. bones together with in situ overlying muscles) are well understood for the neognath forelimb. However, scale effects on neognath gross hindlimb morphology remain largely unexplored. To broaden our understanding of avian whole limb morphology, I investigated the scaling of hindlimb inertial properties in neognath birds, testing empirical scaling relationships against the model of geometric similarity. Inertial property data – mass, moment of inertia, centre of mass distance, and radius of gyration – were collected from 22 neognath species representing a wide range of locomotor specializations. When scaled against body mass, hindlimb inertial properties scale with positive allometry. Thus, in terms of morphology, larger bodied neognaths possess hindlimbs requiring disproportionately more energy to accelerate and decelerate relative to body mass than smaller bodied birds. When scaled against limb length, hindlimb inertial properties scale according to isometry. In the subclade Land Birds (sensu Hackett et al.), hindlimb inertial properties largely scale according to positive allometry. The contrasting results of positive allometry vs. isometry in neognaths are due to how hindlimb length scales against body mass. Negative allometry of hindlimb inertial properties, which would reduce terrestrial locomotion costs, would probably make the hindlimb susceptible to mechanical failure or too diminutive for its many ecological functions. Comparing the scaling relationships of wings and hindlimbs highlights how locomotor costs influence the scaling of limb inertial properties. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 14–31.     

Patterns of fluctuating asymmetry in the limbs of anurans

It has been hypothesized that fluctuating asymmetry (FA) may provide an indication of the functional importance of structures within an organism, with structures that more strongly impact fitness being more symmetric. Based on this idea, we predicted that for tetrapods in which the forelimbs and hindlimbs play an unequal role in locomotion, the less functionally important limb set should display higher levels of FA. We conducted a multispecies test of this hypothesis in anurans (frogs and toads), whose saltatory locomotor mode is powered by the hindlimbs. We also tested whether FA in the forelimbs, which play a more important role during landing, differed between families that differ in the degree of forelimb use in locomotion (Bufonidae vs. Ranidae). We calculated FA from the lengths of humeri and femora measured from disarticulated skeletal specimens of four anuran taxa (Bufonidae: Anaxyrus americanus, Rhinella marina; Ranidae: Lithobates catesbeianus, Lithobates clamitans). Our findings were consistent with the hypothesis that natural selection for increased locomotor performance may influence patterns of FA seen in vertebrate limbs, with all species displaying lower mean FA in the hindlimbs. More subtle functional roles between the forelimbs of bufonids and ranids, however, did not elicit different levels of FA.     

Reduced phenotypic covariation in marsupial limbs and the implications for mammalian evolution

As serially homologous structures, mammalian fore‐ and hindlimbs ancestrally share a common developmental and genetic architecture. As a result, mammalian fore‐ and hindlimbs are predicted to be highly integrated in the absence of selective pressures to form divergent limb morphologies. Marsupials experience such a divergent selective pressure to form a robust forelimb to power a post‐natal crawl to the teat. In this study, phenotypic covariation in marsupials was assessed to determine if specialization for the crawl did indeed reduce integration between their fore‐ and hindlimbs. To explore the evolution of mammalian limb integration, phenotypic covariation in representative eutherians and monotremes was also examined. Phenotypic covariation in limbs was quantified morphometrically, and analysed with correlational and phylogenetic methods. Results indicate that marsupials generally have relatively high levels of within‐limb phenotypic covariation, and low levels between limbs, in contrast to the pattern reconstructed for the mammalian ancestor. Our findings support the hypothesis that pressure to specialize in one limb (either the fore‐ or the hindlimb) can reduce phenotypic covariation between limbs, and that reduced limb phenotypic covariation is derived in marsupials. Further research is needed to test the effect that these differences in limb phenotypic covariation had on the evolution of the major mammalian groups. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 22–36.     

Exoskeleton and Systematics: A Historical Problem in the Classification of Glyptodonts

The glyptodonts (Mammalia: Cingulata) are characterized by an exoskeleton comprising most notably an armored tail and an immobile dorsal carapace formed by a large number of osteoderms. In 1889, Florentino Ameghino published the first phylogenetic scenario for the glyptodonts, based on the sequential application of two transformation series related to the morphology of the tail armor and carapace osteoderms. From the early to mid 1900s, several authors used Ameghino’s transformation series subordinated to a model of evolution in which derived glyptodont groups had arisen independently from separate pre-middle Miocene ancestors. This approach, in which the morphological states of Ameghino’s series were considered analogous rather than homologous, provided different phylogenetic scenarios and the paraphyletic classification still in use. Two recent cladistic analyses based on cranial and postcranial (including caudal tube) characters support the monophyly of glyptodonts and suggest novel intra-clade relationships. However, neither analysis included the classic osteoderm characters used by earlier authors. Therefore, we propose new osteoderm and carapace characters and evaluate their performance in a new cladistic analysis. We found that: a) some osteoderm characters used by earlier authors to support ancestor-descendent hypotheses are in fact fully homoplastic autapomorphies (e.g., multiplication of the number of rows of peripheral figures); b) characters previously believed to have originated independently in several groups (e.g., presence of caudal tube) are synapomorphies at a wider hierarchical level; c) some ancestor–descendant pre-cladistic hypotheses are incompatible with the topology and synapomorphy distribution obtained; and d) there is no reason to favor exoskeletal characters in glyptodont systematics.     

Influences of limb proportions and body size on locomotor kinematics in terrestrial primates and fossil hominins

During locomotion, mammalian limb postures are influenced by many factors including the animal's limb length and body mass. Polk (2002) compared the gait of similar-sized cercopithecine monkeys that differed limb proportions and found that longer-limbed monkeys usually adopt more extended joint postures than shorter-limbed monkeys in order to moderate their joint moments. Studies of primates as well as non-primate mammals that vary in body mass have demonstrated that larger animals use more extended limb postures than smaller animals. Such extended postures in larger animals increase the extensor muscle mechanical advantage and allow postures to be maintained with relatively less muscular effort (Polk, 2002; Biewener 1989). The results of these previous studies are used here to address two anthropological questions. The first concerns the postural effects of body mass and limb proportion differences between australopithecines and members of the genus Homo. That is, H. erectus and later hominins all have larger body mass and longer legs than australopithecines, and these anatomical differences suggest that Homo probably used more extended postures and probably required relatively less muscular force to resist gravity than the smaller and shorter-limbed australopithecines. The second question investigates how animals with similar size but different limb proportions differ in locomotor performance. The effects of limb proportions on gait are relevant to inferring postural and locomotor differences between Neanderthals and modern Homo sapiens which differ in their crural indices and relative limb length. This study demonstrates that primates with relatively long limbs achieve higher walking speeds while using lower stride frequencies and lower angular excursions than shorter-limbed monkeys, and these kinematic differences may allow longer-limbed taxa to locomote more efficiently than shorter-limbed species of similar mass. Such differences may also have characterized the gait of Homo sapiens in comparison to Neanderthals, but more experimental data on humans that vary in limb proportions are necessary in order to evaluate this question more thoroughly.     

Allometric and Group Differences in the Xenarthran Femur

The aim of this study is to analyze shape variation in the xenarthran femur to gain insights into their behavior and locomotion. Specimens of both Cingulata (armadillos and glyptodonts) and Pilosa (anteaters and sloths) were studied and within each group body mass varies by several orders of magnitude. The main focus of the analysis was allometric variation in femoral shape in the three groups studied, armadillos, glyptodonts, and pilosans. Three dimensional coordinates were recorded for 40 homologous landmarks on each of 51 xenarthran femurs. The data were analyzed by geometric morphometric methods, and form space analysis was used to identify the allometric variation in each of the three groups. Across all groups, larger specimens tended to have larger articular surfaces, more robust femora generally, and the shape of the femoral condyles was more suited to extended postures. In addition, in larger specimens the medial condyle was much larger than the lateral condyle and the third trochanter was located more distally. The larger armadillo femora had a greater trochanter located considerably proximal to the femoral head and this is thought to improve femoral extension, but in glyptodonts and pilosans the larger specimens had a greater trochanter that was far lateral to the femoral head and this is interpreted as enhancing femoral rotation.     

The scaling of skeletal microanatomy in non-human primates

The study of scale-correlated changes in the external dimensions and cross-sectional geometry of primate long bones is fundamental to our understanding of primate limb bone structural adaptation. To date, however, there have been no studies of the effects of mechanical loading on patterns of skeletal scaling at the microstructural level. To remedy this, we analysed patterns of microanatomical scaling in the humeri and femora of 107 adult primates belonging to the families Galagonidae and Cercopithecidae. Seven species were included in our analysis. Proximal, midshaft, and distal sections of humeri and femora of each individual were examined and secondary osteonal and cortical area were measured. Secondary osteonal area scales positively allometrically with cortical cross-sectional area and with body mass. This pattern holds generally for humeri and femora—both within and across families. However, there are striking dissimilarities in the relative strengths of the allometric coefficients for humeri and femora measured for different families. These distinctions appear to be related to differences in the ways in which fore- and hindlimbs are loaded. Such differences highlight the promise of microstructural data and the importance of examining the confounding effects of locomotory behaviour in studies of skeletal scaling.     

Lens formation from the cornea following implantation into hindlimbs of larval Xenopus laevis: the influence of limb innervation and extent of differentiation

Corneal fragments of larval Xenopus laevis at stage 48 (according to Nieuwkoop and Faber, '56), were implanted into sham denervated unamputated hindlimbs, denervated unamputated hindlimbs, amputated and sham denervated hindlimbs, and amputated and denervated hindlimbs of larvae at stages 52 and 57. The results show that unamputated limbs at stage 52, either innervated or denervated, manifest a weak capacity to promote the first lens-forming transformations of the outer cornea. This capacity is absent in both limb types at stage 57. After amputation, limbs of both early and late stages form a regenerative blastema and support lens formation from the outer cornea. Denervation of early stage limbs has no appreciable effect on blastema formation and lens-forming transformation of corneal implants. However, denervation of late stage limbs inhibits both processes. These results indicate that the limb tissues of the early stage limbs contain non-neural inductive factors at a low level and that after limb amputation and blastema formation the level of these factors becomes high enough to promote lens formation from implanted cornea, even after denervation. In contrast, the limb tissues of late stage limbs do not contain a suitable level of non-neural inductive factors.     

Effects of phosphoramide mustard and acrolein, cytotoxic metabolites of cyclophosphamide, on mouse limb development in vitro

Phosphoramide mustard and acrolein are toxic and reactive metabolites of the widely used anticancer drug and known teratogen cyclophosphamide. To study the mechanism(s) involved and to determine which of the active metabolites of cyclophosphamide is responsible for the production of limb malformations, the effects of exposure of cultured limb buds to phosphoramide mustard and acrolein were investigated. Fore- and hindlimbs were excised from ICR mice on day 12 of gestation and cultured in roller bottles for 6 days. Limbs were exposed to either phosphoramide mustard or acrolein (10 or 50 micrograms/ml) for the first 20 hours of the culture period. Exposure to phosphoramide mustard produced limb reduction malformations in both the fore- and hindlimbs; total limb bone area was greatly reduced, while the relative contribution of the paw to this area in forelimbs was increased. There was a fourfold reduction in both DNA and RNA; protein content was reduced only by one-half. Alkaline phosphatase activity was significantly decreased in fore- and hindlimbs exposed to phosphoramide mustard, whereas creatine phosphokinase activity was only reduced in hindlimbs in the limbs exposed to the higher concentration of phosphoramide mustard. Exposure to acrolein also produced malformed limbs with a mangled appearance; however, total limb bone area and the relative contribution of the long bones versus paw structures were not altered. Acrolein exposure had little effect on growth parameters such as DNA (decreased only in hindlimbs exposed to 50 micrograms/ml), RNA (increased in hindlimbs exposed to 50 micrograms/ml), or protein content. Alkaline phosphatase and creatine phosphokinase activities were not altered in acrolein-exposed fore- or hindlimbs. Thus, phosphoramide mustard and acrolein have dramatically different effects on developing limbs in vitro; this observation may indicate that they have different targets and/or mechanisms of action as teratogens in the limb. The effects of phosphoramide mustard are very similar to those of "activated" cyclophosphamide (4-hydroperoxycyclophosphamide).     

Body proportions in ancient Andeans from high and low altitudes

Living human populations from high altitudes in the Andes exhibit relatively short limbs compared with neighboring groups from lower elevations as adaptations to cold climates characteristic of high-altitude environments. This study compares relative limb lengths and proportions in pre-Contact human skeletons from different altitudes to test whether ecogeographic variation also existed in Andean prehistory. Maximum lengths of the humerus, radius, femur, and tibia, and femoral head breadth are measured in sex-specific groups of adult human skeletons (N = 346) from the central (n = 80) and the south-central (n = 123) Andean coasts, the Atacama Desert at 2,500 m (n = 102), and the southern Peruvian highlands at 2,000-3,800 m (n = 41). To test whether limb lengths vary with altitude, comparisons are made of intralimb proportions, limb lengths against body mass estimates derived from published equations, limb lengths against the geometric mean of all measurements, and principal component analysis. Intralimb proportions do not statistically differ between coastal groups and those from the Atacama Desert, whereas intralimb proportions are significantly shorter in the Peruvian highland sample. Overall body size and limb lengths relative to body size vary along an altitudinal gradient, with larger individuals from coastal environments and smaller individuals with relatively longer limbs for their size from higher elevations. Ecogeographic variation in relation to climate explains the variation in intralimb proportions, and dietary variation may explain the altitudinal cline in body size and limb lengths relative to body size. The potential effects of gene flow on variation in body proportions in Andean prehistory are also explored.     

Quadrupedal locomotion in squirrel monkeys (Cebidae: Saimiri sciureus): a cineradiographic study of limb kinematics and related substrate reaction forces

Quadrupedal locomotion of squirrel monkeys on small-diameter support was analyzed kinematically and kinetically to specify the timing between limb movements and substrate reaction forces. Limb kinematics was studied cineradiographically, and substrate reaction forces were synchronously recorded. Squirrel monkeys resemble most other quadrupedal primates in that they utilize a diagonal sequence/diagonal couplets gait when walking on small branches. This gait pattern and the ratio between limb length and trunk length influence limb kinematics. Proximal pivots of forelimbs and hindlimbs are on the same horizontal plane, thus giving both limbs the same functional length. However, the hindlimbs are anatomically longer than the forelimbs. Therefore, hindlimb joints must be more strongly flexed during the step cycle compared to the forelimb joints. Because the timing of ipsilateral limb movements prevents an increasing amount of forelimb retraction, the forelimb must be more protracted during the initial stance phase. At this posture, gravity acts with long moment arms at proximal forelimb joints. Squirrel monkeys support most of their weight on their hindlimbs. The timing of limb movements and substrate reaction forces shows that the shift of support to the hindlimbs is mainly done to reduce the compressive load on the forelimb. The hypothesis of the posterior weight shift as a dynamic strategy to reduce load on forelimbs, proposed by Reynolds ([1985]) Am. J. Phys. Anthropol. 67:335-349; [1985] Am. J. Phys. Anthropol. 67:351-362), is supported by the high correlation of ratios between forelimb and hindlimb peak vertical forces and the range of motion of shoulder joint and scapula in primates.     

The influence of denervation on grafted hindlimb regeneration of larval Xenopus laevis

The aim of the present research is to ascertain whether in larval Xenopus laevis nerve-independence for the regeneration of early stage limbs and nerve-dependence of late stage limbs observed in a previous work (Filoni and Paglialunga, '90) is related to extrinsic (systemic) factors or to intrinsic changes taking place in the limb cells themselves during development. In this paper the regenerative capacity of early and late stage hindlimbs under the same extrinsic conditions, insofar as both are grafted onto the denervated hindlimbs of host larvae at the same developmental stage, is studied. All the grafted limbs are amputated after the host larvae have reached stage 57-58 (according to Nieuwkoop and Faber, '56). In experiment I, the grafted limb is amputated at stage 52, at the thigh level; in experiment II, the grafted limb is amputated at stage 54-55, at the tarsalia level; in experiment III the grafted limb is amputated at stage 57, at the tarsalia level. In all three experiments, together with the grafted limb, also the host limb is amputated at the tarsalia level. The results show that while grafted limbs amputated at stages 52 and 54-55 regenerate in the absence of nerves, grafted limbs amputated at stage 57 cannot. The failure of late stage grafted limbs to regenerate cannot be explained in terms of an immune-type inhibiting reaction since it has been observed also in denervated autografted limbs and in the host limbs. Since all the grafted limbs are in the same environmental conditions, the results show that in larval Xenopus laevis nerve-independence for regeneration of early stage limbs and nerve-dependence of late stage limbs are not related to factors extrinsic to the limb but to intrinsic changes taking place in the limb cells themselves during development.     

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.     

The relationship of innervation and differentiation to regenerative capacity in the reamputated hindlimb of larval Xenopus laevis

Regenerated hindlimbs of larval Xenopus laevis were reamputated at critical larval stages and levels, viz when amputation of the control limb at the same larval stage and level is followed by reduced regeneration. Reamputations were performed at the level of (1) the original plane of amputation, (2) the early regenerate (cone/palette stage), (3) the late regenerate (digit stage). Reamputation increased both the percentage rate of regeneration and the morphological complexity of the regenerates in all experimental series. Cell counts in lateral motor columns and spinal ganglia innervating the hindlimb, together with histological observations and mitotic index and labelling index determinations in reamputated and control limbs showed that improved regeneration in the reamputated limb was related to an increase in undifferentiated and proliferating cells in the stump. We did not find any evidence suggesting that renewed regeneration in reamputated anuran limbs results from an increase in innervation, as has previously been hypothesized. We support our conclusions by demonstrating an improvement in regenerationen in the reamputated and denervated hindlimbs.     

An Overview of the Presence of Osteoderms in Sloths: Implications for Osteoderms as a Plesiomorphic Character of the Xenarthra

The presence of osteoderms in the skin of some extinct sloths and in cingulates (armadillos, pampatheres, and glyptodonts) has often been considered a pleisomorphic character of the Xenarthra. While osteoderms are known from the earliest cingulates, they are absent in most sloths including the two extant taxa and only appear late in their fossil record. Osteoderms are currently only reported from five genera of mylodonts and two megatheres, out of the over 100 currently recognized genera of sloths. Consequently, rather than a plesiomorphic character of the Xenarthra, which has been secondarily lost in sloths, it is more likely that osteoderms in sloths are the result of parallel evolution to the cingulates that independently evolved in one, possibly two different sloth clades.