Body size and scaling of the hands and feet of prosimian primates

The hands and feet of primates fulfill a variety of biological roles linked with food acquisition and positional behavior. Current explanations of shape differences in cheiridial morphology among prosimians are closely tied to body size differences. Although numerous studies have examined the relationships between body mass and limb morphology in prosimians, no scaling analysis has specifically considered hand and foot dimensions and intrinsic proportions. In this study, we present such an analysis for a sample of 270 skeletal specimens distributed over eight prosimian families. The degree of association between size and shape was assessed using nonparametric correlational techniques, while the relationship between each ray element length and body mass (from published data and a body mass surrogate) was tested for allometric scaling. Since tarsiers and strepsirrhines encompass many taxa of varying degrees of phylogenetic relatedness, effective degrees of freedom were calculated, and comparisons between families were performed to partially address the problem of statistical nonindependence and "phylogenetic inertia." Correlational analyses indicate negative allometry between relative phalangeal length (as reflected by phalangeal indices) and body mass, except for the pollex and hallux. Thus, as size increases, there is a significant decrease in the relative length of the digits when considering all prosimian taxa sampled. Regression analyses show that while the digital portion of the rays scales isometrically with body mass, the palmar/plantar portion of the rays often scales with positive allometry. Some but not all of these broadly interspecific allometric patterns remain statistically significant when effective degrees of freedom are taken into account. As is often the case in interspecific scaling, comparisons within families show different scaling trends in the cheiridia than those seen across families (i.e., lorisids, indriids, and lemurids exhibit rather different allometries). The interspecific pattern of positive allometry that appears to best characterize the metapodials of prosimians, especially those of the foot, parallels differences found in the morphology of the volar skin. Indeed, relatively longer metapodials appear to covary with flatter and more coalesced volar pads, which in turn slightly improve frictional force for animals that are at a comparative disadvantage while climbing because of their larger mass. Despite the essentially isometric relationship found between digit length and body mass across prosimians, examination of the residual variation reveals that tarsiers and Daubentonia possess, relative to their body sizes, remarkably long fingers. Such marked departures between body size and finger length observed in these particular primates are closely linked with specialized modes of prey acquisition and manipulation involving the hands.     

Athletic anurans: the impact of morphology,ecology and evolution on climbing ability in invasive cane toads

Although primarily terrestrial, cane toads (Rhinella marina) sometimes climb near‐vertical surfaces (tree‐trunks, cliffs, fences) during foraging or dispersal activities. We scored climbing ability (in laboratory trials) of 288 cane toads from four regions in Australia, plus two sites on the island of Hawai'i. We found strong divergence in climbing ability associated not only with sex and relative limb length, but also population of origin. Within each population, longer‐limbed individuals (and hence, males rather than females) were better climbers, although the geographical divergence in climbing ability remained significant even when sex and limb length were included in multivariate regression models. The geographical difference in climbing ability (but not morphology) disappeared when the progeny were raised in captivity under identical conditions, without climbing opportunities. Although influenced by morphology, climbing ability in wild‐caught cane toads appears to be driven primarily by local environmental conditions that facilitate and/or reward arboreal activity.     

The volar skin of primates: its frictional characteristics and their functional significance.

Friction of volar skin on wood is not proportional to load in human beings and prosimians, but to load raised to a fractional power. This meets theoretical expectations for the frictional characteristics of convex elastic surfaces. Although this enables small clawless primates to cling to steeper slopes and larger vertical supports than would otherwise be possible, the magnitude of the effect is not great enough to overcome the disadvantages of clawlessness in climbing vertical or steeply-sloping tree trunks and branches. In human subjects, friction appears to be more nearly proportional to load than in small prosimians used as experimental subjects. It is suggested that this is due to the fact that the small animals have discrete, elevated volar pads. Pad coalescence in large primates may be an adaptation for increasing the power to which load must be raised to become proportional to friction; increasing this exponent yields more friction per unit of adduction force on supports that are small relative to the animals' dimensions and weights.     

Sifaka positional behavior: ontogenetic and quantitative genetic approaches

In many primate species, hands and feet are large relative to neonatal body weight, and they subsequently exhibit negative allometric growth during ontogeny. Here, data are presented showing that this pattern holds for a wild population of lemur, Verreaux's sifaka (Propithecus verreauxi verreauxi). Using morphometric data collected on this population, it is shown that younger animals possess relatively large hands and feet. This ontogenetic pattern suggests a simple behavioral test: do juvenile animals with their larger, almost adult‐sized hands and feet locomote on similarly sized substrates as adult animals? Using locomotor bout sampling, this question was tested by collecting positional behavior data on this population. Results from this test find no differences in locomotor behaviors or substrate use between yearlings and adult animals. To place these results in a broader evolutionary context, heritabilities and selection gradients of hands, feet, and other limb elements for animals in this population were estimated. Among limb elements, heritabilities range from 0.16–0.44, with the foot having the lowest value. Positive directional selection acts most strongly on the foot (directional selection gradient = 0.119). The low heritability and positive selection coefficient indicate that selection has acted, and continues to act, on foot size in young animals. These results are interpreted within a functional context with respect to the development of locomotor coordination: larger feet enable young animals to use “adult‐sized” substrates when they move through their habitat. It is suggested that the widespread pattern of negative allometry of the extremities in sifaka and other primates is maintained by selection, and does not simply reflect a primitive developmental pathway that has no adaptive basis. Am J Phys Anthropol 131:261–271, 2006. © 2006 Wiley‐Liss, Inc.     

Positional behavior of long-tailed macaques (Macaca fascicularis) in northern Sumatra

Although the majority of extant primates are described as "quadrupedal," there is little information available from natural habitats on the locomotor and postural behavior of arboreal primate quadrupeds that are not specialized for leaping. To clarify varieties of quadrupedal movement, a quantitative field study of the positional behavior of a highly arboreal cercopithecine, Macaca fascicularis, was conducted in northern Sumatra. At least 70% of locomotion in travel, foraging, and feeding was movement along continuous substrates by quadrupedalism and vertical climbing. Another 14-25% of locomotion was across substrates by pronograde clambering and vertical clambering. The highest frequency of clambering occurred in foraging for insects, and on the average smaller substrates were used in clambering than during quadrupedal movement. All postural behavior during foraging and feeding was above-substrate, largely sitting. Locomotion across substrates requires grasping branches of diverse orientations, sometimes displaced away from the animal's body. The relatively low frequency of across-substrate locomotion appears consistent with published analyses of cercopithecoid postcranial morphology, indicating specialization for stability of limb joints and use of limbs in parasagittal movements, but confirmation of this association awaits interspecific comparisons that make the distinction between along- and across-substrate forms of locomotion. It is suggested that pronograde clambering as defined in this study was likely a positional mode of considerable importance in the repertoire of Proconsul africanus and is a plausible early stage in the evolution of later hominoid morphology and locomotor behavior.     

Evolution and development of mammalian limb integumentary structures

The adaptive radiation of mammalian clades has involved marked changes in limb morphology that have affected not only the skeleton but also the integumentary structures. For example, didelphid marsupials show distinct differences in nail and claw morphology that are functionally related to the evolution of arboreal, terrestrial, and aquatic foraging behaviors. Vespertilionoid bats have evolved different volar pad structures such as adhesive discs, scales, and skin folds, whereas didelphid marsupials have apical pads covered either with scales, ridges, or small cones. Comparative analysis of pad and claw development reveals subtle differences in mesenchymal and ectodermal patterning underlying interspecific variation in morphology. Analysis of gene expression during pad and claw development reveals that signaling molecules such as Msx1 and Hoxc13 play important roles in the morphogenesis of these integumentary structures. These findings suggest that evolutionary change in the expression of these molecules, and in the response of mesenchymal and ectodermal cells to these signaling factors, may underlie interspecific differences in nail, claw, and volar pad morphology. Evidence from comparative morphology, development, and functional genomics therefore sheds new light on both the patterns and mechanisms of evolutionary change in mammalian limb integumentary structures.     

Locomotion of the Eurasian nuthatch on vertical and horizontal substrates

Legged locomotion of the Eurasian nuthatch Sitta europaea on horizontal and vertical substrates was examined using field observations and experiments. Although previous studies have reported that nuthatches use 'walking' on vertical substrates, we found that they usually used 'hopping' on both vertical and horizontal substrates. When climbing up a vertical substrate, the feet were staggered in position and small phase differences were observed between the left and right leg movements in the gait. In upward climbing, the body was inclined towards the substrate during the first stance phase similar to other tree-trunk climbers, but the tail was not used for helping body rotation unlike most tree-trunk climbers. The staggered position of the feet may allow the legs to play different roles in pulling towards and pushing away from the substrate. In downward climbing, the feet were staggered in position, but the phase difference was quite small. In field observations, the Eurasian nuthatch preferred to move vertically, rather than in an inclined direction.     

Interpreting the Role of Climbing in Primate Locomotor Evolution: Are the Biomechanics of Climbing Influenced by Habitual Substrate Use and Anatomy?

Vertical climbing is widely accepted to have played an important role in the origins of both primate locomotion and of human bipedalism. Yet, only a few researchers have compared climbing mechanics in quadrupedal primates that vary in their degree of arboreality. It is assumed that primates using vertical climbing with a relatively high frequency will have morphological and behavioral specializations that facilitate efficient climbing mechanics. We test this assumption by examining whether time spent habitually engaged in climbing influences locomotor parameters such as footfall sequence, peak forces, and joint excursions during vertical climbing. Previous studies have shown that during climbing, the pronograde and semiterrestrial Macaca fuscata differs in these parameters compared to the more arboreal and highly specialized, antipronograde Ateles geoffroyi. Here, we examine whether a fully arboreal, quadrupedal primate that does not regularly arm-swing will exhibit gait and force distribution patterns intermediate between those of Macaca fuscata and Ateles geoffroyi. We collected footfall sequence, limb peak vertical forces, and 3D hindlimb excursion data for Macaca fascicularis during climbing on a stationary pole instrumented with a force transducer. Results show that footfall sequences are similar between macaque species, whereas peak force distributions and hindlimb excursions for Macaca fascicularis are intermediate between values reported for M. fuscata and Ateles geoffroyi. These results support the notion that time spent climbing is reflected in climbing mechanics, even though morphology may not provide for efficient mechanics, and highlight the important role of arboreal locomotor activity in determining the pathways of primate locomotor evolution.     

Arboreal substrates influence foraging in tropical ants

1. Physically complex substrates impart significant costs on cursorial central‐place foragers in terms of time spent outside the nest and total distance travelled. Ants foraging in trees navigate varied surfaces to access patchy resources, thus providing an appropriate model system for examining interactions between foraging efficiency and substrates. 2. We expected that the speed of recruitment, body size distribution and species richness of foraging arboreal ants would differ predictably among common substrate types occurring on tropical tree trunks. We measured changes in ant abundance and species composition over time at baits placed on bare tree bark, moss‐covered bark, and vine‐like vegetation appressed to bark. We also measured average body size and body size frequency on the three substrate types. Ants discovered baits sooner and accumulated at baits relatively faster when using vine substrates as the primary foraging trail. Average body size was smaller on vine substrates than on bark. Experimental removal of vine and moss substrates nullified these differences. Contrary to our predictions, species richness and body size distributions did not differ among the three substrate types, due in part to the frequent presence of a few common ground‐nesting species at baits on bare bark. 3. Our results collectively indicate that linear substrates facilitate access of foraging ants to patchy resources. Ant use of vine‐like substrates appears to be opportunistic; vine use is not confined to certain species nor constrained by body size.     

Palmar and plantar pads and flexion creases of genetic polydactyly mice (Pdn)

Attempts to gain a better understanding of the relationship between the epidermal ridge patterns (dermatoglyphics) and flexion creases on the volar aspects of human hands and feet and specific medical disorders led to a search for a suitable animal model, allowing studies of the fetal development of the pertinent structures. A common experimental animal, the rat (Rattus norvegicus), was found to be an excellent candidate, owing to the strong resemblance of the volar pads and flexion creases on its palmar and plantar surfaces to those of human subjects. A hereditary preaxial polydactyly mouse (Pdn) provides an opportunity to study the effects of this malformation on the surrounding morphological structures and, specifically, on the volar pads, i.e., the sites over which the dermatoglyphic patterns develop. The hands and feet of the wild‐type (+/+) mice show no anomalies, and their major pad and flexion crease configurations correspond to those of normal rats. The heterozygous (Pdn/+) mice, in spite of having a thumb/big toe with a duplicated distal phalanx on their hands/feet, did not display any alterations in palmar/plantar pads. The homozygous (Pdn/Pdn) mice have a protrusion in the thenar area and one to three supernumerary digits on the preaxial portion of both the hands and feet. The effect of these anomalies was found to be limited to the pad and flexion crease configurations in the preaxial areas; the postaxial sites were not affected. The original number of pads on the thenar/first interdigital areas of Pdn/Pdn mice was apparently identical to that of the +/+ and Pdn/+ mice. The preaxial protrusion, however, affected the number, size, and location of the pads observed in the newborn mice, resulting in varying pad configurations, such as fused and scattered pads or a pad cluster formed by gathering the neighboring pads. These pad modifications were induced by the preaxial plantar/palmar protrusion only and were not affected by the presence of supernumerary preaxial digits. In view of the similarities in the morphology and fetal development of human and mouse distal limbs, the present study is relevant to human subjects, particularly to the understanding of the significance of dermatoglyphic variations in individuals with specific medical disorders. Future studies of naturally occurring or experimentally induced limb malformations in mice or rats should provide valuable insights into the development of human hands and feet and into factors contributing to their congenital anomalies. J. Morphol. 239:87–96, 1999. © 1999 Wiley‐Liss, Inc.     

They Wallop Like They Gallop: Audiovisual Analysis Reveals the Influence of Gait on Buttress Drumming by Wild Chimpanzees (Pan troglodytes)

The role learning plays in the acquisition of communicative gestures by wild chimpanzees (Pan troglodytes) is unclear. We aimed to evaluate the likelihood that social experience influences the structure of chimpanzee buttress drumming displays by examining whether individuals differed in the way they used their hands and feet to strike trees. We analyzed digital video recordings of 245 bouts by 9 adult males from Gombe National Park, Tanzania, frame by frame in conjunction with acoustic analysis. We investigated 1) how limb sequences used to approach drumming trees influenced limb use during drumming, 2) the relative use of hands vs. feet in drumming, and 3) the relative amplitude of beats produced by hands vs. feet. We found that the chimpanzees most often approached trees at a gallop and usually initiated drumming bouts with limb sequences that were identical to gait limb sequences. All individuals produced more beats with their feet than with their hands, and foot beats were higher in relative amplitude than hand beats. In only one instance did an individual produce a bout with hands only, whereas in three of nine observations of drumming on resonant camp equipment, the individuals primarily used their hands rather than their feet. We suggest that although chimpanzees may, by observing others, learn to use buttresses as tools to generate loud sounds, it is unlikely that learning influences the structure of displays because they result from innately determined gait patterns deployed to generate sound from comparatively nonresonant substrates.     

Scale architecture of the palmar and plantar epidermis of Polychrus acutirostris Spix, 1825 (Iguania, Polychrotidae) and its relationship to arboreal locomotion

In tetrapod squamates, the diversity of micro-ornamentations of the epidermis of the contact areas of hands and feet is generally associated with constraints and modalities related to locomotion. Polychrus acutirostris is a medium-sized lizard that occurs in open heterogeneous habitats in South America, such as the cerrados, caatingas, and fallow lands. It progresses slowly on branches of various diameters in its arboreal environment. It can also move more rapidly on the ground. The hands and feet are prehensile and may be considered an adaptation for grasping and climbing. Epidermal surfaces from the palmar and plantar areas of the hands and feet of P. acutirostris were prepared for SEM examination, and studied at various magnifications. They show three major levels of complexity: (1) scale types, organized in gradients of size and imbrication, (2) scalar ornamentations, organized by increasing complexity and polarity, and (3) presence of Oberhäutchen showing typically iguanian honeycomb micro-ornamentations. The shape and surface structure of the scales with their pattern of micro-ornamental peaks, which improve grip, and the grasping hands and feet indicate that P. acutirostris is morpho-functionally specialized for arboreality.     

Locomotion and posture in Lagothrix lagotricha

This long-term study of woolly monkey (Lagothrix) locomotor and postural behaviour employs methods identical to those used during a previous study of the locomotion and posture of two species of Ateles, allowing a detailed comparison between the two genera, which are strong competitors in extensive parts of the Amazon basin and northern Andes. As in Ateles, Lagothrix locomotion can be divided into five patterns, based on limb usage: quadrupedal walking and running, suspensory locomotion, climbing, bipedalism (very rare in wild woolly monkeys) and leaping. Lagothrix differs from Ateles primarily in its greater reliance on quadrupedal locomotion during both travel and feeding and on its de-emphasis of the use of suspensory locomotion as compared to Ateles, while the use of climbing and leaping is roughly equal in the two genera. Lagothrix exhibits more generalised (primitive) locomotive behaviour in accordance with its morphology, in comparison to the more specialised Ateles. The generic differences reflect differences in habitat use and particularly foraging ecology.     

What does "arboreal locomotion" mean exactly and what are the relationships between "climbing", environment and morphology?

The characteristics of "climbing" in the sense of locomotion or posture on three-dimensional substrates are discussed from a biomechanical viewpoint. For this purpose, the mechanical conditions of the most widely spread modes of locomotion or gaits used in arboreal surroundings are reviewed. This allows precise identification of morphological characteristics of traits that are advantageous, and therefore have a positive selective value. Further, at least some of the environmental and substrate characteristics that need to be present for using a specific gait, are noted. It turns out that the extremity which is placed lower on the substrate, has to carry a higher load. If this extremity is consistently the hindlimb--which actually is the case in primates, because of understandable, though complex reasons--a division of labor is likely to occur between the limbs: the hindlimb becoming stronger and the forelimb weaker, but more versatile. A very specific, and advantageous feature of the primates is their possession of prehensile hands and feet. That means the autopodia are able (1) to produce by themselves, without the aid of body weight, very high frictional resistance, and (2) to transmit tensile forces as well as torsional moments on the substrate. The above-mentioned division of labor between fore- and hindlimbs implies that the former make the first contacts with and explore the properties of parts of the environment. As a next step, prehensile hands on long arms may easily replace length and mobility of the neck in getting hold of food items. So very characteristic traits of human body shape can be derived to a large extent from the necessities of arboreal locomotion: Prehensile hands, long arms, concentration of body weight on the hindlimbs, shortness of the trunk in comparison to limb length.     

Tribological Behavior of Gampsocleis Gratiosa Foot Pad Against Vertical Flat Surfaces

Some tribological behavior between mature Gampsocleis gratiosa foot pads and vertical flats of different materials were studied in this work. stereomicroscope （SMS） and scanning electron microscope （SEM） were used to measure the morphology of the Gampsocleis gratiosa foot pads. An atomic force microscope （AFM） was used to measure the morphologies of the surfaces of glass and a wall doped with calcium carbonate material. The attaching behavior of Gampsocleis gratiosa feet on the two vertical surfaces was observed. The attaching force （perpendicular to the vertical surface） and the static frictional force （along the direction of gravitation） of Gampsocleis gratiosa foot pads on a vertical glass were measured. It was shown that the average attaching force is 50.59 mN and the static frictional force is 259.10 mN. The physical models of the attaching interface between Gampsocleis gratiosa foot pads and the two vertical surfaces were proposed. It was observed that the foot pads are smooth in macroscale; however, the pad surface is composed by approximate hexagonal units with sizes of 3 μm to 7 μm in microscale; the adjacent units are separated by nanoscale grooves. The Observations showed that the Gampsocleis gratiosa can not climb the vertical calcium carbonate wall; in contrast, they can easily climb the vertical glass surface. Based on the features of the geometrical morphologies of the foot pads and the glass surface, we speculate that the attaching force and strong static frictional force are attributed to the interinlays between the deformable Gampsocleis gratiosa foot pads and the nanoscale sharp tips of the glass surface.     

Autopodial skeletal diversity in hystricognath rodents: Functional and phylogenetic aspects

Metapodials and phalanges of the second to fourth digital ray were measured for the hands and feet of 214 specimens belonging to 45 extant species of hystricognath rodents, encompassing members of all major clades of the radiation. Principal components analysis (PCA), the phalangeal index of the third digital ray in the hands and feet, and the relationship between second and fourth digital ray were used to investigate intrinsic autopodial proportions as well as to provide a base for comparisons between hands and feet. PCA separated cursorial Hystricognathi from arboreal ones, but lead to little distinction in other locomotory modes. Cursors have longer metapodials and shorter phalanges, particularly in their hind limb, while arboreal species have relatively longer manual and pedal phalanges. Terrestrial, scansorial, fossorial, and semi-aquatic species were not clearly distinguished, but there is a tendency towards elongated manual digits and relatively short feet in most fossorial species. Closely related species with similar locomotory habits tend to group together in PCA morphospace, and also have similar phalangeal indices. The results are in agreement with current hypotheses on locomotory adaptations of the hand and foot, and concur with many previous findings on autopodial proportions in arboreal, cursorial, and fossorial species. They also highlight the limited use of autopodial proportions for inferring systematic affinities. The lack of distinction in the majority of species is likely related to the lack of highly specialized locomotory types in Hystricognathi.     

A dynamic force analysis system for climbing of large primates

Registering substrate reaction forces from primates during climbing requires the design and construction of customized recording devices. The technical difficulties in constructing a reliable apparatus hinder research on the kinetics of primate locomotion. This is unfortunate since arboreal locomotion, especially vertical climbing, is an important component of the hominoid locomotor repertoire. In this technical paper, we describe a custom-built climbing pole that allows recordings of dynamic 3-dimensional forces during locomotion on horizontal and sloping substrates and during vertical climbing. The pole contains an instrumented section that can readily be modified and enables us to register forces of a single limb or multiple limbs in a broad range of primates. For verification, we constructed a similar set-up (which would not be usable for primates) using a conventional force plate. Data for a human subject walking on both set-ups were compared. The experimental set-up records accurate and reliable substrate reaction forces in three orthogonal directions. Because of its adjustability, this type of modular set-up can be used for a great variety of primate studies. When combining such kinetic measurements together with kinematic information, data of great biomechanical value can be generated. These data will hopefully allow biological anthropologists to answer current questions about primate behaviours on vertical substrates.     

Optimal Gait for Bioinspired Climbing Robots Using Dry Adhesion:A Quasi-Static Investigation

Legged robots relying on dry adhesives for vertical climbing are required to preload their feet against the wall to increase contact surface area and consequently maximize adhesion force. Preloading a foot causes a redistribution of forces in the entire robot, including contact forces between the other feet and the wall. An inappropriate redistribution of these forces can cause irreparable detachment of the robot from the vertical surface. This paper investigates an optimal preloading and detaching strategy that minimizes energy consumption, while retaining safety, during locomotion on vertical surfaces. The gait of a six-legged robot is planned using a quasi-static model that takes into account both the structure of the robot and the characteristics of the adhesive material. The latter was modelled from experimental data collected for this paper. A constrained optimization routine is used, and its output is a sequence of optimal posture and motor torque set-points.     

Rheotaxis of Biomphalaria glabrata on vertical substrates and its role in the recolonization of habitats treated with molluscicides

The authors observed specimens of Biomphalaria glabrata climbing up the vertical wall of a ditch against the current. The snails that showed this behavior during application of a molluscicide in the breeding site survived and probably played a role in repopulation, which was observed three months later. These observations motivated field and laboratory investigations which led the authors to conclude that: a) this species is able to climb vertical surfaces both in field and laboratory situations; b) the current of water, as a physical stimulus, is sufficient to trigger this behavior (rheotaxis); c) rheotaxis on vertical surfaces depends on the presence of a necessarily moderate current; d) there are indications that B. glabrata may undergo habituation with respect to rheotaxis on vertical walls; e) the relationship between rheotaxis and habituation should be considered as a factor causing snail grouping in water bodies which may contribute to their localization in the field; f) rheotaxis on vertical surfaces may facilitate population dispersal, and its occurrence should be considered when campaigns for the control of schistosomiasis transmission are planned. The authors present some proposals to avoid the manifestation of this behavior in some field situations.     

Gripping during climbing of arboreal snakes may be safe but not economical

On the steep surfaces that are common in arboreal environments, many types of animals without claws or adhesive structures must use muscular force to generate sufficient normal force to prevent slipping and climb successfully. Unlike many limbed arboreal animals that have discrete gripping regions on the feet, the elongate bodies of snakes allow for considerable modulation of both the size and orientation of the gripping region. We quantified the gripping forces of snakes climbing a vertical cylinder to determine the extent to which their force production favoured economy or safety. Our sample included four boid species and one colubrid. Nearly all of the gripping forces that we observed for each snake exceeded our estimate of the minimum required, and snakes commonly produced more than three times the normal force required to support their body weight. This suggests that a large safety factor to avoid slipping and falling is more important than locomotor economy.