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The Journal of Experimental Biology 204, 1233–1246 (2001) 1233 Printed in Great Britain © The Company of Biologists Limited 2001 JEB3375 SPATIO-TEMPORAL GAIT CHARACTERISTICS OF LEVEL AND VERTICAL LOCOMOTION IN A GROUND-DWELLING AND A CLIMBING GECKO A. ZAAF *, R. VAN DAMME, A. HERREL AND P. AERTS Laboratory for Functional Morphology, Department of Biology, University of Antwerp, Universiteitsplein 1, B-2610 Wilrijk, Belgium *e-mail: firstname.lastname@example.com Accepted 16 January; published on WWW 15 March 2001 Summary The effects of incline (vertical versus horizontal) on substratum. Moreover, gait characteristics differ little spatio-temporal gait characteristics (stride and step between the species despite the clear differences in length, frequency, duty factor, degree of sprawling) were ecological niche. Higher level or climbing speeds are measured over a range of speeds in a ground-dwelling realized mainly (or exclusively in the case of level (Eublepharis macularius) and a climbing (Gekko gecko) locomotion in G. gecko) by increasing stride frequency. species of gecko. Surprisingly, the climbing species also Stride lengths and duty factors vary with speed in the performs very well when moving on the horizontal ground-dweller, but not in the climbing species. Step length substratum. In the present experiments, climbing speeds and the degree of sprawling are speed-independent (except ranged from 0.6 to 1.2 m s−1, whereas speeds for level for hind-limb sprawling in G. gecko on the level). It is locomotion were between 0.6 and 1.8 m s−1. In contrast, the argued that this common strategy suits climbing (fixed vertical climbing capacities of the ground-dweller are spatial variables, no floating phases) rather than level limited (speeds below 0.1 m s−1 versus level speeds between locomotion. 0.2 and 1.1 m s−1). In general, we demonstrate that very little adjustment in gait characteristics is made by either Key words: Gekko gecko, Eublepharis macularius, kinematics, species when they are forced to move on their non-habitual locomotion, climbing, level locomotion, gait. Introduction Many studies of locomotion have documented the presumably beneficial for similar reasons. Claws, adhesive importance of trade-offs in evolutionary adaptation. Different pads, suction cups, a sculptured skin and a flattened body shape environments place different, often conflicting, demands on the are obvious examples of morphological adaptations that allow locomotor apparatus of animals, and species are predicted a scansorial life-style. to evolve morphologies and physiologies that allow a high Besides such obvious morphological adaptations, natural performance capability in their preferred micro-habitat selection might also adjust locomotor behaviour (posture and (Moermond, 1979; Losos, 1990). This may come at the gait) to meet the altered demands imposed by climbing. expense of performance capability in other contexts. For This can occur through subtle design changes (e.g. in joint instance, bats foraging in densely vegetated areas tend to have morphology, musculo-skeletal mechanics, muscle physiology) short, broad wings designed for high manoeuvrability, while and/or through alterations in motor patterns. For instance, bats foraging in open spaces have long, narrow wings to reduce arborealism in chameleons is reflected in their muscle power requirements (Norberg, 1994). architecture and composition (Peterson, 1984; Abu-Ghalyun, For terrestrial tetrapods, the degree of inclination of the 1995) together with a highly specialised pattern of limb substratum may constitute an important environmental design movement (Peterson, 1984; Losos et al., 1993). To achieve factor: an animal moving on a steep or vertical substratum must insight into adaptations and potential trade-offs at this level, generate propulsive forces, not only to overcome inertia (in the one can carry out in-depth morphological, kinesiological and case of unsteady motion) and environmental resistance (from mechanical analyses in an effort to reveal all the mechanistic the air and the substratum), but also to counter gravity (e.g. aspects of the behaviour (e.g. climbing). Alternatively, one can Cartmill, 1985; Zaaf et al., 1999). On a very steep or vertical approach the problem from another perspective. Apart from an incline, the front legs must pull on the substratum to avoid essential descending higher-level control and peripheral backwards tumbling when the hind legs push to provide feedback, locomotor patterns and behaviour are moulded to a propulsion (e.g. Alexander, 1992; Cartmill, 1985), and the large extent by the intrinsic (physical) properties of the entire placement of the centre of mass close to the substratum is locomotor system (electrophysiological dynamics of the 1234 A. ZAAF AND OTHERS neuromuscular components; mechanical dynamics of the avoided, and front leg patterns might differ from those of the musculoskeletal components, etc.) and its interactions with the hind legs, given their different action in vertical climbing (see environment (see also, for instance, Diedrich and Warren, above). A ground-dwelling species can safely modulate its 1995; Diedrich and Warren, 1998a; Diedrich and Warren, speed by changing both the spatial and temporal variables of 1998b; Full and Kubow, 1998; Holt et al., 1990; Holt et al., its gait. (ii) If the climbing strategy is an expression of the 1991; Kugler and Turvey, 1987; Latash, 1998; Schöner et al., intrinsic properties of the system of the climbing species, then 1990; Stewart, 1995; Stewart and Golubitsky, 1992; Thelen we hypothesise that the latter will retain this style of speed and Smith, 1994). The spatio-temporal characteristics and modulation when moving on a level surface. This potentially patterns of the locomotor cycles can therefore be considered as constrains its performance on the level. (iii) Given the higher the collective result of these intrinsic properties (design) and flexibility in speed modulation strategy, the ground-dwelling the dynamics of the locomotor system (e.g. Aerts et al., 2000; species has the opportunity to adjust its locomotor strategy Latash, 1998; McFadyen and Bélanger, 1997; Peck and when climbing. In this case, gravity might affect its Turvey, 1997; Van Damme et al., 1998; Verstappen and Aerts, performance drastically if modulation of the behaviour is 2000; Zernicke and Smith, 1996). Studying and comparing insufficient. spatio-temporal gait characteristics (stride frequency, stride To evaluate these hypotheses, we will test how the spatio- and step length, duty factor, relative phase, etc.) of species with temporal gait characteristics of Eublepharis macularius and widely different locomotor habits (ground-dwelling versus Gekko gecko change with speed, whether they differ between climbing) can thus reveal whether specific differences in species, between substratum gradients (vertical versus 'design' and/or control strategies are present without requiring horizontal) and (given the difference in possible role while an examination of all the intricate details of the morphology or climbing) between the fore and hind legs. the dynamics and laws of all the underlying processes. Lizards of the Gekkota group present ideal opportunities for the study of potential trade-offs between a cursorial Materials and methods (ground-dwelling) and a scansorial (climbing) lifestyle. The Three Gekko gecko (L.) and three Eublepharis macularius Eublepharidae represent the ancestral condition for gekkotans (Blyth) were used in this study. All individuals had similar and are almost exclusively ground-dwelling (Grismer, 1988; snout–vent lengths (Table 1). The animals were obtained from Russell, 1976; Russell, 1979). Many Gekkonidae, in contrast, a commercial dealer and housed in pairs in separate terraria are specialised climbers, living almost exclusively on vertical (60 cm×100 cm×40 cm) on a 12 h:12 h light:dark photoperiod. structures. We have studied two species that differ widely in Ambient temperature varied from 26 °C during the day to 20 °C microhabitat use: the ground-dwelling Eublepharis macularius at night. A heating lamp provided a basking place at a higher and the scansorial Gekko gecko. These two species clearly temperature (40 °C). The animals were provided with food differ in general body shape and posture in relation to their (crickets, mealworms and grasshoppers) and water ad libitum. preferred habitat. Whereas G. gecko has the typical dorso- Spatio-temporal gait variables of climbing and level ventrally flattened shape of climbers, E. macularius has a more locomotion were compared for the specimens. For the cylindrical body shape (Fig. 1). Moreover, G. gecko keeps experiments, a track was constructed consisting of two its body close to the substratum while E. macularius, when removable wooden boxes at each end of a glass tunnel moving, shows a more erect posture (Fig. 1). Furthermore, previous analyses have identified a number of functional Table 1. Morphometric data for the specimens used in this differences in appendicular musculature that were interpreted study as adaptations to their respective microhabitats (Zaaf et al., 1999). Snout–vent Fore-limb Hind-limb The present study focuses on the spatio-temporal gait length Mass length length characteristics (stride length, step length, stride frequency, duty (cm) (g) (cm) (cm) factor, relative phase) to investigate whether the extreme Gekko gecko differences in lifestyle between the two species are reflected in Specimen 1 13.03 57.92 2.97 3.91 these characteristics. The following features are examined. (i) Specimen 2 13.29 61.78 3.45 4.52 Does the manner in which the specialist climber modulates its Specimen 3 12.36 40.01 3.30 3.90 locomotor speed differ from the ancestral strategy displayed by Mean ± S.D. 12.89±0.48 53.24±11.62 3.24±0.25 4.11±0.36 the ground-dwelling species? When climbing vertically, the Eublepharis macularius consequences of an erratic step are likely to be severe, and Specimen 1 13.33 60.11 3.31 4.05 climbing might thus require more precision in terms of leg Specimen 2 12.39 45.31 3.10 3.57 positioning compared with level locomotion. Therefore, it Specimen 3 12.72 43.11 3.25 3.89 would appear to be advantageous to control speed by altering Mean ± S.D. 12.81±0.48 49.51±9.25 3.22±0.11 3.84±0.24 the temporal aspects (frequency, duty factor) only, keeping the spatial variables (stride and step length) constant. A floating Further morphometric data for these species can be found in Zaaf phase (no legs in contact with the substratum) should be et al., 1999. Gait characteristics of geckos 1235 (140 cm×20 cm×15 cm) fixed on a wooden support. The tunnel respectively; see Russell, 1975). We therefore decided to place was large enough to permit free limb and body movements, markers at the level of the metatarsus and metacarpus (needed and its floor was covered with a layer of cork. To study to determine step length and stride length; see below). climbing, the tunnel was mounted vertically. The animals were Level locomotion and climbing sequences were recorded in placed in the box at the bottom of the tunnel and induced to dorsal view using a NAC-1000 high-speed video system set at climb through it towards the other box fixed at the top. When 500 frames s−1. Animals were given at least 15 min rest an animal entered the top box, the boxes were switched and between successive trials. One of the E. macularius specimens the procedure was repeated. To study level locomotion, the was also videotaped while moving along the treadmill with the same arrangement was used, but with the tunnel in a horizontal belt at zero speed. This experiment allowed us to test whether position. To increase the speed range, E. macularius were also the treadmill affects gait variables in these lizards. placed on an adjustable-speed treadmill. For each condition, Only sequences in which the animals moved straight and at animals were trained for 1 week before filming. For the a constant speed were retained for further analysis. From the experiments, all lizards were marked with white non-toxic positions of the marker on the snout tip early and late in these paint dots on the tip of the snout, on the centre of the pectoral sequences, an approximate estimate of speed was obtained. On and pelvic girdles and on the mid-forefoot and mid-hindfoot. the basis of these estimates, nine climbing sequences were In G. gecko, the mid-forefoot and mid-hindfoot are the first selected for two specimens of G. gecko, representing a velocity limb segments that contact the substratum and the last to leave range as wide as possible. Inducing horizontal locomotion was it (before and after the digits touch and leave the substratum, more difficult in this species because the animals often Fig. 1. (A) The ground-dweller Eublepharis macularius (snout–vent length 12.39 cm) and (B) the specialist climber Gekko gecko (snout–vent length 13.03 cm). 1236 A. ZAAF AND OTHERS preferred to move on the side-walls of the corridor or simply Results sat on the side-walls enclosing the belt of the treadmill. Effects of experimental arrangement and inter-individual We obtained useful results for five trials and two trials, differences respectively, for the two specimens used for climbing and eight Preliminary analyses revealed no differences in gait from an additional individual. For the three Eublepharis characteristics between E. macularius moving on the treadmill macularius specimens, seven, five and five level locomotion or in the tunnel in the absence of the treadmill (ANOVA, all sequences were used. One specimen refused to climb. For the P>0.10). As was to be expected from their similarity in overall other two specimens, four and six climbing sequences were size and limb dimensions (Table 1), differences among selected. individuals within species were also not significant (ANOVA, For each of these sequences, the marked body points were all P>0.05). Therefore, we combined data from different digitised frame by frame over a complete locomotor cycle. A experimental arrangements and individuals for further more precise estimate of speed was obtained from the slope of analyses. the forward displacement of the tip of the snout against time The mean body sizes and limbs dimensions of all individuals (linear regression; r2 values were always well above 0.97, used here are very similar (Table 1), so inter-specific indicating that speed was fairly constant throughout the comparisons of spatio-temporal gait variables can be measured stride). Stride length (the distance travelled by the performed without normalisation (i.e. dynamic similarity body during an entire cycle), step length (the distance travelled conditions apply; e.g. Alexander, 1992). when a specific leg contacts the ground), stride frequency (the number of cycles per second) and duty factor (the fraction of Multivariate analyses the cycle that a particular foot is on the ground) were Multivariate analysis of covariance revealed a highly determined according to the methods of Van Damme et al. (Van Damme et al., 1998). Limb angles at touch-down and at lift-off were calculated as 0.18 the angle between a line connecting the foot/hand with the centre of the respective girdle and an axis through the girdle 0.16 perpendicular to the mid-sagittal plane (straight posture). Limb 0.14 angles in front of the perpendicular axis are considered Stride length (m) positive; behind this axis, they are negative. 0.12 The degree of sprawling was determined by measuring the distance between the markers on the pelvic or pectoral girdle 0.10 and the mid-hindfoot/forefoot in stance when the forefoot and the hindfoot are directly lateral to the shoulder and hip, 0.08 respectively. 0.06 The relative phase was calculated for the fore-limb, using touch-down of the ipsilateral hind-limb as the reference time 0.04 (relative phase 0 °). The relative phase was expressed as the 0 1 2 relative timing (within the cycle) of fore-limb touch-down 12 multiplied by the stride frequency. To analyse differences in limb kinematics between species and substrata, we first ran a multiple analysis of covariance 10 (MANCOVA) with the gait characteristics as the dependent Frequency (Hz) variables, species and substratum as the factors and speed as a 8 covariate. Testing all characteristics simultaneously was not possible because of a lack of degrees of freedom. We therefore 6 performed two MANCOVAs, one for each pair of limbs. Because we also wanted to estimate species and substratum 4 differences in speed modulation strategies, we proceeded with univariate tests on each gait characteristic. Relationships 2 between speed and spatio-temporal gait characteristics were established by means of least-squares linear regression 0 0 1 2 analysis. When gait characteristics changed with velocity, analysis of covariance (ANCOVA, velocity entered as Velocity (m s-1) covariate) was used to assess inter-specific differences, Fig. 2. Stride lengths and stride frequencies for Gekko gecko differences between fore- and hind-limbs and the effect of (triangles) and Eublepharis macularius (circles) moving at different inclination (horizontal versus vertical). Otherwise, differences velocities on vertical (filled symbols) and horizontal (open symbols) were tested using t-tests. substrata. Gait characteristics of geckos 1237 significant species × substratum interaction effect in both hind- (Table 2; Fig. 5). The relative phase of the fore-limb did not limb (Wilk's λ=0.34, d.f.=7,53, P<0.0001) and fore-limb change with speed and equalled 0.5 (i.e. diagonal pairs move (Wilk's λ=0.18, d.f.=7, 53, P<0.0001) gait characteristics. This in synchrony; Table 2; Fig. 6). suggests that the effect of inclination on the kinematics of On the vertical substratum, both stride length and stride locomotion differed between the two species. frequency increased with increased velocity (Table 2; Fig. 2), The significant species × substratum interaction effect could but the change in stride length was relatively small in be a statistical artefact of the extremely low velocities attained comparison with the change in stride frequency. For instance, by E. macularius on the vertical substratum. We therefore as calculated from the regression equations in Table 2, the present the results of univariate tests for each species below. stride frequency of climbing G. gecko changed by 75 % as speed increased from 0.5 to 1.0 m s−1. Over the same velocity Gait characteristics and speed interval, stride length increased by 13 %. As for level We were able to measure the gait characteristics of G. gecko locomotion, step length, limb angle at lift-off and touch-down moving at velocities on the level between 0.6 and 1.8 m s−1 and and duty factor were independent of speed in climbing G. for climbing between 0.6 and 1.2 m s−1. For E. macularius, we gecko (Table 2; Figs 3, 4). The hind-limbs were placed more obtained level data velocities between 0.24 and 1.05 m s−1. laterally at higher speeds (i.e. increased sprawling), but this Eublepharis macularius proved to be a poor climber, and we was not the case for the fore-limbs (Table 2; Fig. 5). The were therefore able to assess climbing gait characteristics for relative fore-limb phase (0.5) was independent of speed a small range of low velocities (0.025–0.085 m s−1) only. (Table 2; Fig. 6). On the level, G. gecko increased its velocity by increasing Ground-dwelling E. macularius increased both stride its stride frequency. Stride length, step length, limb angle at frequency and stride length to increase velocity (Table 3; lift-off and at touch-down and duty factor did not change with Fig. 2). From the equations in Table 3, a velocity change from speed (Table 2; Figs 2–4). The hind-limbs were placed more 0.5 to 1.0 m s−1 involved an 18 % increase in stride length and sagittally at higher speeds (i.e. a reduction in the degree of a 69 % increase in stride frequency. Duty factor in ground- sprawling), but this was not the case for the fore-limbs dwelling E. macularius decreased with velocity (Table 3; Table 2. Relationships between gait characteristics and velocity for Gekko gecko moving on horizontal and vertical substrata Level locomotion (N=15) Climbing (N=18) r2 a b r2 a b Stride length (m) Hind-limbs 0.018 − − 0.29* −0.802±0.006 0.191±0.074 Step length (m) Fore-limbs 0.02 − − 0.00001 − − Hind-limbs 0.02 − − 0.00004 − − Stride frequency (Hz) Hind-limbs 0.90*** 0.805±0.012 0.945±0.085 0.87*** 0.798±0.006 0.753±0.072 Duty factor Fore-limbs 0.09 − − 0.11 − − Hind-limbs 0.02 − − 0.008 − − Sprawling (m) Fore-limbs 0.036 − − 0.130 − − Hind-limbs 0.293* −1.489±0.015 −0.237±0.102 0.336* −1.379±0.006 0.199±0.070 Limb angle at touch-down (degrees) Fore-limbs 0.005 − − 0.034 − − Hind-limbs 0.0003 − − 0.019 − − Limb angle at lift-off (degrees) Fore-limbs 0.066 − − 0.034 − − Hind-limbs 0.0563 − − 0.024 − − Relative phase 0.185 − − 0.156 − − Intercepts and slopes (±S.E.M.) of least-squares regressions [(log10(gait characteristic)=a+blog10(velocity)] are given for those relationships that had significant r2 values. Velocity is expressed in m s−1, stride length and step length in m and stride frequency in Hz. *Significant at P=0.05; ***Significant at P=0.001. 1238 A. ZAAF AND OTHERS 0.10 0.10 Fore-limbs Hind-limbs 0.09 0.09 Step length (m) 0.08 Step length (m) 0.08 0.07 0.07 0.06 0.06 0.05 0.05 0.04 0.04 0 1 2 0 1 2 1.0 1.0 0.9 0.9 0.8 0.8 Duty factor Duty factor 0.7 0.7 0.6 0.6 0.5 0.5 Fig. 3. Step lengths and duty factors for Gekko gecko (triangles) and 0.4 0.4 Eublepharis macularius (circles) 0.3 0.3 moving at different velocities on 0 1 2 0 1 2 vertical (filled symbols) and horizontal (open symbols) substrata. Velocity (m s-1) Velocity (m s-1) Fig. 3). Step length and limb angle at lift-off and touch-down hind-limbs were 4.58 % smaller than those of the fore-limbs of did not change with velocity, and neither did the degree of the same cycle. For the duty factor, this difference amounted sprawling (Table 3; Figs 3–5). Like the climbing species, the to an average of 8.55 %. For level locomotion in G. gecko, the relative fore-limb phase (0.5) did not change with speed (Table degree of sprawling was consistently higher in the hind-limbs 3; Fig. 6). than in the fore-limbs (t14=5.78, P<0.0001). For climbing G. Because of the limited velocity range obtained in our gecko, the degree of sprawling was consistently higher in the experiments with climbing E. macularius, we will not examine fore-limbs than in the hind-limbs (t17=2.70, P=0.01). the effects of speed on gait characteristics for climbing in this For level data in E. macularius, hind-limb step lengths were species quantitatively. consistently larger than those for the fore-limbs of the same cycle (paired t-tests, t21=3.22, P=0.004). The average within- Hind-limb versus fore-limb gait characteristics cycle difference amounted to 2.7 %. Despite the larger hind- For symmetrical gaits, steady locomotion should normally limb steps, the fore-limb angle at touch-down was larger than yield identical stride lengths and frequencies for the fore- and that of the hind-limb (t-test, t21=6.36, P<0.0001), but the hind-limb cycles (this was confirmed in preliminary tests). limb angle at lift-off was similar (P>0.8). Duty factors did Here, we compare the relevant gait characteristics of the hind- not differ between fore- and hind-limbs in ground-dwelling and fore-limbs within one cycle. For level locomotion in G. E. macularius (paired t-tests, both P>0.18). In climbing gecko, most gait characteristics for the hind-limb did not differ E. macularius, the step length of the hind-limb differed significantly from those for the fore-limb (paired t-tests, all significantly from the step length of the fore-limb (paired t-test: P>0.06). Only the hind-limb angle at lift-off was significantly t9=2.36, P=0.04). On average, the step lengths of the hind- smaller than that observed for the fore-limb (t1,14=8.09, limbs were 9.8 % smaller than those of the fore-limbs of the P<0.0001). During climbing, however, limb angles at lift-off same cycle. This is because hind-limb angle at touch-down and and touch-down did not differ between the front and hind leg lift-off tended to be smaller than those of the fore-limb when (t-test, both P>0.1). Moreover, the step lengths and duty factors climbing (paired t-test, t9=5.46, P<0.0001 for limb angle at of the fore-limbs were consistently larger than those of the touch-down, and t9=2.58, P=0.03 for limb angle at lift-off). hind-limbs (paired t-tests, step length, t17=3.91, P=0.001; duty In both vertical and horizontal locomotion, the degree of factor, t17=6.94, P<0.0001). On average, the step lengths of the sprawling was higher in the hind-limbs (paired t-test, Gait characteristics of geckos 1239 90 90 Hind-limbs Limb angle at touch-down (degrees) Fore-limbs Forelimbs Hindlimbs Limb angle at touch-down (degrees) 80 80 70 70 60 60 50 50 40 40 30 30 20 20 10 10 0 1 2 0 1 2 -20 -20 Limb angle at lift-off (degrees) Limb angle at lift-off (degrees) -30 -30 -40 -40 -50 -50 -60 -60 Fig. 4. Limb angles at touch-down (upper graphs) and at lift-off (lower -70 -70 graphs) for the fore- (left side) and hind-limbs (right side) of Gekko -80 -80 gecko (triangles) and Eublepharis macularius (circles) while climbing -90 -90 0 1 2 0 1 2 (filled symbols) and moving on a horizontal substratum (open symbols). Velocity (m s-1) Velocity (m s-1) horizontal, t21=−7.51, P<0.0001; vertical, t21=−6.26, The stride frequency of the hind-limb increased slightly P<0.0001). more rapidly with speed during level locomotion than during climbing (ANCOVA, difference between slopes F1,27=4.27, Climbing versus level locomotion P=0.048). For G. gecko, regression lines relating hind-limb stride Hind-limb step lengths did not differ between climbing and length and velocity for vertical and horizontal locomotion did level locomotion in G. gecko (t-test, t31=0.77, P=0.45), but the not differ in slope (ANCOVA, F1,27=0.89, P=0.35), but there fore-limb step lengths of climbing animals were larger was a difference in intercepts (F1,28=6.87, P=0.014). On than those measured for animals moving on the horizontal average, stride lengths were smaller for horizontal than for substratum (t31=2.62, P=0.01). Fore-limb angles at touch- vertical locomotion. down and hind-limb angles at lift-off appear not to be affected 0.050 0.050 Fore-limbs Hind-limbs 0.045 0.045 0.040 0.040 Sprawling (m) Sprawling (m) 0.035 0.035 0.030 0.030 0.025 0.025 Fig. 5. The degree of sprawling 0.020 0.020 of Gekko gecko (triangles) and Eublepharis macularius (circles) 0.015 0.015 moving at different velocities on vertical (filled symbols) 0.010 0.010 0 1 2 0 1 2 and horizontal (open symbols) substrata. Velocity (m s-1) Velocity (m s-1) 1240 A. ZAAF AND OTHERS 1.0 Table 3. Relationships between gait characteristics and 0.9 velocity for Eublepharis macularius moving on a horizontal substratum 0.8 Level locomotion (N=22) 0.7 Relative phase r2 a b 0.6 Stride length (m) 0.5 Fore-limbs 0.72*** −0.832±0.011 0.2140±0.030 0.4 Hind-limbs 0.71*** −0.812±0.012 0.241±0.034 0.3 Step length (m) Fore-limbs 0.004 − − 0.2 Hind-limbs 0.007 − − 0.1 Stride frequency (Hz) 0 1 2 Fore-limbs 0.97*** 0.819±0.011 0.758±0.031 Velocity (m s-1) Hind-limbs 0.97*** 0.819±0.011 0.757±0.031 Fig. 6. Relative phase of the fore-limbs versus velocity for Gekko Duty factor gecko (triangles) and Eublepharis macularius (circles) while Fore-limbs 0.26** −0.240±0.017 −0.126±0.047 climbing (filled symbols) and moving on a horizontal substratum Hind-limbs 0.61*** −0.247±0.013 −0.201±0.036 (open symbols). Sprawling (m) Fore-limbs 0.025 − − Hind-limbs 0.079 − − by inclination (t-test, P>0.2), but both fore-limb angle at lift- off and hind-limb angle at touch-down are larger during level Limb angle at touch-down (degrees) locomotion (t-test, t30=6.62, P<0.0001 for fore-limb angle at Fore-limbs 0.015 − − lift-off, and t30=2.75, P<0.01 for hind-limb angle at touch- Hind-limbs 0.000015 − − down). Limb angle at lift-off (degrees) The degree of sprawling of the fore-limbs is conspicuously Fore-limbs 0.007 − − smaller on the level surface than during climbing (t-test, Hind-limbs 0.0007 − − t31=12.25, P<0.0001). The regression lines relating hind-limb Relative phase 0.166 − − sprawling and velocity differ in intercept (ANCOVA, F1,28=33.23, P<0.0001) but not in slope (F1,27=0.89, P=0.35). Slopes and intercepts (±S.E.M.) of least-squares regressions On average, the hind-limbs are more spread on the vertical [log10(gait characteristic)=a+blog10(velocity)] are given for those substratum than on the horizontal one (Table 4). relationships that had significant r2 values. The duty factors of climbing G. gecko were on average Velocity is expressed in m s−1, stride length and step length in m lower than those of animals moving on the horizontal and stride frequency in Hz. substratum (hind-limbs, t31=3.22, P=0.003; fore-limbs, **Significant at P=0.01; ***Significant at P=0.001. t31=2.26, P=0.03). Although the relative fore-limb phase was lower on the level (t-test, t31=2.45, P=0.02) than on the vertical greater on the horizontal substratum than when climbing (t- substratum, relative phase values were close to 0.5 in both test: t30=2.50, P=0.01), but that of the hind-limbs did not differ conditions (Table 4). between the locomotor substrata (t-test: P>0.3). The limited velocity range obtained for climbing E. macularius precludes statistical analysis of differences in Inter-specific differences in gait characteristics speed modulation strategies between horizontal and vertical Because of the large difference in velocity ranges for which locomotion, but Figs 2 and 3 suggest that an animal moving we obtained data on climbing gait characteristics in both on the horizontal substratum at speeds as low as those realized species, we did not analyse inter-specific differences in stride during climbing would show stride lengths, stride frequencies length and stride frequency for climbing. and duty factors similar to those measured for climbing. The When moving on the horizontal substratum, the mean step lengths of the fore-limbs were smaller during horizontal step length (t-test, fore-limbs, t35=0.22, P=0.83; hind-limbs, locomotion than when climbing (t-test: t30=2.041, P=0.05). t35=2.02, P=0.051) and the degree of sprawling (t-tests, fore- The opposite was found for hind-limb step length (t-test: limbs, t35=0.44, P=0.69; hind-limbs, t35=−0.67, P=0.50) did t30=2.57, P=0.02). The relative fore-limb phase also differed not differ between the species (Table 4). between level locomotion and climbing (t-test: t30=3.16, The slopes of the regressions of stride length on speed (both P=0.003). When climbing, E. macularius mainly shifts to a log-transformed) differed significantly between the two species 'lateral single foot' gait pattern (see Hildebrand, 1985) using (ANCOVA: hind-limbs, F1,33=7.29, P=0.011). While E. a relative phase between 0.15 and 0.35 (mean phase 0.28; macularius increased its stride length to increase its speed on Table 4; Fig. 6). The degree of sprawling of the fore-limbs was the level, stride length was unchanged in G. gecko. Gait characteristics of geckos 1241 Table 4. Comparison between the gait characteristics of the fore- and hind-limbs in Gekko gecko and Eublepharis macularius while climbing and moving on a horizontal substratum E. macularius G. gecko Fore-limbs Hind-limbs Fore-limbs Hind-limbs Level Stride length (m) 0.127±0.015 0.130±0.017 0.147±0.014 0.145±0.013 Step length (m) 0.066±0.007 0.072±0.006 0.066±0.005 0.068±0.004 Stride frequency (Hz) 4.076±1.436 4.063±1.143 7.063±2.477 6.880±2.313 Duty factor 0.632±0.063 0.657±0.074 0.579±0.036 0.548±0.045 Sprawling (m) 0.025±0.004 0.034±0.005 0.025±0.005 0.033±0.005 Limb angle at touch-down (degrees) 62.88±62.879 51.0±9.533 56.73±10.048 58.452±9.608 Limb angle at lift-off (degrees) −61.93±12.372 −62.29±5.739 −64.51±7.880 −50.975±7.527 Relative phase 0.470±0.067 0.456±0.052 Climbing Stride length (m) 0.075±0.009 0.084±0.015 0.156±0.007 0.155±0.009 Step length (m) 0.072±0.009 0.065±0.007 0.070±0.003 0.067±0.002 Stride frequency (m) 0.537±0.019 0.553±0.226 5.830±0.822 5.857±0.784 Duty factor 0.881±0.030 0.865±0.074 0.555±0.023 0.507±0.027 Sprawling (m) 0.020±0.004 0.032±0.005 0.043±0.002 0.041±0.002 Limb angle at touch-down (degrees) 67.44±8.198 42.24±3.52 52.34±4.133 47.57±12.78 Limb angle at lift-off (degrees) −66.62±12.81 −59.65±15.06 −51.49±2.47 −48.93±12.983 Relative phase 0.280±0.103 0.490±0.024 Values are means ± S.E.M. (not log-transformed). The slopes of the regressions of stride frequency on level avoided). If not, the force of gravity will reduce or even reverse speed (both log-transformed) also differed between the two the upward momentum of the lizard's body in every stride. species (ANCOVA: hind-limbs, F1,33=5.83, P=0.021). Stride From a theoretical point of view, it can even be argued that at frequency increased more rapidly with speed in G. gecko than least one front leg must exert pulling forces at any time to avoid in E. macularius. backwards tumbling due to gravity. Ground-dwelling species, Finally, for the hind-limbs, the slopes of the regressions of in contrast, do not have these constraints. In theory, they can duty factor on speed (both log-transformed) differed between safely modulate their speed by changing both the spatial and the species (ANCOVA: F1,33=10.46, P=0.003). The duty temporal variables of their gait. factor decreased with increasing speed in E. macularius, but According to Russell (Russell, 1976; Russell, 1979), ground remained constant in G. gecko. Unlike the hind-limbs, no dwelling represents the ancestral state within the gekkotans. differences were found between the slopes (ANCOVA: Therefore, we will first evaluate the locomotion strategy F1,33=0.78, P=0.38) or the intercepts (F1,34=0.02, P=0.88) of displayed by E. macularius on the horizontal and compare it the regression equations of duty factor of the forelimb on speed with data available from other non-specialist climbers in order in the two species. to compare this strategy with that of the specialist climber. Legged animals can increase their velocity by taking larger strides, by increasing their stride frequency or by using a Discussion combination of these two strategies. The degree to which stride Three working hypotheses are tested in this study (see length and/or frequency are modulated with respect to speed Introduction). First, does the speed modulation strategy of the seems to vary among different non-climbing lizard species climber G. gecko differ from that of the ground-dwelling E. (White and Anderson, 1994; see Fig. 7). Although a direct macularius? It is postulated that in the climber only the inter-specific comparison is hampered by methodological temporal aspects (frequency, duty factor) of the gait will differences and by differences in the range of velocities over change with speed, whereas the spatial variables (stride and which the gait characteristics were measured (Fig. 7), all step length) will remain constant. Such a speed modulation previously studied lizards seem to modulate their speed using strategy might ensure the accuracy of leg positioning when a combination of changes in stride frequency and stride length moving faster, a requirement that can be assumed to be (Avery et al., 1987; White and Anderson, 1994; Reilly and particularly important when climbing vertical structures. Delancey, 1997; Fieler and Jayne, 1998; Irschick and Jayne, Moreover, throughout the stride, at least one of the legs should 1998; Van Damme et al., 1998; Fig. 7). always remain in contact with the substratum to deliver an On its preferred horizontal substratum, E. macularius shows upward force to counter gravity (i.e. a floating phase must be a speed modulation strategy comparable with those of the other 1242 A. ZAAF AND OTHERS 0.5 20 Pa 0.4 Dd Ph 15 Us Stride frequency (Hz) Dd Stride length (m) Cd Cd 0.3 Gg Pp Dd Ve Us 10 0.2 Pp Dd Em Em Gg Sc Sc 5 0.1 Ph Pa Ve 0 0 0 1 2 3 4 5 0 1 2 3 4 5 0.16 0.9 Ve 0.14 0.8 Dd 0.12 0.7 Ve Step length (m) 0.10 0.6 Em Duty factor Sc Gg Dd 0.8 Cd 0.5 Em Pa Pp 0.6 Gg 0.4 Ph Pp Dd 0.4 Pa Us 0.3 Ph 0.2 Dd Cd 0.2 Us 0 0.1 0 1 2 3 4 5 0 1 2 3 4 5 Velocity (m s-1) Velocity (m s-1) Fig. 7. Gait characteristics for moving on a horizontal substratum at different speeds for Gekko gecko (Gg) and Eublepharis macularius (Em) compared with data for other lizards (Ve, Varanus exanthematicus, Jayne et al., 1990; Dd, Dipsosaurus dorsalis, Fieler and Jayne, 1998; Sc, Sceloporus clarkii, Reilly and Delancey, 1997; Pa, Podarcis hispanica atrata, Van Damme et al., 1998; Ph, Podarcis hispanica hispanica, Van Damme et al., 1998). Also indicated are the gait characteristics of Phrynosoma platyrhinos (Pp), Uma scoparia (Us), Callisaurus draconoides (Cd) and Dipsosaurus dorsalis (Dd) running at maximal speed (Irschick and Jayne, 1999). ground-dwelling lizards (Fig. 7). Both stride length and stride see Fig. 7) and also other tetrapods (e.g. walking cats and dogs, frequency increase significantly with speed, although faster McMahon, 1984) show a constant step length over a range of locomotion is achieved mainly by increasing the stride speeds. frequency. To increase stride length, step length may be Combined with the constant step length, however, the increased and/or the duty factor may be reduced (less overlap absence of a floating phase in this ground-dweller is in the support by several legs, leading to the incorporation of remarkable because this seriously constrains stride length and, a floating phase in the stride). E. macularius keeps its step hence, maximal speed. In ground-dwelling animals, no reasons length constant (see Table 3) and avoids floating phases in its seem to exist to exclude floating phases from the running strides. Only occasionally do duty factors drop somewhat strides, and many lizards do show floating, especially at higher below 0.5. The fact that step length does not change with speed speeds (e.g. Van Damme et al., 1998; P. Aerts and R. Van suggests that this variable is maximised at all speeds, being set Damme, personal observations). At first glance, the by morphological and mechanical constraints (leg length, evolutionary retention of 'climbing control' (in which contact girdle rotation, maximal excursions at the joints between leg with the substratum is essential; see above) cannot be put segments). Such a constant step length is not exceptional. forward as a possible explanation because it is assumed that E. Other lizard species (e.g. lacertids, Van Damme et al., 1998; macularius represents the ancestral, ground-dwelling condition Gait characteristics of geckos 1243 of the gekkotans. This implies that E. macularius is either constant and probably avoids floating phases in its strides. unwilling to include a floating phase into its limb cycle (at least Extrapolation of the regression equation for duty factor in our experiments) or cannot do so because of biomechanical (Table 3) to the maximal speed of 1.8 m s−1 predicts a duty constraints (floating phases will only occur when the push-off factor of 0.5! These two features are attributable to a specialist forces exceed a certain level). This may at least partly explain climber (see above). So, does the locomotor style of E. why the maximal velocity on the level is relatively low for E. macularius resemble the presumed ancestral state of a macularius compared with the maximal velocities attained by horizontal runner or were the ancestral gekkotans (as suggested other species of similar size (as illustrated in Fig. 7). As an by Russell, 1976; Russel, 1979) climbers, although not alternative explanation, it could be argued that the absence of necessarily as specialised as G. gecko, rather than ground- a floating phase is an artefact of the actual measured velocity dwellers? It could also be argued that the strategy used by both range being too narrow and that further stride length increase species represents a basic non-specialised locomotor behaviour, by including floating might have occurred if higher speeds had which can be considered as an exaptation for highly specialised been recorded. The maximum speed of E. macularius we have climbing. In this respect, it is worthwhile considering the recorded was approximately 1.8 m s−1 (measured on a race- modulation strategy of the two Podarcis species plotted in track; A. Zaaf and R. Van Damme in preparation), which is Fig. 7 (see also Van Damme et al., 1998). Both species higher than the maximal speeds attained during the present resemble the gekkotans of the present study fairly well: stride experiments (approximately 1.1 m s−1). length modulation is small compared with frequency This unexpected aspect of the speed modulation strategy of modulation and step length is kept contant. Nevertheless, one the ground-dwelling species means that nearly all the spatio- species is a ground-dweller (P. h. atrata), whereas the other temporal variables of the specialist-climber's strides resemble must be considered as a climber (P. h. hispanica). However, it the presumed ancestral style rather well when performing on is still possible that climbing represents the primitive state in the preferred vertical substratum! Both stride length and this case too (see Van Damme et al., 1998). frequency increase significantly when G. gecko climbs faster. The second working hypothesis formulated above stated that Moreover, this occurs in a very similar manner to that observed G. gecko retains its climbing style of speed modulation when for the velocity increase by the ground-dwelling species moving on a level surface because this style is an expression (compare the regression constants of the equations relating of the intrinsic properties of the system, leaving no room for stride length and frequency to speed; see Tables 2, 3). behavioural modulation. From this perspective, these intrinsic Furthermore, step length remains constant in both species. properties are considered to be tuned by natural selection to a Therefore, it is likely that the invariability of the duty factor, specialised climbing life-style. In general, we must conclude as it appears from the regression model (Table 2), is a that G. gecko performs on the level surface in the same way as statistical artefact resulting from the fact that we did not record when it climbs: i.e. frequency-modulated, with a constant step low-speed climbing by G. gecko in our experiments (Fig. 3; length and keeping at least one (fore) limb always on the Table 2). Indeed, it is impossible to increase stride length while ground. keeping both step length and duty factor constant. However, as Compared with climbing, stride lengths are somewhat in E. macularius, duty factors rarely drop below 0.5. For the smaller on the level substratum, while frequencies are higher. fore-limbs, they were always above 0.5, whereas those of the But, in this case and in contrast to the climbing results, stride hind-limb were below 0.5 for only a minor fraction of the cycle length does not change with speed. The constant step lengths (less than 4 %; see Fig. 3). This conforms to our predictions are identical to those used in climbing, which provides further concerning the constraints on climbing: a front leg is always support for the suggestion that this variable is maximized at all in contact with the substratum, and floating phases are absent. speeds and is set by morphological and mechanical constraints In practice, two diagonal legs are always in contact with the (leg length, girdle rotation, maximal excursions at the joints substratum, except for those rare cases in which hind-limb duty between leg segments, etc.). On only a few occasions do the factors fall below 0.5. duty factors of the hind legs drop below 0.5. In all other cases, Thus, our results show that G. gecko uses a speed modulation even at the highest speeds, they remain well above this value strategy, both during climbing and when moving on level (see Fig. 3). The degree of sprawling differs between climbing surface, that is very similar to that of the level runner E. and level locomotion (see Table 4). On the level, sprawling is macularius. This is because the latter species seems to apply a reduced, with the front legs, especially, placed nearer the body. locomotor strategy that conforms more to that predicted for a As a result, the body is presumably lifted from the substratum. climber than to that for a level runner. Indeed, although stride To climb vertically, it is essential to keep the body close to the length increases significantly with speed in both species, it is substratum (see above). Since gravity acts parallel to a vertical mainly increases in cycle frequency that achieve speed surface, this does not necessarily imply significant frictional increases: on their preferred substratum, both species apply a forces. On the level, the body weight would induce frictional gain factor of 1.26 to stride length, but a factor of 2.28 to forces unless G. gecko lifts its body from the ground. frequency, when the speed triples (from 0.5 to 1.5 m s−1; That smaller stride lengths were observed in G. gecko when obtained from the regression equations of the hind legs in moving on the horizontal is remarkable because, theoretically, Tables 2, 3). Moreover, E. macularius keeps its step length there are no constraints on stride length when moving on a 1244 A. ZAAF AND OTHERS horizontal substratum (see above). Therefore, stride length range 2.5–8.5 cm s−1 gives predicted stride lengths ranging would be expected to be at least equal to that for similar from 6 to 9 cm, frequencies between 0.4 and 1.02 Hz and duty climbing speeds. Zaaf et al. (Zaaf et al., 1999) showed that the factors of 0.92–0.79. These predicted values coincide muscular system of G. gecko appears to be fine-tuned to its strikingly well with our measurements made for climbing (see climbing life-style. It is therefore not inconceivable that the Figs 2, 3). Step lengths are independent of speed but change altered leg configuration required to lift the body above the slightly with substratum orientation (the step lengths of the substratum during horizontal movement constrains muscle fore-limbs were longer when climbing, those of the hind-limbs performance (or joint motions) in the stride direction, resulting were shorter; see Table 4). in a decrease in the stride length. Since step length does not A major difference seems to be that E. macularius shifts change with substratum orientation, this must express itself from a walking trot to a 'lateral single foot' sequence when through an extended overlap in ground contact of diagonal moving from a horizontal to a vertical substratum. This is pairs of legs. Indeed, the average duty factors are somewhat obvious from the relative phase between the fore- and hind- larger during level locomotion (see Table 4). limbs. Again, however, this apparent change in style might be So, concerning this second hypothesis, we must conclude the result of the very low speeds recorded rather then being that the basic coordination of the limb movements does not induced by the difference in incline. When moving slowly, change when G. gecko changes to a novel substratum most tetrapods (including lizards and salamanders) use this orientation. This basic coordination resembles that of the sequence of footfalls because it provides the highest stability ground-dwelling E. macularius, since climbing in the former (Hildebrand, 1985; Hildebrand, 1988). Only when moving species was fairly similar to level locomotion in the latter faster do they change to a walking or running trot because the (see above). Provided that level locomotion is the ancestral time then available for the centre of mass to move away from locomotor state of the gekkotans (Grismer, 1988; Russell, the diagonal line of support becomes too short. 1976; Russell, 1979) and that the strategy applied by both Unlike in G. gecko, sprawling in E. macularius appears to species represents a basic non-specialised locomotor behaviour decrease slightly on the vertical substratum (not statistically (see above), G. gecko (or specialist climbers in general) significant for the hind-limb; see Table 4). A qualitative apparently retained this strategy with minor alterations when evaluation of the video sequences of climbing E. macularius adopting their new life-style. In this context, the ancestral suggests that this reduction is because the specimens try to locomotor state must be considered an exaptation for climbing minimize the distance between the body and the substratum in allowing for superb performance as soon as adhesive pads an alternative way. The intra-leg configurations during stance emerged. However, assuming a scansorial life-style as the differ from those observed during level locomotion: the legs ancestral state (Grismer, 1988; Russell, 1976; Russell, 1979; are kept close to the sides of the body, with the shoulder and see above), it can be hypothesized that the intrinsic properties hip lowered below the elbow and knee respectively (i.e. the of the locomotor system are so attuned to this life-style that the elbows and knees point upwards). It is surprising that this specific spatio-temporal behaviour (i.e. the collective result drastic change does not affect the step length to a larger extent. of these properties) is inevitably exhibited on whatever This provides support for the suggestions that gekkotans substratum the animals perform. This suggestion would be maximize their step length when possible and that this step disproved if E. macularius alters its spatio-temporal behaviour length is determined mainly by rotational constraints at the when climbing (see below). level of the hip and shoulder joint. The ground-dwelling species, E. macularius, has potentially To summarise, it appears that the overall pattern of more flexibility in its speed modulation strategy since the coordination in climbing E. macularius does not differ from its constraints associated with vertical climbing are absent. This level locomotion strategy. This is not really unexpected increased flexibility should allow this species to adjust its because the level locomotion style already corresponds to that strategy when climbing. From the results and the above presumably appropriate for climbing (see above). However, it discussion, it is clear that this does not occur: overall, ground- is obvious that a vertical incline presents E. macularius with dwelling E. macularius modulate their speed in a similar way serious problems. When observing animals climbing, it is clear to G. gecko climbing or moving on a level surface. So, how that the low speeds achieved are not a matter of motivation but does E. macularius alter its locomotor behaviour when are a direct consequence of the lack of adhesive structures. climbing? As mentioned above, the narrow, slow speed range So, despite the fact that it has been shown in the literature obtained for climbing in the present study precludes statistical that substratum inclination can affect absolute sprint comparison. However, visual inspection of the frequency, performance (for lizards over 40 g; Huey and Hertz, 1984), the stride length and duty factor data (Figs 2, 3) suggests that, if net cost of transport (Farley and Emshwiller, 1996) and the level-surface locomotion had been recorded for E. macularius detailed limb kinematics and gait characteristics (Irschick and at such slow speeds, frequency, stride length and duty factor Jayne, 1998; Jayne and Irschick, 1999) we demonstrate here, would probably have been very similar to those measured for for two gecko species with clearly different preferred habitats, climbing. This suggestion is reinforced when the log/log that only slight adjustments in gait characteristics are made regressions for level locomotion presented in Table 3 are when they are forced to move on a non-habitual substratum. In extrapolated to the speeds obtained for climbing. The speed addition, the gait characteristics differ little between these two Gait characteristics of geckos 1245 species. Given the large niche differences and assuming Avery, R. A., Mueller, C. F., Smith, J. A. and Bond, D. J. (1987). the existence of selective pressure on spatio-temporal gait The movement patterns of lacertid lizards: speed, gait and pause in variables, this resemblance is probably dictated by historical Lacerta vivipara. J. Zool., Lond. 211, 47–63. (phylogenetic) constraints. If level locomotion represents the Cartmill, M. (1985). Climbing. In Functional Vertebrate Morphology ancestral state for gekkotans (Russell, 1976; Russell, 1979), it (ed. M. Hildebrand, D. M. Bramble, K. F. Liem and B. D. Wake), pp. 73–88. Cambridge, MA: Harvard University Press. is surprising to find that this common strategy suits climbing Diedrich, F. J. and Warren, W. H. (1995). Why change in gait? (fixed spatial variables, no floating phases) rather than level Dynamic of walk–run transition. J. Exp. Psychol. 21, 183–202. locomotion. Presumably, constraints other than those strictly Diedrich, F. J. and Warren, W. H. (1998a). The dynamics of gait coupled to locomotion must be considered to explain this. transition: effect of grade and load. J. Motor Behav. 30, 60–78. Ancestral gekkotans are thought to be nocturnal (Autumn et Diedrich, F. J. and Warren, W. H. (1998b). Dynamics of gait al., 1994; Autumn et al., 1997). The accuracy of foot placement transitions. In Timing of Behaviour. Neural, Psychological and (through fixed spatial gait variables) might be essential when Computational Perspectives (ed. D. A. Rosenbaum and C. E. visual inspection of the substratum is hindered by low- or no- Collyer), pp. 323–343. Cambridge, MA: MIT Press. light conditions. From this point of view, nocturnality could be Farley, C. T. and Emshwiller, M. (1996). Efficiency of uphill considered an exaptation for climbing. The development of locomotion in nocturnal and diurnal lizards. J. Exp. Biol. 199, adhesive pads allowed G. gecko, as well as other true geckos, 587–592. Fieler, C. L. and Jayne, B. C. (1998). Effect of speed on the hindlimb to exploit the climbing niche to its extremes (i.e. on smooth kinematics of the lizard Dipsosaurus dorsalis. J. Exp. Biol. 201, vertical or overhanging surfaces) without affecting their 609–622. performance on less-demanding substrata (sub-vertical, Full, R. J. and Kubow, T. (1998). The role of the mechanical system horizontal). This could explain why the level performance of in control. In Motion System (ed. R. Blickhan, A. Wisser and W. the specialist climber is not inferior to that of the ground- Nachtigall), pp. 215–216. Jena: Gustav Fischer. dwelling gekkotan (the basic control strategy is applied in all Grismer, L. L. (1988). Phylogeny, taxonomy, classification and circumstances), but also why the two species we studied biogeography in eubelpharid geckos. In Phylogenetic Relationships apparently move considerably slower than other (non- of the Lizard Families (ed. R. Estes and G. Pregill), pp. 369–469. climbing) lizards of similar size (see Fig. 7; i.e. the basic Stanford: Stanford University Press. control strategy is not optimized for running). This further Hildebrand, M. (1985). Walking and running. In Functional implies that the poor climbing performance of E. macularius Vertebrate Morphology (ed. M. Hildebrand, D. L. Bramble, K. F. Liem and D. B. Wake), pp. 38–57. Cambridge, MA: Harvard is probably a consequence of the extreme experimental University Press. conditions (vertical, smooth surface), rather than being an Hildebrand, M. (1988). Form and function in vertebrate feeding and expression of an intrinsic inability to climb. Detailed locomotion. Am. Zool. 28, 727–738. performance analyses, including tests on sub-vertical inclines, Holt, K. G., Hamill, J. and Anders, R. O. (1990). The force-driven could provide insight into this problem. Such analyses are the harmonic oscillator as a model for human locomotion. Human subject of our current research. Movement Sci. 9, 55–68. Holt, K. G., Hamill, J. and Anders, R. O. (1991). Predicting the Test specimens were provided by the Royal Zoological minimal energy costs of human walking. Med. Sci. Sport. Exerc. Society of Antwerp. The study is supported by GOA BOF 23, 491–498. 1999-2003 grant to P.A. and R.V.D. P.A. is a Research Huey, R. B. and Hertz, P. E. (1984). Effects of body size and slope on acceleration of a lizards (Stellio stellio). J. Exp. Biol. 110, Director and R.V.D and A.H. are post-doctoral researchers of 113–123. the FWO-Flanders. Irschick, D. J. and Jayne, B. C. (1998). Effect of incline on speed, acceleration, body posture and hindlimb kinematics in two species of lizard Callisaurus draconoides and Uma scoparia. J. Exp. Biol. References 201, 273–287. Abu-Ghalyun, Y. (1995). Histochemistry, capillarization and Irschick, D. J. and Jayne, B. C. (1999). Comparative three- mitochondrial densities of muscles fibres isolated from the dimensional kinematics of the hindimb for high-speed bipidal and iliobularis muscle of Agama pallida. Israel J. Zool. 41, 193–204. quadrapedal locomotion of lizards. J. Exp. Biol. 202, 1047–1065. Aerts, P., Van Damme, R., Van Elsacker, L. and Duchene, V. Jayne, B. C., Bennett, A. F. and Lauder, G. V. (1990). Muscle (2000). Spatio-temporal gait characteristics of the hind-limb cycles recruitment during terrestrial locomotion: how speed and during voluntary bipedal and quadrapedal walking in bonobos (Pan temperature affect fibre type use in a lizard. J. Exp. Biol. 152, paniscus). Am. J. Phys Anthropol. 111, 503–517. 101–128. Alexander, R. McN. (1992). Exploring Biomechanics: Animals in Jayne, B. C. and Irschick, D. J. (1999). Effect of incline and speed Motion. New York: Scientific American Library. on the three-dimensional hindlimb kinematics of a generalized Autumn, K., Farley, C. T., Emshwiller, M. and Full, R. J. (1997). iguanian lizard (Dipsosaurus dorsalis). J. Exp. Biol. 202, 143–159. Low cost of locomotion in the banded gecko: a test of nocturnal Kugler, P. N. and Turvey, M. T. (1987). Information, Natural Law hypothesis. Physiol. Zool. 70, 660–669. and the Self-Assembly of Rhythmic Movement: Theoretical and Autumn, K., Weinstein, R. B. and Full, R. J. (1994). Low cost of Experimental Investigations. Hillsdale, NJ: Erlbaum. locomotion increases performance at low temperature in a Latash, M. L. (1998). Neurophysiological Basis of Movement. nocturnal lizard. Physiol. Zool. 67, 238–262. Champaign, IL: Human Kinetics. 1246 A. ZAAF AND OTHERS Losos, J. B. (1990). The evolution of form and function: Morphology Russell, A. P. (1976). Some comment concerning interrelationships and locomotor performance in West indian Anolis lizards. among gekkonine geckos. In Morphology and Biology of Reptiles Evolution 44, 1189–1203. (ed. A. d'A. Bellairs and C. B. Cox), pp. 217–244. London: Losos, J. B., Walton, B. M. and Bennett, A. F. (1993). Trade-off Academic Press. between sprinting and clinging ability in Kenyan chameleons. Russell, A. P. (1979). Parallelism and integrated design in the foot Funct. Ecol. 7, 281–286. structure in the gekkonine and diplodactine geckos. Copeia 1, 1–21. McFadyen, B. J. and Bélanger M. (1997). Neuromechanical Schöner, G., Jiang, W. Y. and Kelso, J. A. S. (1990). A synergetic concepts for the assessment of control of human gait. In Three- theory of quadrupedal gaits and gait transition. J. Theor. Biol. 4, Dimensional Analysis of Human Locomotion (ed. P. Allard, A. 359–391. Cappozzo, A. Lundberg and C. L. Vaughan), pp. 49–66. Stewart, I. (1995). Nature's Numbers. Discovering Order and Chichester: John Wiley & Sons, Ltd. Pattern in the Universe. London: Weidenfeld & Nicolson. McMahon, T. A. (1984). Muscle, Reflexes and Locomotion. Stewart, I. and Golubitsky, M. (1992). Fearful Symmetry. Oxford: Princeton, NJ: Princeton University Press. Blackwell Publishers. Moermond, T. C. (1979). Habitat constraints on the behaviour, Thelen, E. and Smith, L. B. (1994). A Dynamic Systems Approach morphology and community structure of Anolis lizards. Ecology 60, to the Development of Cognition and Action. Cambridge, MA: MIT 152–164. Press. Norberg, U. (1994). Wing design, flight performance and habitat use Van Damme, R., Aerts, P. and Vanhooydonck, B. (1998). Variation in bats. In Ecological Morphology: Integrative Organismal Biology in morphology, gait characteristics and speed of locomotion in two (ed. P. C. Wainwright and S. M. Reilly), pp. 205–239. Chicago, populations of lizards. Biol. J. Linn. Soc. 63, 409–427. London: The University of Chicago Press. Verstappen, M. and Aerts, P. (2000). Terrestrial locomotion in the Peck, A. J. and Turvey, M. T. (1997). Coordination dynamics of black-billed magpie. I. Spatio-temporal gait characteristics. Motor bipedal galloping pattern. J. Motor Behav. 29, 311–325. Control 4, 150–164. Peterson, J. A. (1984). The locomotion of Chamaeleo (Reptilia: White, T. D. and Anderson, R. A. (1994). Locomotory patterns and Sauria) with particular reference to the forelimb. J. Zool., Lond. cost as related to body size and form in teiid lizards. J. Zool., Lond. 202, 1–42. 233, 107–128. Reilly, S. M. and Delancey, M. J. (1997). Sprawling locomotion in Zaaf, A., Herrel, A., Aerts, P. and De Vree, F. (1999). Morphology the lizard Sceloporus clarkii: the effect of speed on gait, hindlimb and morphometrics of the appendicular musculature in geckos with kinematics and axial bending during walking. J. Zool., Lond. 243, different locomotor habit (lepidosaurians). Zoomorph. 219, 11–22. 417–433. Zernicke, R. F. and Smith, L. J. (1996). Biomechanical insights to Russell, A. P. (1975). A contribution to the functional analysis of the neural control of movement. In Handbook of Physiology (ed. L. foot of the tokay Gekko gecko (Reptilia: Gekkonidae). J. Zool., Rowell and J. Shepard), pp. 293–330. New York: Oxford Lond. 176, 437–476. University Press.
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The RSPCA today
Since its humble beginning in 1824 - as the Society for the Prevention of Cruelty to Animals - the RSPCA has worked tirelessly to promote kindness and prevent cruelty to animals.
The RSPCA is a registered charity (no. 219099) that receives no lottery or state aid. Its £82 million annual running costs are funded exclusively by voluntary donations and legacies.
How YOU can help:
. If you know about an animal that is injured or being treated cruelly, call the RSPCA's national 24-hour cruelty and advice line on 0300 1234 999.
. Campaign for new animal welfare laws.
. Make a donation to the RSPCA to help the Society achieve even more for animals.
. Become a member of the RSPCA and support its work.
. Become a volunteer at your local RSPCA branch and help out with practical animal welfare work, or fundraising.
Action for animals
There are now 323 uniformed RSPCA inspectors and 146 Animal Collection Officers (ACOs) in England and Wales working tirelessly for animals in distress. In 2003 inspectors investigated 105,932 animal cruelty complaints which resulted in 1,829 convictions.
Preventing abuse
The courts are a last resort for inspectors, who prefer to educate rather than prosecute. They watch the treatment of animals in transit, in markets, pet shops, boarding kennels and farms and offer help and advice about their care.
Cruelty casebook
Every 25 seconds someone somewhere in England and Wales dials 0300 1234 999 - the RSPCA's national cruelty and advice line - for help. In 2003 the RSPCA removed 182,570 animals from danger or abuse. Inspectors and ACOs are also an emergency service for injured, trapped, or stranded animals and in 2003 they carried out 11,806 rescues.
Animal care
The RSPCA rehomed 69,787 animals in 2004 - mostly through the Society's network of 175 branches. Branches are separately registered charities operating subject to RSPCA and branch rules. They work for animal welfare locally and many provide services including subsidised veterinary treatment for those in need, neutering and rehoming schemes. Together, branches are responsible for 38 branch-run clinics, 37 animal centres and five animal welfare centres.
The RSPCA (national Society) is responsible for four animal hospitals, three specialist wildlife centres, 15 animal centres and seven clinics throughout England and Wales
All animals
The RSPCA's influence covers the whole range of animal protection. The Society is involved in practical welfare, law enforcement as well as high-profile campaigning and education. It employs veterinary experts and consultants in the care and treatment of farm livestock, wildlife, domestic pets and animals used in research.
Traditions maintained
The RSPCA's law-reforming origins are an important element of its work today. Much UK animal welfare legislation owes its existence to RSPCA campaigning, and the Society works for animal welfare at Westminster through its professional parliamentary lobbyists. Ministers and MPs frequently seek the advice and opinions of the RSPCA on legislative matters.
The RSPCA's reach extends into Europe through Eurogroup for Animal Welfare - a consortium of animal welfare organisations from each EU member state - that campaigns for improvements in animal welfare legislation.
Public eye
RSPCA advertising campaigns, television exposure, direct mail fundraising, promotional videos, magazines, booklets, leaflets and posters keep the Society at the forefront of public awareness about animal welfare.
Next generation
The RSPCA's education service also plays a key role. The education website has a range of National Curriculum-linked resources. Teachers can download lesson plans for primary and secondary pupils and order free photo packs online. Five regional education advisers lead a team of education officers to support teachers. To see if this service is available in your area, contact your local REA
International aid
The RSPCA's overseas fund distributed more than ?100,000 to animal welfare organisations in 18 different countries in 2003. There are more than 200 animal welfare organisations in 65 countries associated to the RSPCA.
Click here to visit RSPCA Online
What's new
New RSPCA Assured food label hits supermarket shelves
To help millions of farm animals.
RSPCA welcomes 'organ-on-a-chip' technology award of 'Design of the Year'
RSPCA warmly endorses new technology seeking to revolutionise the way medicines are developed and tested - without the use of animals.
The A to Z of RSPCA rescues from... axolotl to zebra finch
To mark the start of RSPCA Week, the charity releases a new list of the most unusual animals they rescued in 2014.
'Lack of ban on wild animals in circuses is outdated'
RSPCA brand the omission of ban of circus animals in Queen's Speech 'unacceptable'.
RSPCA response to news that the UK Government has NOT set out any plans to repeal the Hunting Act
David Bowles: 'The RSPCA is pleased to learn that the Government currently has no plans to repeal the Hunting Act and reintroduce such barbaric practices into the British countryside'.


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Your bathroom is one of the only rooms in your home that relaxes your cares away and can invigorate you to start anew the next the day. There is nothing better than bringing new innovations into your bathing and pampering experience and this year at the Kitchen and Bath Industry Show (KBIS) in Las Vegas, Nevada there was no lack of cool gadgets and gorgeous finds. Whether you are thinking about remodeling your bathroom or you just want to start creating your "wish list" here are 10 spectacular bathroom innovations for your modern home.
Image via: Kohler Faucets
1. Traditional Materials in Innovative Applications:
Bathrooms have always been a room where we can relax our cares away from the world and there is nothing like showcasing beautiful finishes in your bathroom. There is definitely a return to showcasing traditional materials such as wood, rock, seashells and other organic finishes in sinks such as Kohler Faucets above, countertops, backsplashes and more. While the materials are traditional they are being used for applications where you will see teak wood aesthetics in the gorgeous bathroom below in this article featuring Brizo Faucets.
Image via: Kohler Faucets
2. Emphasis on water conservation combined with technology
Over the years sustainability and water conservation have been on the forefront of all our lives, especially in the bathroom. At KBIS there was a reemergence of ensuring modern fixtures look sleek and fashionable but still are helping conserve water. Low flow toilets and the manufacturers focusing on greener ways to save water was seen in many toilet brands. One of the manufacturer's – Toto toilets has always been on the cutting edge of technology and their innovative process of using water to disinfect makes these automatic toilets the most impressive seat in your bathroom. This is one of the most impressive bathroom innovations of the entire show.
"The bath space consumes the most water in the home. In fact, toilets alone account for nearly 30 percent of indoor water consumption. Therefore, the toilet represents one of the greatest opportunities to decrease water consumption in the home. This can also be said of restrooms in many types of commercial buildings. With the right high efficiency fixtures and fittings, saving water becomes second nature." – Toto Toilets
3. New finishes appeal to international markets
Depending on your design style there is an appeal of a variety of finishes and colors. Gold and brass has become more apparent and is making a comeback in bathroom finishes. Everywhere from plumbing fixtures, to shower controls, lighting fixtures and even ceramic tile. Many middle eastern countries have an attraction to gold finishes while western countries prefer the matte chrome and oil bronzed finishes.
While there isn't one general finish for all markets its nice to see the bath industry starting to add more colors to their lines. While chrome and brushed nickel will never go out of style, copper, gold and rose gold finishes are making a great presence this year at the Kitchen and Bath Industry Show.
Image via: Brizo
4. Bathroom Innovations: Creative faucets to make your life easier
In this year's kitchen and bath show there was a definite sense of creative faucets that enhance your bathroom but also make life easier. From hand held faucets that grace the freestanding tub to deck mounted sinks that also offer the hand held sprayers. The sense of bringing kitchen faucet amenities to the bathroom is alive and well. Bathroom faucet manufacturers are starting to realize that the bathroom has the need for convenience and variety too.
5. Cabinetry handles and pulls in innovative materials
In years past cabinetry handles and pulls were strictly functional and came in only a few finishes and design styles. At KBIS there was an reemergence of understanding that pulls and handles are more than utilitarian. In fact the ability to showcase textures, patterns, colors, and even "jewels" has become a trend in modern bathrooms and kitchens.
Instead of thinking of knobs as an afterthought, pulls are being chosen when cabinetry fronts are designed. Top Knobs says they are the 'jewelry of the cabinetry' and rightfully so. There is no wonder that accessories such as handles and pulls are becoming more popular in today's bathrooms.
Image via: Top Knobs
6. Bathroom organization with creative amenities
Whether your bathroom is lacking good organization or you are seeking more, there was no shortage of creative amenities to help every home owner. Cabinetry that is divided into sections for personal styling aids, toiletries, cosmetics and more is becoming increasingly popular.
A number of bathroom cabinetry manufacturers are appealing to homeowners lifestyles and ensuring storage keeps up with the demands of busy households. Bathroom cabinets that close with a touch of a finger, have illuminated LED interiors and have divided interiors are a must for every modern home.
7. Metallic finishes in vanities and faucets that wow!
While metallic finishes have always been the main material for making plumbing fixtures, bathroom manufacturers are finding new ways to wow your bathroom interiors. From hammered sink details that resemble artisan crafts rather than a typical vanity sink to faucets that beautifully unveil the release of water similar to a gorgeous fountain, the options for your bathroom are limitless.
8. Vanity sinks that feel organic in nature
Porcelain and ceramic sinks have long been the materials of vanity sinks in our bathrooms but bathroom manufacturers are looking to organic materials that feel one with nature. From wood and sea shells to resin and concrete materials that can be made to look like materials right out of nature.
Teak, walnut, bamboo, and mahogany woods are big this year in natural and faux finishes. Also stained ceramics and tiles that are made to look like wood are becoming increasingly popular for sinks as well as backsplashes, walls and floors. They prove to be a trend to watch as they are water resistant and can often be cheaper than many hardwood varieties.
9. Shower controls are more sleek and modern than ever before
When rain shower heads were introduced years back some manufacturers only made one style and homeowners got tired of the same look. Today the relaxing shower head is still a must in any luxury bathroom and there was a gorgeous array at KBIS this year. From different sprayers and control options to low profile and luxurious details that will make your shower become a spa experience.
10. There is no limit to imagination of bath manufacturers
If there is one thing I learned by attending the Kitchen and Bath Industry show this year is that bathroom manufacturers are always trying to push the envelope to ensure your experience is the best. From showers and bathtubs that give you the ultimate bathing experience with smart technology, aromatherapy, chromo lights and sounds, there is very little you can't do while relaxing your cares away!
Also the technology to save the Earth's precious resource of water has become more of priority in all the brands. While beauty and style is important its nice to observe that bath manufacturers are using innovation to save water and reuse water in other parts of your home when applicable. There is no doubt that these 10 innovations will brighten up your bathroom this year!
Which of the innovations are you most excited about testing out in your bathroom?
3 comments
These seem like really great bathroom innovations. I really like the idea of having water conservation combined with technology installed in my bathroom. Does it take a special kind of plumber to install that kind of technology in a home? It would be really nice to have an automatic toilet installed in all of my bathrooms at home.
Thanks for the pictures! I'm feeling more inspired than ever to remodel my bathroom. I bought an older home, and I'm excited to update the interior. It's pretty boring to look at right now.
My mom has been wanting to remodel her bathroom for a while now. She has been looking at ideas lately and I think that I will show her some of these ideas because they are the same style that she is looking for. Do you have any tips or advice of things to do before starting the actual remodel? I can't wait for her to finally start and finish this project because she has wanted to do this for a really long time and I'm sure it is going to turn out great.
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Reviewed By Richard K (Alstead, NH) Reviewed for a 2011 GMC Sierra —6/24/2015 7:08 PM
All I will say is the whole experience was fun. All went great. The costumer service rep was Great.. The cover was a Fit.. Thanks Auto Anything..
Reviewed By Timothy S (GLOUCESTER, VA) Reviewed for a 2011 Ford Mustang —6/23/2015 1:48 PM
Haven't owned car cover through any severe weather yet. Car parked outside, excellent fit and quality. Applied cable along underside, but don't really believe its needed. Will definitely buy again.


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Adam Reach was sat in a classroom at Gateshead College five years ago. Now, he is one game away from the Premier League. He was plucked from further education by Middlesbrough aged 18.
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Middlesbrough boss Aitor Karanka and his captain Grant Leadbitter are dreaming of a trip to Wembley. The Smoggies currently lead Brentford 2-1 on aggregate in the play-off semi-final.
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On-loan Chelsea striker may stay if club are promoted as Middlesbrough go into Friday's Championship play-off second leg against Brentford with 2-1 lead
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St. Louis, MO (Sports Network) - Ryan Reaves scored two goals and the St. Louis Blues took a 4-1 win over the Calgary Flames on Thursday.
The win vaulted St. Louis from sixth to fourth place in the Western Conference standings. With 58 points, the team is one ahead of Los Angeles and San Jose, both of which were off on Thursday. All three teams have one game to play.
Andy McDonald and Alexander Steen each posted a goal while Brian Elliott stopped 18 shots for the Blues, who have won five of six.
"I don't care if its 4-1 or whatever," said St. Louis head coach Ken Hitchcock. "We had too many sloppy plays in the third period that could have ended up in our net. Our goalie made four great saves, and you can't make those mistakes either."
Sven Baertschi scored the lone goal of the game for the Flames, who have lost their past two. Joey MacDonald gave up all four goals on 20 shots in the loss.
"You can see how much it takes to be a playoff team," said Baertschi. "They had an outstanding season and you can tell there were some mistakes that we made that they took advantage of. They made some mistakes, but we couldn't capitalize."
After a scoreless first period, Reaves scored 3:21 into the second while Baertschi answered 4 1/2 minutes later with his third of the year to tie the game.
McDonald, though, scored on a snap shot from the slot off a feed from Chris Stewart for a 2-1 lead exactly halfway through the second period.
Skating on the power play, the Blues made it a 3-1 game as a left circle faceoff win saw the puck come down low and Steen backhanded his second attempt into the net.
Reaves made it a three-goal game 2:10 into the third as Chris Porter's wraparound attempt from the right side saw the rebound come out front where Reaves buried it for his fourth of the year.
Elliott stopped all eight shots he faced in the third to give St. Louis the big victory.
St. Louis closes its regular season at home against Chicago on Saturday ...
Calgary plays its final game of the season in Chicago on Friday ... St. Louis
took two of three from Calgary ... St. Louis was 1-for-5 on the power play
while Calgary was 0-for-4.
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The Houston Texans have signed defensive lineman Brandon Deaderick, who played last season with the New Orleans Saints.
The Toronto Blue Jays activated catcher Dioner Navarro from the 15-day disabled list following a 2-0 loss to the Washington Nationals in the opener of Tuesday's doubleheader.
Sacramento Kings forward Carl Landry underwent surgery on Tuesday to repair a torn ligament in his right wrist, the team said.
The Montreal Canadiens revealed on Tuesday that defenseman Jeff Petry has agreed to terms on a six-year contract extension.


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canvas prints, trout canvas prints, thom glace canvas prints, fish canvas prints, watercolor canvas prints, nature canvas prints
Corner Detail: Stretched canvas print with 1.5" stretcher bars and mirrored image sides. Also available with black sides, whites sides, and 5/8" stretcher bars.
Bring your artwork to life with the texture and depth of a stretched canvas print. Your image gets printed on one of our premium canvases and then stretched on a wooden frame of 1.5" x 1.5" stretcher bars (gallery wrap) or 5/8" x 5/8" stretcher bars (museum wrap). All stretched canvases ship within 3 - 4 business days and arrive "ready to hang" with pre-attached hanging wire, mounting hooks, and nails.
Stretched canvas prints look beautiful with or without frames.
Fine Art America is one of the largest, most-respected giclee printing companies in the world with over 40 years of experience producing museum-quality prints. All of our prints are produced on state-of-the-art, professional-grade Epson printers. We use acid-free papers and canvases with archival inks to guarantee that your prints last a lifetime without fading or loss of color.
'Study of a Greenback Cutthroat Trout' is part of a large series of North American Trout by Thom Glace. You can see more of his nature paintings at: www.thomglacewatercolors.com.
Thom Glace is a nature artist, specializing in transparent watercolor studies of fish(especially trout & salmon) and dragonfly species- with an occasional landscape thrown in for fun. Thom had to retire young due to a heart condition. He has been able to enjoy his love of painting since then. He had an interesting career in International Marketing that took he and his wife, Mary, to various locations around the world. After living in locations such as The Netherlands, St. Croix USVI, New Hampshire, North Carolina (Graduate of UNC- Chapel Hill) -he ended up back in Central Pennsylvania, where he was raised. So, after traveling to over 60 countries and 40 + states, Thom has been forced to end his wanderlust lifestyle. Now he lives...
Our return policy is very simple:
If you're not happy with a purchase that you made on FineArtAmerica.com, for any reason, you can return it to us within 30 days of the order date. As soon as it arrives, we'll issue a full refund for the entire purchase price. Please note - Fine Art America does not reimburse the outgoing or return shipping charges unless the return is due to a defect in quality.
Fine Art America sells thousands of pieces of artwork each month - all with a 100% money-back guarantee.
If you've hesitated to purchase artwork online in the past, why not give us a try? You have nothing to lose!
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Alumni filmmakers / Followers
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To all video makers. High quality footage material shop VMALL is here ! http://vfxmall.com VFX Twitter http://www.twitter.com/powsource Our Facebook http://www.facebook.com/profile.php?id=100001573008775 We at PYROTECH furnish the TV, Movie, Events, Stage, live Concerts with our specialized…
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Charles Breiner is a writer, director and editor currently based in Los Angeles. He has written and directed 7 short films including: "Wedding Night" "Art Sale" "The Landlord" "Oceans Apart and "Breaking In Is Hard To Do". He has also wrote, directed…
Calvin College PRO Grand Rapids, MI
Calvin College was established in 1876 in Grand Rapids, Michigan. From modest beginnings the college has grown to now include 4,000 students from around the world and more than 300 committed and caring Christian faculty members. Calvin offers 100 majors and programs on its 400-acre campus. Learn more…
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kids - Page 9
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If you want to get your kids outside, away from their Xbox or Wii, and interested in some of your backyard garden projects, you'll probably need to be somewhat inventive with your gardening ideas. Pulling weeds and planting flowerbeds just won't cut it for most of children. Think incentive here.
We have some ideas.
Mommy and Me Expo Coming Back To Amarillo For 2013
Should The U.S. Adopt Finland's 'Baby Box' For New Parents?
When it comes to maternity leave, The United States earns a pretty dismal ranking when compared to other countries, being the only first-world country, and one of only three countries (the others being Lesotho, Swaziland and Papua New Guinea) that does not offer paid maternity leaves. Not only do other countries have better systems in place for maternity leave, but they have some pretty great welcoming gifts for their newest citizens. In Finland, for example, new parents can choose between receiving the above maternity package filled with essentials for baby's first few months or a tax-free lump sum of 140 euros as a maternity grant.
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Last month, Michael Garcia made a lot of people smile. The 45-year-old, who is a waiter at Laurenzo's Restaurant in Houston, Texas, refused to serve a family who insulted Milo Castillo, a five-year-old with Down Syndrome, by suggesting that "special needs children need to be special someplace else."


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Avant Première Screening Schedule is online
69 companies from all over the world present 550 film productions all together: the exclusive preview to buyers, production - and distribution companies and broadcasters.
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If you participated at Avant Premère 2015 you can contact other participants through the Virtual Market!
Knowledge transfer at its best!
Young filmmakers get in touch with experts in these coaching and feedback sessions!
Technical innovations for the Arts: TIME Lab by Fraunhofer HHI
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Abu Dhabi's TAQA says not holding merger talks
Gold languishes below $1,170 ahead of U.S. jobs data
Iman bin Laden, the daughter of Al Qaeda head Osama bin Laden, has been given…
December 29, 2009 4:22 by kippreport
Iman bin Laden, the daughter of Al Qaeda head Osama bin Laden, has been given a "travel permit" from the Saudi Arabian embassy in Iran, reports Arabic-language paper Asharq Al-Awsat.
The paper reported last week that a number of Osama bin Laden's children sought refuge in the Saudi embassy in Tehran following the September 11 attacks on the US. It is believed they entered the country carrying false passports.
Iman bin Laden hopes to travel to her native Saudi Arabia on either Tuesday or Wednesday, the newspaper reports.
Dubai second most popular travel hub for Britons
December 29 2009
Why Gulf News? Why?
December 30 2009


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The Motion Photography Prize At Saatchi Gallery
Are animated GIFs art?
Are animated GIFs art?
Explore the detrimental effects of dizzying fame with humour and ...
Ants clamber the walls and bricks curl up into a ball.
Chilean architect Smiljan Radic offers something odd.
Find out how rockets work and what a space engineer actually does.
When did you last visit Olympia?
A second crop of new talent.
Lisa Dwan is terrifying and captivating.
How did a space ship to the moon end up in South Kensington?
Eight days of Christmas celluloid.
Representative art is making a comeback.
Brompton Road reportedly sold for £50 million.
Is this everything you'd ever want to know about one of the greatest ...
Breathtaking ancient artefacts next to their modern counterparts


text:
FIG. 1 is a perspective view of a slideable power adapter showing our new design, in a closed position;
FIG. 2 is a front elevation view thereof;
FIG. 3 is a back elevation view thereof;
FIG. 4 is a bottom plan view thereof;
FIG. 5 is a top plan view thereof;
FIG. 6 is a left side elevation view thereof; and,
FIG. 7 is a right side elevation view thereof.
The broken lines shown represent unclaimed subject matter and form no part of the claimed design.