Hindlimb Morphology Research Articles

Scaling of the ankle extensor muscle-tendon units and the biomechanical implications for bipedal hopping locomotion in the post-pouch kangaroo Macropus fuliginosus.

Bipedal hopping is used by macropods, including rat-kangaroos, wallabies and kangaroos (superfamily Macropodoidea). Interspecific scaling of the ankle extensor muscle-tendon units in the lower hindlimbs of these hopping bipeds shows that peak tendon stress increases disproportionately with body size. Consequently, large kangaroos store and recover more strain energy in their tendons, making hopping more efficient, but their tendons are at greater risk of rupture. This is the first intraspecific scaling analysis on the functional morphology of the ankle extensor muscle-tendon units (gastrocnemius, plantaris and flexor digitorum longus) in one of the largest extant species of hopping mammal, the western grey kangaroo Macropus fuliginosus (5.8-70.5 kg post-pouch body mass). The effective mechanical advantage of the ankle extensors does not vary with post-pouch body mass, scaling with an exponent not significantly different from 0.0. Therefore, larger kangaroos balance rotational moments around the ankle by generating muscle forces proportional to weight-related gravitational forces. Maximum force is dependent upon the physiological cross-sectional area of the muscle, which we found scales geometrically with a mean exponent of only 0.67, rather than 1.0. Therefore, larger kangaroos are limited in their capacity to oppose large external forces around the ankle, potentially compromising fast or accelerative hopping. The strain energy return capacity of the ankle extensor tendons increases with a mean exponent of ~1.0, which is much shallower than the exponent derived from interspecific analyses of hopping mammals (~1.4-1.9). Tendon safety factor (ratio of rupture stress to estimated peak hopping stress) is lowest in the gastrocnemius (< 2), and it decreases with body mass with an exponent of -0.15, extrapolating to a predicted rupture at 160 kg. Extinct giant kangaroos weighing 250 kg could therefore not have engaged in fast hopping using 'scaled-up' lower hindlimb morphology of extant western grey kangaroos.

The oldest Asian hesperornithiform from the Upper Cretaceous of Japan, and the phylogenetic reassessment of Hesperornithiformes

Asian hesperornithiforms are extremely rare in contrast to the much more abundant record from North America. In Asia, these fossil birds are only known from fragmentary materials from Mongolia. Here we describe the skeletal remains of a new hesperornithiform Chupkaornis keraorum gen. et sp. nov. from the Upper Cretaceous Kashima Formation (Coniacian to Santonian) of the Yezo Group in Mikasa City, Hokkaido, Japan. This is the best-preserved hesperornithiform material from Asia and it is the first report of hesperornithiforms from the eastern margin of the Eurasian continent. Chupkaornis has a unique combination of characters: finger-like projected tibiofibular crest of femur, deep, emarginated lateral excavation with a sharply defined edge of the ventral margin of the thoracic vertebrae, and the heterocoelous articular surface of the thoracic vertebrae. Our new phylogenetic analysis revises the phylogenetic relationships of Hesperornithiformes. In contrast to previous studies, Enaliornis is assigned as the most basal taxon and Baptornis is positioned as more derived than Brodavis. Chupkaornis is a sister taxon to the clade of Brodavis and higher taxa. Parahesperornis and Hesperornis are positioned within Hesperornithidae, the derived Hesperornithiformes. Many of the skeletal character changes are concentrated at the base of Hesperornithidae (Parahesperornis and more derived taxa), and involve the modification of the pelvic girdle and hind limb morphology (e.g. dorsal directed antitrochanter of pelvis, short and sprawled femur, including probable lobe-toed feet suggested by the specialized distal articular surface of first digit of fourth toe, and predominantly robust digit IV phalanges). These skeletal modifications are likely adaptations for foot-propelled diving behaviour.http://zoobank.org/urn:lsid:zoobank.org:pub:FB783237-E565-4B74-9386-EADF8E12DFD4

Scaling and functional morphology in strigiform hind limbs

Strigiformes are an order of raptorial birds consisting exclusively of owls: the Tytonidae (barn owls) and the Strigidae (true owls), united by a suite of adaptations aiding a keen predatory lifestyle, including robust hind limb elements modified for grip strength. To assess variation in hind limb morphology, we analysed how the dimensions of the major hind limb elements in subfossil and modern species scaled with body mass. Comparing hind limb element length, midshaft width, and robusticity index (RI: ratio of midshaft width to maximum length) to body mass revealed that femoral and tibiotarsal width scale with isometry, whilst length scales with negative allometry, and close to elastic similarity in the tibiotarsus. In contrast, tarsometatarsus width shows strong positive allometry with body mass, whilst length shows strong negative allometry. Furthermore, the tarsometatarsi RI scales allometrically to mass0.028, whilst a weak relationship exists in femora (mass0.004) and tibiotarsi (mass0.004). Our results suggest that tarsometatarsi play a more substantial functional role than tibiotarsi and femora. Given the scaling relationship between tarsometatarsal width and robusticity to body mass, it may be possible to infer the body mass of prehistoric owls by analysing tarsometatarsi, an element that is frequently preserved in the fossil record of owls.

Comparative Morphometrics in Leptodactyline Frogs (Anura, Leptodactylidae, Leptodactylinae): Does Burrowing Behavior Relate to Sexual Dimorphism?

Abstract Fossorial habits occur in many animal lineages and usually involve both morphological and physiological adaptations that may evolve independently. Burrowing behavior in some species of the anuran subfamily Leptodactylinae involves the specialized use of the hind limbs and/or head. The aim of this study was to identify the morphometric characters associated with burrowing behavior in species of this subfamily. We then hypothesized that, as this habit is usually associated with males, we would find sexual dimorphism in head and hind-limb morphology in the burrowing species but not in the nonburrowing species. We compared 500 specimens from 24 species using phylogenetic statistical analyses and phylogenetic mapping of sexually dimorphic characters. The results demonstrated the following: 1) There was no correlation between the measurements of the limbs, head, or tarsal tubercle and burrowing behavior in the analyzed species; 2) there was no sexual dimorphism related to burrowing behavior reflected i...

Microgeographic variation in locomotor traits among lizards in a human-built environment.

Microgeographic variation in fitness-relevant traits may be more common than previously appreciated. The fitness of many vertebrates is directly related to their locomotor capacity, a whole-organism trait integrating behavior, morphology, and physiology. Because locomotion is inextricably related to context, I hypothesized that it might vary with habitat structure in a wide-ranging lizard, Podarcis erhardii, found in the Greek Cyclade Islands. I compared lizard populations living on human-built rock walls, a novel habitat with complex vertical structure, with nearby lizard populations that are naive to human-built infrastructure and live in flat, loose-substrate habitat. I tested for differences in morphology, behavior, and performance. Lizards from built sites were larger and had significantly (and relatively) longer forelimbs and hindlimbs. The differences in hindlimb morphology were especially pronounced for distal components—the foot and longest toe. These morphologies facilitated a significant behavioral shift in jumping propensity across a rocky experimental substrate. I found no difference in maximum velocity between these populations; however, females originating from wall sites potentially accelerated faster over the rocky experimental substrate. The variation between these closely neighboring populations suggests that the lizards inhabiting walls have experienced a suite of trait changes enabling them to take advantage of the novel habitat structure created by humans.

Hind limb morphometry of terror birds (Aves, Cariamiformes, Phorusrhacidae): functional implications for substrate preferences and locomotor lifestyle

ABSTRACTThe hind limbs of birds have long been considered a key feature in the conquest of different environments. However, the high level of morphological diversity encountered complicates the foundation of a good theoretical correlation between morphology, locomotor habits and substrate preference and this, in turn, complicates palaeobiological interpretations. Phorusrhacids (Aves, Cariamiformes) are a good example, since they have been unequivocally categorised as terrestrial birds due to their reduced forelimbs; and as apex predators with the ability to pursue prey based only on their hind limb morphology. Multivariate techniques (PCA and discriminant analysis), based on traditional metrics and geomorphometrics of the hind limb and pelvis, were applied in order to explore terrestriality and cursoriality in phorusrhacids. Although several groups of birds could be identified, when looking solely at hind limb metrics, some phorusrhacids appear to be associated with walking birds, while others are associated with cursorial birds. However, the pelvis separates cursorial birds and phorusrhacids from walking and wading birds. This scenario is complicated further by a lack of clear definition of the different locomotor modes and substrate preferences in extant birds, and this makes it difficult to confirm phorusrhacid cursoriality based solely on morphometrics. However, some qualitative features of the pelvis and foot make the picture a little clearer. To study limb adaptations in fossil birds, a more holistic study, with an emphasis on qualitative features of the whole posterior locomotor module, is necessary, since morphometrics leaves some issues unresolved. A comparison with the wings is also needed, in order to make a more complete analysis of locomotor behaviour.

Differences in hindlimb morphology of ducks and chickens: effects of domestication and selection.

BackgroundPoultry account for the most numerous species farmed for meat and have been subject to intense selection over approximately 60 generations. To assess morphological changes which have occurred in the avian leg due to selection for rapid growth and high meat yields, divergent lines of chicken (Gallus gallus) and duck (Anas platyrhynchos) were studied between 3 and 7 weeks of age. For each line, femoral and tibiotarsal morphology was recorded using computed tomography scanning and tibiotarsal bone quality measures (stiffness, bending stress and porosity) were assessed.ResultsIn chicken and duck, divergence in hindlimb morphology has occurred in the commercial meat lines compared to their lighter conspecifics. As expected, the differences were largest between species. Leg development nears completion much earlier in ducks than in chickens. Duck tibiotarsi showed a large degree of lateral curvature, which is expected to affect foot position during swimming and walking, and thus to influence gait. All lines have adapted their tibiotarsal morphology to suit the loading forces they experience; however bone quality was found to be poorer in chickens.ConclusionsWe demonstrate that intensive selection for growth rate in both chickens and ducks has resulted in leg morphology changes, which are likely to influence gait. Ducks represent an interesting compromise of adaptation for efficient locomotion in two media—on land and in water. Some aspects of bone morphology in the duck, such as lateral curvature of the tibiotarsus, may result from adaptation to swimming, which potentially imposes limitations on terrestrial locomotion.Electronic supplementary materialThe online version of this article (doi:10.1186/s12711-015-0166-9) contains supplementary material, which is available to authorized users.

Locomotor Mode and the Evolution of the Hindlimb in Western Mediterranean Anurans

The evolutionary association between morphology, locomotor performance and habitat use is a central element of the ecomorphological paradigm, and it is known to underlie the evolution of phenotypic diversity in numerous animal taxa. In anuran amphibians the hindlimb acts as the propulsive agent, and as such, it is directly associated with jumping performance. In this study we combine individual- and species-level analyses to examine the effects of locomotor mode on body size and hindlimb morphology of Western Mediterranean anurans. In addition to the commonly studied hindlimb traits, we also examine the ratio between tibiofibula and femur length. Body size shows no signs of an evolutionary association to locomotor mode. Instead, hindlimb traits are significantly differentiated between locomotor groups, both at the individual and species levels. Specifically, we observe a gradient of tibiofibula to femur ratio values that matches biomechanical predictions. The analysis of adult static allometries indicates that these differences arise early in ontogeny. By comparing the fit of distinct evolutionary models we provide evidence that the locomotor mode adopted by each species to match the requirements of the habitat it frequents has shaped the evolution of the hindlimb, but not body size.

Walking on five legs: investigating tail use during slow gait in kangaroos and wallabies

Pentapedal locomotion is the use of the tail as a fifth leg during the slow gait of kangaroos. Although previous studies have informally noted that some smaller species of macropodines do not engage in pentapedal locomotion, a systematic comparative analysis of tail use during slow gait across a wide range of species in this group has not been done. Analysis of relative movement of the pelvis, tail, and joint angles of the lower limbs during slow gait, using 2D landmark techniques on video recordings, was carried out on 16 species of Macropodinae. We also compared the relative lengthening of the tibia using crural index (CI) to test whether hindlimb morphology was associated with pentapedal locomotion. Pentapedal locomotion was characterised by three features: the presence of the ‘tail repositioning phase’, the constant height of the pelvis and the stationary placement of the distal tail on the ground during the hindlimb swing phase. The mean CI of pentapedal species was significantly greater than that of non-pentapedal species (1.71 versus 1.36; P &lt; 0.001). This lends support to the hypothesis that the use of pentapedal locomotion is associated with the relative lengthening of the hindlimb, which in turn is associated with body size and habitat preference within the Macropodinae.

Comparative functional anatomy of hindlimb muscles and bones with reference to aquatic adaptation of the sea otter.

Although the sea otter (Enhydra lutris) is a complete aquatic species, spending its entire life in the ocean, it has been considered morphologically to be a semi-aquatic animal. This study aimed to clarify the unique hindlimb morphology and functional adaptations of E. lutris in comparison to other Mustelidae species. We compared muscle mass and bone measurements of five Mustelidae species: the sea otter, Eurasian river otter (Lutra lutra), American mink (Neovison vison), Japanese weasel (Mustela itatsi) and Siberian weasel (M. sibirica). In comparison with the other 4 species, E. lutris possessed significantly larger gluteus, popliteus and peroneus muscles, but smaller adductor and ischiopubic muscles. The popliteus muscle may act as a medial rotator of the crus, and the peroneus muscle may act as an abductor of the fifth toe and/or the pronator of the foot. The bundles of the gluteus superficialis muscle of E. lutris were fused with those of the tensor fasciae latae muscle and gluteofemoralis muscles, and they may play a role in femur abduction. These results suggest that E. lutris uses the abducted femur, medially rotated crus, eversion of the ankle and abducted fifth digit or extended interdigital web as a powerful propulsion generator. Therefore, we conclude that E. lutris is a complete aquatic animal, possessing differences in the proportions of the hindlimb muscles compared with those in other semi-aquatic and terrestrial mustelids.

Trunk orientation causes asymmetries in leg function in small bird terrestrial locomotion.

In contrast to the upright trunk in humans, trunk orientation in most birds is almost horizontal (pronograde). It is conceivable that the orientation of the heavy trunk strongly influences the dynamics of bipedal terrestrial locomotion. Here, we analyse for the first time the effects of a pronograde trunk orientation on leg function and stability during bipedal locomotion. For this, we first inferred the leg function and trunk control strategy applied by a generalized small bird during terrestrial locomotion by analysing synchronously recorded kinematic (three-dimensional X-ray videography) and kinetic (three-dimensional force measurement) quail locomotion data. Then, by simulating quail gaits using a simplistic bioinspired numerical model which made use of parameters obtained in in vivo experiments with real quail, we show that the observed asymmetric leg function (left-skewed ground reaction force and longer leg at touchdown than at lift-off) is necessary for pronograde steady-state locomotion. In addition, steady-state locomotion becomes stable for specific morphological parameters. For quail-like parameters, the most common stable solution is grounded running, a gait preferred by quail and most of the other small birds. We hypothesize that stability of bipedal locomotion is a functional demand that, depending on trunk orientation and centre of mass location, constrains basic hind limb morphology and function, such as leg length, leg stiffness and leg damping.

Morphoecological Peculiarities of Pelvis in Several Genera of Rails with Some notes On systematic Position Of The Coot, Fulica Atra (Rallidae, Gruiformes)

Abstract Morphoecological Peculiarities of Pelvis in Several Genera of Rails with Some Notes on Systematic Position of the Coot, Fulica atra (Rallidae, Gruiformes). Bogdanovich, I. A. - Some features of pelvic and hindlimb morphology of several ecologically different species of rallids are studied. Adaptive significances of these features are revealed. A special attention is paid to the hindlimb skeleton of the Coot (Fulica atra, Linnaeus, 1758), which retained the locomotor universalism despite adaptations to swimming and diving.

The phylogenetic position of the Pterosauria within the Archosauromorpha re-examined

A previous analysis of the phylogenetic position of the Pterosauria argued that pterosaurs were not closely related to dinosaurs as is generally accepted, but rather were outside the crown group Archosauria. However, that study was dismissed for the use of inappropriate methods. Here, the data set from that analysis was divided into five partitions: one with characters associated with cursorial digitigrade bipedal locomotion and the other four with characters from the skull and mandible, postcranial axial skeleton, forelimb and hindlimb, respectively. The partitions were subjected to homogeneity testing, and the Cursorial partition was found to be incongruent with other partitions and all other characters at the α = 0.01 probability level. Deletion of the Pterosauria removed all significant incongruence, demonstrating that the incongruence results from the coding of pterosaurs for the cursorial characters. The cause of the incongruence was interpreted as homoplasy in hindlimb morphology, and after re-evaluating and reformulating the characters of the Cursorial partition, the revised data set was tested for homogeneity and no significant incongruence was found. Lastly, the data set was updated with additional characters and taxa from recent analyses, tested as before, and when analysed suggested that the Pterosauria were basal archosauriforms well outside the crown group Archosauria.

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

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

The ontogeny of escape behavior, locomotor performance, and the hind limb in Sceloporus woodi

Flight initiation distance describes the distance at which an animal flees during the approach of a predator. This distance presumably reflects the tradeoff between the benefits of fleeing versus the benefits of remaining stationary. Throughout ontogeny, the costs and benefits of flight may change substantially due to growth-related changes in sprint speed; thus ontogenetic variation in flight initiation distance may be substantial. If escape velocity is essential for surviving predator encounters, then juveniles should either tolerate short flight initiation distances and rely on crypsis, or should have high flight initiation distances to remain far away from their predators. We examined this hypothesis in a small, short-lived lizard (Sceloporus woodi). Flight initiation distance and escape velocity were recorded on an ontogenetic series of lizards in the field. Maximal running velocity was also quantified in a laboratory raceway to establish if escape velocities in the field compared with maximal velocities as measured in the lab. Finally a subset of individuals was used to quantify how muscle and limb size scale with body size throughout ontogeny. Flight initiation distance increased with body size; larger animals had higher flight initiation distances. Small lizards had short flight initiation distances and remained immobile longer, thus relying on crypsis for concealment. Escape velocity in the field did not vary with body size, yet maximum velocity in the lab did increase with size. Hind limb morphology scaled isometrically with body size. Isometric scaling of the hind limb elements and its musculature, coupled with similarities in sprint and escape velocity across ontogeny, demonstrate that smaller S. woodi must rely on crypsis to avoid predator encounters, whereas adults alter their behavior via larger flight initiation distance and lower (presumably less expensive) escape velocities.

Morphological integration versus ecological plasticity in the avian pelvic limb skeleton

Understanding patterns and distributions of morphological traits is essential for discerning underpinning processes of morphological variation. We report on the variation in the avian pelvic limb skeleton. Length and width variables were measured in the skeletons of 236 avian species in order to examine the importance of body mass, ecological factors, phylogeny and integration in the formation of specific hindlimb morphology. Scaling relationships with body mass were analyzed across Aves and in individual avian subclades. Principal component analysis and multiple regressions were performed to examine the relationship between morphology, ecology, and phylogeny. Finally, the occurrence of within-limb morphological integration was tested by partial correlation analysis of the residuals from element lengths vs. body mass and correlation analysis of avian hindlimb proportions. Body mass is the greatest contributor to variation, and it strongly influences variation in avian skeletal lengths. Lengthening of the leg typically comes from disproportionate increases in tibiotarsal and tarsometatarsal length. Partial correlation analysis showed that only these two elements are distinctly integrated consistently across all bird taxa, whereas relation of femur and third toe to other limb elements displays no clear pattern. Hence, morphological integration of all elements is not a prerequisite for limb design, and variation between taxa is mainly to be found in femoral and digital length. Furthermore, variation in tibiotarsal relative length is much lower than in other elements likely due to geometric constrains. Clear ecological adaptations are obscured by multifunctionality of the avian hindlimb, and phylogeny significantly constrains the morphology. Finally, when looking at relative lengths segmented limbs meet the requirements of many-to-one-mapping of phenotype to functional property, in line with a common concept of evolvability of function and morphology.

Pitx1 broadly associates with limb enhancers and is enriched on hindlimb cis-regulatory elements

Extensive functional analyses have demonstrated that the pituitary homeodomain transcription factor Pitx1 plays a critical role in specifying hindlimb morphology in vertebrates. However, much less is known regarding the target genes and cis-regulatory elements through which Pitx1 acts. Earlier studies suggested that the hindlimb transcription factors Tbx4, HoxC10, and HoxC11 might be transcriptional targets of Pitx1, but definitive evidence for direct regulatory interactions has been lacking. Using ChIP-Seq on embryonic mouse hindlimbs, we have pinpointed the genome-wide location of Pitx1 binding sites during mouse hindlimb development and identified potential gene targets for Pitx1. We determined that Pitx1 binding is significantly enriched near genes involved in limb morphogenesis, including Tbx4, HoxC10, and HoxC11. Notably, Pitx1 is bound to the previously identified HLEA and HLEB hindlimb enhancers of the Tbx4 gene and to a newly identified Tbx2 hindlimb enhancer. Moreover, Pitx1 binding is significantly enriched on hindlimb relative to forelimb-specific cis-regulatory features that are differentially marked by H3K27ac. However, our analysis revealed that Pitx1 also strongly associates with many functionally verified limb enhancers that exhibit similar levels of activity in the embryonic mesenchyme of forelimbs and hindlimbs. We speculate that Pitx1 influences hindlimb morphology both through the activation of hindlimb-specific enhancers as well as through the hindlimb-specific modulation of enhancers that are active in both sets of limbs.

Pelvic and hindlimb myology of the basal archosaur Poposaurus gracilis (archosauria: Poposauroidea)

The discovery of a largely complete and well preserved specimen of Poposaurus gracilis has provided the opportunity to generate the first phylogenetically based reconstruction of pelvic and hindlimb musculature of an extinct nondinosaurian archosaur. As in dinosaurs, multiple lineages of basal archosaurs convergently evolved parasagittally erect limbs. However, in contrast to the laterally projecting acetabulum, or "buttress erect" hip morphology of ornithodirans, basal archosaurs evolved a very different, ventrally projecting acetabulum, or "pillar erect" hip. Reconstruction of the pelvic and hindlimb musculotendinous system in a bipedal suchian archosaur clarifies how the anatomical transformations associated with the evolution of bipedalism in basal archosaurs differed from that of bipedal dinosaurs and birds. This reconstruction is based on the direct examination of the osteology and myology of phylogenetically relevant extant taxa in conjunction with osteological correlates from the skeleton of P. gracilis. This data set includes a series of inferences (presence/absence of a structure, number of components, and origin/insertion sites) regarding 26 individual muscles or muscle groups, three pelvic ligaments, and two connective tissue structures in the pelvis, hindlimb, and pes of P. gracilis. These data provide a foundation for subsequent examination of variation in myological orientation and function based on pelvic and hindlimb morphology, across the basal archosaur lineage leading to extant crocodilians.

Divergence in locomotor performance, ecology, and morphology between two sympatric sister species of desert-dwelling gecko

The functional and evolutionary implications of morphological diversification have been addressed for several groups of vertebrates. Although the mechanisms of gecko adhesion have received considerable attention, little is known regarding locomotor performance of geckos in nature, and how this might relate to morphological diversity. In this study we examine locomotor performance of two sympatric sister species of geckos of the genus Rhoptropus (Rhoptropus afer and Rhoptropus bradfieldi) found in the coastal desert regions of Namibia. One species (R. afer) commonly runs on sandy substrates and moves between isolated rock sheets, whereas the other species (R. bradfieldi) commonly lives and runs on isolated boulders. The morphology of R. afer is extremely divergent from its sister species and all other species in the genus. We initially recorded the inclination of the substrate in which the lizards were found in order to characterize the habitat of each species. We then quantified maximum speed and acceleration on a level 1-m trackway, and also during escapes in the field. We found that R. bradfieldi occupies steeper surfaces than those occupied by R. afer. On the trackway and in nature, R. afer runs faster than R. bradfieldi, although the differences in locomotor performance between the species are greater for the field measurements. Rhoptropus afer commonly runs for more than 2 or 3 m, whereas R. bradfieldi commonly runs less than 50 cm during an escape sprint. Our main conclusions are that: (1) R. afer attains higher maximum speeds when escaping under controlled and field conditions, although the magnitude differs between conditions; and (2) hindlimb morphology correlates with maximum running speed in R. afer, but not in R. bradfieldi. Similar to the well-studied Anolis lizards, we propose that these two gecko species represent distinct and highly divergent ecomorphs. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 860‐869.

Mutation in the Bmpr1b gene affects optic nerve development

Abstract Purpose Systematic analysis of the ocular defects in a new mouse mutant carrying a Bmpr1b mutation. Methods The Bmpr1b mutants were analyzed by slit lamp, funduscopy, laser interference biometry, electroretinography, virtual drum, histology and in‐situ hybridization. Results The new Bmpr1b mutant is characterized by a T‐&gt;G transversion in the splice donor site of exon 10. The mutants show irregular forelimb and hindlimb morphology, brachydactyly, irregular cartilage development and alterations in the reproductive system. The lens is clear, but in homozygous mutants, an enlarged optic disc was observed by funduscopy. The eyes are slightly larger, which is mainly due to an increased aqueous humor. Electroretinography demonstrated a more rapid decline of the b‐wave indicating a less efficient transmission of electric stimuli within the retina. Moreover, homozygous mutant mice show also almost no response to moving stripes as compared to the wild type mice. Histological analysis indicated dramatic changes in the optic nerve head. Analysis of the embryonic development of the eye showed that the expression area of Pax6 expands posterior, and concomitantly, Pax2 expression (which is regular at embryonic day 11.5) is repressed at embryonic days 15.5 and 17.5 at the optic nerve head. Conclusion The T‐G mutation in the splice donor site of exon 10 of the Bmpr1b gene leads to a recessive hypoplasia of the optic nerve head, which is mainly caused by repression of Pax2 expression in the late phase of embryogenesis.