The functional morphometrics of the hip and thigh in leaping prosimians

Examination of the form of the hip and thigh through discriminant function analysis of a series of 15 osteometric indices taken upon 289 specimens of 20 sets of leaping prosimians distinguishes four groups. Around a centrally located group of the various species of Lemur are three separate rays: one containing the three indriid genera (Propithecus, Indri, and Avahi), a second including the galagines and Tarsius, and a third comprising the two species of Cheirogaleus and the genus Microcebus. The lemurine genus Lepilemur lies intermediately between the central group of lemurs and the indriids. The lemurine genus Hapalemur lies intermediately between, on the one hand, the centrally located group of lemurs, and on the other, each of the two groups, cheirogaleines and galagines-plus-tarsiers. The arrangements of the particular species within each of these last two groups separately is from those that leap least to those that leap most. Given that the structure of the hip and thigh is related to the biomechanical demands of leaping, the existence of these separate morphological groups implies the existence of different biomechanical modes of leaping. Such meager taxonomic and behavioral information as is available seems to support this idea, although it is not impossible that other aspects of the behaviors of the animals may also be associated. This information sets up a series of new behavioral hypotheses that might be tested by better field and laboratory studies aimed more precisely at locomotion. And this information may be useful in helping to make assessments of particular fossils and in helping determine how various prosimian locomotor modes may have evolved.     

Locomotor behavior and skeletal morphology of two sympatric pitheciine monkeys,Pithecia pithecia and Chiropotes satanas

Field observations of two sympatric pitheciine species reveal that the positional repertoire of the white-faced saki, Pithecia pithecia, is dominated by leaping behaviors, whereas the bearded saki, Chiropotes satanas, is predominantly quadrupedal. Examination and comparison of the postcranial skeletal morphologies and limb proportions of these species display numerous features associated with their respective locomotor behaviors. These observations accord with associations found in other primate and mammalian groups and with predictions based on theoretical and experimental biomechanics. Preliminary observations of the skeletal morphology of Cacajao calvus demonstrate a marked similarity to that of Chiropotes. The fossil platyrrhine Cebupithecia sarmientoi displays greater similarity to Pithecia, suggesting that its positional repertoire also included significant leaping and clinging behaviors.     

A new crocodylomorph specimen from the Araripe Basin (Crato Member,Santana Formation), northeastern Brazil

The fossil fauna of the Santana Formation (Early Cretaceous) comprises many distinct taxa, but crocodylomorphs are poorly understood. Here we describe a new specimen (MPSC-R1137) that consists of a complete hind limb found in the Crato Member, the basal section of the Santana Formation. Based on the characteristics of the fibula (e.g., pronounced variation of the shaft width) and length proportions of the femur and tibia, this specimen can be distinguished from Caririsuchus camposi and Araripesuchus gomesii, which are known from the Romualdo Member (the upper lithostratigraphic unit of the Santana Formation). The only crocodylomorph formally described from the Crato Member is Susisuchus anatoceps, whose holotype lacks elements of the hind limb. On the basis of a comparative anatomical study of the hind limb, which shows no similarities between MPSC-R1137 and other crocodylomorphs from the Araripe Basin, we tentatively classify this new specimen as cf. Susisuchus sp., and provide new anatomical information for this rather derived crocodylomorph.      

Phylogenetic Separation in Limb Use in Captive Gibbons (Hylobatidae): A Comparison Across the Primate Order

Although there have been few studies of self‐scratching in primates, some have reported distinct differences in whether hands or feet are used, and these variations seem to reflect the evolutionary history of the Order. Monkeys and prosimians use both hands and feet to self‐scratch while African great apes use hands almost exclusively. Gibbons represent an evolutionary divergence between monkeys and great apes and incidental observations at the Gibbon Conservation Center pointed to a difference in self‐scratching among the four extant gibbon genera (Hoolock, Nomascus, Symphalangus, and Hylobates). To validate and further explore these preliminary observations, we collected systematic data on self‐scratching from 32 gibbons, including nine species and all four genera. To supplement gibbon data, we also collected self‐scratching information from 18 great apes (four species), five prosimians (two species), 26 New World Monkeys (nine species) and 20 Old World Monkeys (seven species). All monkeys and some prosimians used both hands and feet to self‐scratch, whereas one prosimian species used only feet. All African great apes used hands exclusively (orangutans were an exception displaying occasional foot‐use). This appears to represent a fundamental difference between monkeys and great apes in limb use. Interestingly, there was a clear difference in self‐scratching between the four gibbon genera. Hylobates and Symphalangus self‐scratched only with hands (like all African great apes), while Hoolock and Nomascus self‐scratched with both hands and feet (like monkeys and prosimians). This difference in gibbon behavior may reflect the evolutionary history of gibbons as Hoolock and Nomascus are thought to have evolved before both Hylobates and Symphalangus. What evolutionary pressures led to this divergent pattern is currently opaque; however, this shift in limb preference may result from niche separation across the order facilitating differences in the behavioral repertoire associated with hind and forelimbs. Am. J. Primatol. 74:1035‐1043, 2012. © 2012 Wiley Periodicals, Inc.     

Macrauchenia patachonica Owen, 1838: Limb bones morphology,locomotory biomechanics,and paleobiological inferences

Macrauchenia patachonica Owen, 1838 was among the last and largest litopterns, an extinct order of South American native ungulates. Macrauchenia patachonica had anatomical peculiarities as extremely retracted nasals, enlarged cervical vertebrae, and limb bones proportions without good living analogs that lead to asking about its paleobiology. To quantitatively assess the strange combination of limb bone features in M. patachonica, we constructed an indicator of differences in anatomical adaptations for efficient running between forelimb and hind limb (IDFH). We also made a multivariate analysis using data on osteological ratios of living mammals and two other litopterns. We discuss several biomechanical and paleobiological implications of the striking differences between hind limb and forelimb design in M. patachonica. Our main suggestion is that M. patachonica, during fast locomotion, probably used a posture with the neck in a horizontal position.     

Locomotory abilities and habitat of the Cretaceous bird Gansus yumenensis inferred from limb length proportions

The relative length proportions of the three bony elements of the pelvic (femur, tibiotarsus and tarsometatarsus) and pectoral (humerus, ulna and manus) limbs of the early Cretaceous bird Gansus yumenensis, a well‐represented basal ornithuromorph from China, are investigated and compared to those of extant taxa. Ternary plots show that the pectoral limb length proportions of Gansus are most similar to Apodiformes (swifts and hummingbirds), which plot away from all other extant birds. In contrast, the pelvic limb length proportions of Gansus fall within the extant bird cluster and show similarities with the neornithine families Podicipedidae (grebes), Diomedeidae (albatross) and Phalacrocoracidae (cormorants). Although it does have some of the pelvic limb features of grebes and cormorants, the femur of Gansus is more gracile and is thus more consistent with an albatross‐like shallow‐diving mode of life than a strong foot‐propelled diving movement pattern. The position of Gansus in pectoral limb ternary morphospace is largely due to its elongated manus. In contrast to apodiformes, where the humerus and ulna are short and robust, an adaptation, which provides a stiff wing for their demanding fast agile and hovering flight (respectively), the wing‐bones of Gansus are slender, indicating a less vigorous flapping flight style. The suite of characters exhibited by Gansus mean it is difficult to completely interpret its likely ecology. Nevertheless, our analyses suggest that it is probable that this bird was both volant and capable of diving to some degree using either foot‐propelled or, perhaps, both its wings and its feet for underwater locomotion.     

Variation in the leaping of lemurs

Locomotor diversity among lemurids has been ignored by placing most species into a category of arboreal quadrupeds. Recent field studies have shown that leaping behaviors comprise a relatively large amount of their travel. In this contribution I detail a study of captive lemurs locomoting on a designed support network and/or in outdoor enclosures. Associations between support context, leaping behaviors, and landing kinematics are detailed for Hapalemur griseus, Eulemur rubriventer, Eulemur fulvus, Eulemur mongoz, and Varecia variegata. In-air body position, including the amount and location of trunk flexion, coupled with the extent of shoulder and hip flexion dictates limb use at landing. Limb strike pattern when landing onto the same support type varies interspecifically. The kinematic variation in leaping behaviors may well have implications for functional analyses of the postcranium. © 1996 Wiley-Liss, Inc.     

Submaximal leaping in the grey mouse lemur

In arboreal animals such as the grey mouse lemur (Microcebus murinus Miller, 1777), leaping is the most frequent strategy for predator avoidance. The aim of this study was to characterise the locomotor adaptation in response to the structural constraint of the habitat (i.e., position of the landing substrate). Thus, we characterised the push-off phase by inducing the lemurs to leap up to a range of heights from horizontal to their own individual highest performance. Using uniplanar high-frequency cineradiographs collected in a sagittal plane, the relative contributions of the centre of mass (CoM) velocity vector magnitude and orientation to leaping performance were evaluated. The kinematics of the push-off phase showed that for low landing heights, leaping performance was essentially due to hip and knee extensions. Higher leaps seemed to be related to an increase in ankle contribution. At all leaping heights, the proximal-to-distal sequence of the hind limb joints controlled the orientation and magnitude of the M. murinus CoM velocity vector while pushing off. Finally, the analysis of the velocity vector at the onset of take-off suggested that the optimal solution for predator avoidance was to leap for horizontal distance and not for vertical distance.     

A model for comparison of masticatory effectiveness in primates

A model is presented to express how effectively animals increase the exposed surface area of their food by chewing. It includes a coefficient of masticatory effectiveness (E) the value of which increases with effectiveness of exposing new food surface area with each chew. Humans and other species of primates differ significantly in their values of E; among the nonhuman primates studies, Lemur catta has a higher coefficient than Lemur fulvus, and both have higher coefficients than either Varecia variegatus or Galago crassicaudatus argentatus. The differences among the coefficients to these prosimians are correlated with variations in specific features of the molar morphology. Of six lower molar shearing crests considered, the relative length of the post-metacristid correlates most highly with the coefficient of masticatory effectiveness for the prosimian species. Also, among comparable-sized prosimians, E correlates significantly with the absolute postmetacristid length. Both these findings indicate that the relative size of molar shearing crests is related significantly to how effectively an animal chews its food. There are also implications for an adaptation to a high-fiber diet.     

Phylogenetic relationships within the tribe Janieae (Corallinales,Rhodophyta) based on molecular and morphological data: a reappraisal of Jania1

Generic boundaries among the genera Cheilosporum, Haliptilon, and Jania—currently referred to the tribe Janieae (Corallinaceae, Corallinales, Rhodophyta)—were reassessed. Phylogenetic relationships among 42 corallinoidean taxa were determined based on 26 anatomical characters and nuclear SSU rDNA sequence data for 11 species (with two duplicate plants) referred to the tribe Corallineae and 15 species referred to the tribe Janieae (two species of Cheilosporum, seven of Haliptilon, and six of Jania, with five duplicate plants). Results from our approach were consistent with the hypothesis that the tribe Janieae is monophyletic. Our data indicate, however, that Jania and Haliptilon as currently delimited are not monophyletic, and that Cheilosporum should not be recognized as an independent genus within the Janieae. Our data resolved two well‐supported biogeographic clades for the included Janieae, an Indian‐Pacific clade and a temperate North Atlantic clade. Among anatomical characters, reproductive structures reflected the evolution of the Janieae. Based on our results, three genera, Cheilosporum, Haliptilon, and Jania, should be merged into a single genus, with Jania having nomenclatural priority. We therefore propose new combinations where necessary of some species previously included in Cheilosporum and Haliptilon.     

Birth month associations with height,head circumference,and limb lengths among peruvian children

Associations between season of birth and body size, morbidity, and mortality have been widely documented, but it is unclear whether different parts of the body are differentially sensitive, and if such effects persist through childhood. This may be relevant to understanding the relationship between early life environment and body size and proportions. We investigated associations between birth month and anthropometry among rural highland (n = 162) and urban lowland (n = 184) Peruvian children aged 6 months to 8 years. Stature; head‐trunk height; total limb, ulna, tibia, hand, and foot lengths; head circumference; and limb measurements relative to head‐trunk height were converted to internal age‐sex‐specific z scores. Lowland and highland datasets were then analyzed separately for birth month trends using cosinor analysis, as urban conditions likely provide a more consistent environment compared with anticipated seasonal variation in the rural highlands. Among highland children birth month associations were significant most strongly for tibia length, followed by total lower limb length and stature, with a peak among November births. Results were not significant for other measurements or among lowland children. The results suggest a prenatal or early postnatal environmental effect on growth that is more marked in limb lengths than trunk length or head size, and persists across the age range studied. We suggest that the results may reflect seasonal variation in maternal nutrition in the rural highlands, but other hypotheses such as variation in maternal vitamin D levels cannot be excluded. Am J Phys Anthropol 154:115–124, 2014. © 2014 The Authors. American Journal of Physical Anthropology Published by Wiley Periodicals, Inc.     

Evolutionary aspects of primate locomotion

Both neontological and phylogenetic studies are necessary to interpret primate locomotion. Reference to palaeoprimatology and palaeocology, for instance, will lead to a fuller understanding of the roots of such gaits as the vertical clinging and leaping of Tarsius, Indri and Propithecus. Evolutionary trends in posture and locomotion are discussed. The postural trend has been towards maintenance of trunk verticality and the locomotor trend towards an increasing dependence on the forelimbs among arboreal primates. Three stages are recognized in the phylogenetic course of arboreal locomotor adaptation: Stage A. Vertical clinging and leaping; Stage B. Quadrupedalism; Stage C. Brachiation. The role of prehensility of the hand in the evolution of locomotor types is discussed in relation to forest morphology and, in particular, to stratification. Finally a scheme of evolution, set in the framework of ecology, for Old World Monkey groups is presented.     

MORPHOLOGICAL DIVERSIFICATION AND ADAPTIVE RADIATION: A COMPARISON OF TWO DIVERSE LIZARD CLADES

We compared the morphological diversity (i.e., the amount of morphological space occupied) of two similar clades, the lizard genera Anolis and Sceloporus. These species-rich monophyletic clades are similar in body size, age of origin, and many aspects of their natural history. We examined a number of morphological traits whose variation is likely to represent adaptation to different aspects of the environment, including body size, limb proportions, head dimensions, and tail length. Examination of the position of species in multidimensional space, based on a principal components analysis, indicates that the morphological diversity of Anolis, which we refer to as disparity, is significantly greater than that of Sceloporus. One potential explanation for this pattern is that morphological diversification in Anolis was facilitated by the evolution of subdigital toe-pads, which allow anoles to use the environment in ways not available to Sceloporus. The geographic location of diversification (tropical and subtropical for Anolis, arid for Sceloporus) may also have been important.     

Pilot survey of avahi population (woolly lemurs) in littoral forest fragments of southeast Madagascar

This study presents a pilot survey of the avahis (pair-living, nocturnal prosimians) living in threatened littoral forest fragments of southeastern Madagascar. In the period of October–December 2004, I evaluated the density of adult and newborn avahis (carried by the mothers) by counting the individuals encountered during 58 night walks in seven fragments of Sainte Luce and Mandena forests, along one trail/fragment. Along each trail, I used random plots (5 m²) for a preliminary characterization of the vegetation. The density of the population was not correlated with fragment size and number of plant morphospecies, while it was correlated with large tree availability. Possibly due to a low energy diet based on leaves and to specialized and energetically expensive vertical leaping, the loss of large trees by selective logging seems to affect avahi populations more than other variables.     

Fiber type population in limb muscles ofMicrocebus murinus

Populations and distributions of fiber types were studied in 19 limb muscles ofMicrocebus murinus. Proportions and cross-sectional areas of muscles fiber types were compared with data from the literature for other prosimians (Galago, Lemur, andNycticebus), another primate (Macaca cynomolgus), and the rat. Most muscles are heterogenous, with type I fibers (slow oxidative) localized in the deeper part, near the bone. Type IIA fibers (fast oxidative glycolytic) are more evenly distributed than type I and type IIB (fast glycolytic). The combination of large number and large size of type I fibers results in enhanced slow-twitch and oxidative properties as required for antigravity function of postural muscles. Compared with other primates,Microcebus shows relatively small cross-sectional areas of fibers and less numerous type I fibers, in every muscle, which is probably related to small body mass. The fiber type population of the different components of the quadriceps femoris is also related to the particular mode of locomotion of the mouse-lemur: running and leaping, climbing and hopping. M. vastus medialis and m. vastus lateralis are made up only of fast twitch fibers, IIA and IIB. A possible repercussion of hypothyroidism during the rest season and a decrease in locomotor activity was the subject of investigation of the fiber type proportion and section areas. No difference were found between individuals euthanized during the active period and those at rest period. Either a very low level of thyroxine associated with reduced activity is sufficient to maintain the processes controlling myosin expression, or the effects on muscles fibers of natural hypothyroidism and hypokinesia neutralize each other during the rest season.     

An analysis of birth sex ratio bias in captive prosimian species

The extent to which sex ratio bias is a common reproductive characteristic of prosimians has not been well established. The present study analyzed reproduction in 13 breeding groups of captive prosimians for evidence of birth sex ratio bias. A substantial male bias was demonstrated in nongregarious, but not gregarious, breeding groups. Analyses of birth sex ratios of individual mothers suggested that the observed bias did not result from the tendency of a few mothers to overproduce males, but rather from a small but reliable excess of male births in general. An examination of infant mortality revealed that male Otolemur garnettii and Microcebus murinus infants were more vulnerable to preweaning mortality, whereas female Eulemur fulvus albifrons infants were more vulnerable. An analysis of birth order by sex found that mothers of one group (O. garnettii) tended to produce males initially and females later. Additionally, a distinct pattern of birth seasonality was noted among Malagasy prosimians that was absent in the African prosimians. Greater length of period of sexual receptivity for nongregarious females as compared to gregarious females is proposed as a possible mechanism of male birth sex ratio bias. © 1996 Wiley-Liss, Inc.     

Natural selection in the <Emphasis Type="Italic">TLR-</Emphasis>related genes in the course of primate evolution

The innate immune system constitutes the front line of host defense against pathogens. Toll-like receptors (TLRs) recognize molecules derived from pathogens and play crucial roles in the innate immune system. Here, we provide evidence that the TLR-related genes have come under natural selection pressure in the course of primate evolution. We compared the nucleotide sequences of 16 TLR-related genes, including TLRs (TLR1–10), MYD88, TILAP, TICAM1, TICAM2, MD2, and CD14, among seven primate species. Analysis of the non-synonymous/synonymous substitution ratio revealed the presence of both strictly conserved and rapidly evolving regions in the TLR-related genes. The genomic segments encoding the intracellular Toll/interleukin 1 receptor domains, which exhibited lower rates of non-synonymous substitution, have undergone purifying selection. In contrast, TLR4, which carried a high proportion of non-synonymous substitutions in the part of extracellular domain spanning 200 amino acids, was found to have been the suggestive target of positive Darwinian selection in primate evolution. However, sequence analyses from 25 primate species, including eight hominoids, six Old World monkeys, eight New World monkeys, and three prosimians, showed no evidence that the pressure of positive Darwinian selection has shaped the pattern of sequence variations in TLR4 among New World monkeys and prosimians. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Scaling of cursoriality in mammals

Data on limb bone lengths from 64 mammalian species were combined with data on 114 bovid species (Scott, '79) to assess the scaling of limb lengths and proportions in mammals ranging from 0.002 to 364 kg. We analyzed log-transformed data using both reduced major axis and least-squares regression to focus on the distribution across mammals of two key traits—limb length and metatarsal/femur ratio—associated with cursorial adaptation. The total lengths of both fore and hindlimbs scale in a manner very close to the M^0.33 predicted by geometric similarity. Thus the relative limb lengths of large mammals, including bovids, generally regarded among the most cursorial of mammals, are very similar to those of the rodents and insectivores in this sample. Metatarsal/femur ratio also shows little change with changing mass, although bovids tend to have relatively longer metapodials than do other families in the sample. We argue that many of the remaining morphological traits associated with cursoriality (e.g., reduction in joint mobility and number of distal limb bone elements) promote cursoriality only at large body sizes. These results lead us to question the general perception that cursoriality is most widespread among large mammals. We also suggest that discussions of cursoriality should focus explicitly on the two partially independent aspects of performance that are otherwise confounded under this general term—speed and the ability to cover substantial distance. © 1993 Wiley-Liss, Inc.     

Frequent segmental sequence exchanges and rapid gene duplication characterize the<Emphasis Type="Italic"> MHC</Emphasis> class I genes in lemurs

Major histocompatibility complex (MHC) class I genes have complicated and profound evolutionary histories. To reconstruct and better understand their histories, partial class I genes (exon 2–intron 2–exon 3) were sequenced in a sampling of prosimians (Strepsirhini, Primates). In total, we detected 117 different sequences from 36 Malagasy prosimians (lemurs) and 1 non-Malagasy prosimian (galago) representing 4 families, 7 genera, and 13 species. Unlike the MHC class II genes (MHC-DRB), MHC class I genes show a generally genus-specific mode of evolution in lemurs. Additionally, no prosimian class I loci were found to be orthologous to HLA genes, even at highly conserved loci (such as HLA-E, HLA-F). Phylogenetic analysis indicates that nucleotide diversity among loci was very small and the persistence time of the polymorphisms was short, suggesting that the origin of the lemur MHC class I genes detected in this study was relatively recent. The evolutionary mode of these genes is similar to that of classic HLA genes, HLA-A, HLA-B, and HLA-C, in terms of their recent origin and rarity of pseudogenes, and differs from them with respect to the degree of gene duplications. From the viewpoint of MHC genes evolution, some interlocus sequence exchanges were apparently observed in the lemur lineage upon phylogenetic and amino acid motif analyses. This is also in contrast to the evolutionary mode of HLA genes, where intralocus exchanges have certainly occurred but few interlocus exchanges have taken place. Consequently, the gene conversion model for explaining the generation of the MHC diversity among different loci can be thought to play more important roles in the evolution of lemur MHC class I genes than in that of HLA genes.Electronic Supplementary Material Supplementary material is available in the online version of this article at