Most textbooks and research reports state that the structures of the tetrapod forelimbs and hindlimbs are serial homologues. From this view, the main challenge of evolutionary biologists is not to explain the similarity between tetrapod limbs, but instead to explain why and how they have diverged. However, these statements seem to be related to a confusion between the serial homology of the vertebrate pelvic and pectoral appendages as a whole, and the serial homology of the specific soft- and hard-tissue structures of the tetrapod forelimbs and hindlimbs, leading to an even more crucial and puzzling question being overlooked: why are the skeletal and particularly the muscle structures of the forelimb and hindlimb actually so strikingly similar to each other? Herein we provide an updated discussion of these questions and test two main hypotheses: (i) that the similarity of the limb muscles is due to serial homology; and (ii) that tetrapods that use hindlimbs for a largely exclusive function (e.g. bipedalism in humans) exhibit fewer cases of similarity between forelimbs and hindlimbs than do quadrupedal species. Our review shows that of the 23 arm, forearm and hand muscles/muscle groups of salamanders, 18 (78%) have clear 'topological equivalents' in the hindlimb; in lizards, 14/24 (58%); in rats, 14/35 (40%); and in modern humans, 19/37 (51%). These numbers seem to support the idea that there is a plesiomorphic similarity and subsequent evolutionary divergence, but this tendency actually only applies to the three former quadrupedal taxa. Moreover, if one takes into account the total number of 'correspondences', one comes to a surprising and puzzling conclusion: in modern humans the number of forelimb muscles/muscle groups with clear 'equivalents' in the hindlimb (19) is substantially higher than in quadrupedal mammals such as rats (14), lizards (14) and even salamanders (18). These data contradict the hypothesis that divergent functions lead to divergent morphological structures. Furthermore, as we show that at least five of the 19 modern human adult forelimb elements that have a clear hindlimb 'equivalent' derive from embryonic anlages that are very different from the ones giving rise to their adult hindlimb 'equivalents', they also contradict the hypothesis that the similarity in muscle structures between the forelimb and hindlimb of tetrapods such as modern humans are due to their origin as serial homologues. This similarity is instead the result of phylogenetically independent evolutionary changes leading to a parallelism/convergence due to: (i) developmental constraints, i.e. similar molecular mechanisms are involved (particularly in the formation of the neomorphic hand), but this does not necessarily mean that similar anlages are used to form the similar adult structures; (ii) functional constraints, related to similar adaptations; (iii) topological constraints, i.e. limited physical possibilities; and even (iv) phylogenetic constraints, which tend to prevent/decrease the occurrence of new homoplasic similar...

Superfast muscles power high-frequency motions such as sound production and visual tracking. As a class, these muscles also generate low forces. Using the toadfish swimbladder muscle, the fastest known vertebrate muscle, we examined the crossbridge kinetic rates responsible for high contraction rates and how these might affect force generation. Swimbladder fibers have evolved a 10-fold faster crossbridge detachment rate than fast-twitch locomotory fibers, but surprisingly the crossbridge attachment rate has remained unchanged. These kinetics result in very few crossbridges being attached during contraction of superfast fibers (only Ϸ1͞6 of that in locomotory fibers) and thus low force. This imbalance between attachment and detachment rates is likely to be a general mechanism that imposes a tradeoff of force for speed in all superfast fibers.The superfast fiber type is found where high-frequency contractions are required, such as in vertebrate eye muscles and in both vertebrate and invertebrate synchronous soundproducing muscles. These muscles have a series of modifications for speed, including a large volume of sarcoplasmic reticulum (SR) (1-7) to produce very rapid calcium transients (8) and low-affinity troponin to speed myofilament deactivation after [Ca 2ϩ

Salamanders are generally agreed to represent the primitive tetrapod body plan, as well as a postural analog for early tetrapods. Dissection and description of the muscles of the forelimb, trunk, and hindlimb of the California newt, Taricha torosa, were undertaken to provide baseline data on the locomotor structures in this species. Hypaxial trunk muscles are similar to those of other vertebrates. As in other salamanders, limb muscles show a simple parallel-fibered architecture and often span multiple joints. Several differences in limb musculature were also noted. The extensor iliotibialis muscle possesses a single head in T. torosa, rather than the two heads common in larger salamander species. The ischioflexorius muscle, while divided into proximal and distal sections, is not distinct from the puboischiotibialis in its proximal portion. The femorofibularis is enlarged in this species; it is suggested that the femorofibularis and ischioflexorius muscles may be functionally analogous systems. Forelimb and hindlimb musculature show similar morphological patterns, particularly in distal limb segments, which may provide insight into the primitive arrangement of tetrapod limb muscles.

California newts (Taricha torosa) are capable of locomotion in both aquatic and terrestrial environments. The transition between swimming and terrestrial walking was examined by videotaping individual Taricha walking both up and down a ramp, inclined at 15°to the horizontal, that had its lower end immersed in water and its upper end out of the water. When ascending the ramp, California newts first approached it by swimming, then used their limbs to walk while still in water, and finally left the water using a normal terrestrial walking gait. The reverse of this sequence was observed when individuals descended the ramp. In both directions, Taricha used a lateral sequence walk with a duty factor of approximately 76% when out of the water. Timing of footfalls was more variable in water and featured shorter duty factors, leading to periods of suspension. Comparison of angular and timing variables revealed effects due to direction and degree of immersion. Few timing variables showed differences according to stride within sequence (indicating whether the animal was in or out of the water), suggesting that the basic walking pattern is equally good in both environments.

Moving on land versus in water imposes dramatically different requirements on the musculoskeletal system. Although many limbed vertebrates, such as salamanders and prehistoric tetrapodomorphs, have an axial system specialized for aquatic locomotion and an appendicular system adapted for terrestrial locomotion, diverse extant teleosts use the axial musculoskeletal system (body plus caudal fin) to move in these two physically disparate environments. In fact, teleost fishes living at the water's edge demonstrate diversity in natural history that is reflected in a variety of terrestrial behaviors: (1) species that have only incidental contact with land (such as largemouth bass, Micropterus) will repeatedly thrash, which can roll an individual downhill, but cannot produce effective overland movements, (2) species that have occasional contact with land (like Gambusia, the mosquitofish, which evade predators by stranding themselves) will produce directed terrestrial movement via a tail-flip jump, and (3) species that spend more than half of their lives on land (like the mudskipper, Periopthalmus) will produce a prone-jump, a behavior that allows the fish to anticipate where it will land at the end of the flight phase. Both tail-flip and prone jumps are characterized by a two-phase movement consisting of body flexion followed by extension-a movement pattern that is markedly similar to the aquatic fast-start. Convergence in kinematic pattern between effective terrestrial behaviors and aquatic fast starts suggests that jumps are an exaptation of a neuromuscular system that powers unsteady escape behaviors in the water. Despite such evidence that terrestrial behaviors evolved from an ancestral behavior that is ubiquitous among teleosts, some teleosts are unable to move effectively on land-possibly due to morphological trade-offs, wherein specialization for one environment comes at a cost to performance in the other. Indeed, upon emergence onto land, gravity places an increased mechanical load on the body, which may limit the maximum size of fish that can produce terrestrial locomotion via jumping. In addition, effective terrestrial locomotor performance may require a restructuring of the musculoskeletal system that directly conflicts with the low-drag, fusiform body shape that enhances steady swimming performance. Such biomechanical trade-offs may constrain which teleost species are able to make the evolutionary transition to life on land. Here, we synthesize the current knowledge of intermittent terrestrial locomotion in teleosts and demonstrate that extant fishes represent an important model system for elucidating fundamental evolutionary mechanisms and defining the physiological constraints that must be overcome to permit life in both the aquatic and terrestrial realms.

Salamanders are acknowledged to be the closest postural model of early tetrapods and are capable of walking both in a terrestrial environment and while submerged under water. Nonetheless, locomotion in this group is poorly understood, as is underwater pedestrian locomotion in general. We, therefore, quantified the movements of the body axis and limbs of the California newt, Taricha torosa, during steady-speed walking in two environments, both of which presented a level surface: a treadmill and a trackway that was submerged in an aquarium. For treadmill walking at a relative speed of 0.63 snout-vent lengths (SVL)/sec, newts used a diagonal couplets lateral sequence walk with a duty factor of 77%. In contrast, submerged speeds were nearly twice as fast, with a mean of 1.19 SVL/sec. The submerged gait pattern was closer to a trot, with a duty factor of only 41%, including periods of suspension. Environment appears to play a critical role in determining gait differences, with reduction of drag being one of the most important determinants in increasing duration of the swing phase. Quantitative analysis of limb kinematics showed that underwater strides were more variable than terrestrial ones, but overall were strikingly similar between the two environments, with joint movement reversals occurring at similar points in the step cycle. It is suggested that the fundamental walking pattern appears to function well under multiple conditions, with only minor changes in motor control necessary.

Based on similarity of motor patterns of lizards, crocodiles, birds and mammals, various authors have concluded that a number of homologous muscles across these taxa demonstrate neuromuscular conservatism. This hypothesis remains untested for more basal taxa. Therefore, a quantitative electromyographic study of the hind limb during treadmill walking (mean speed of 0.75 SVL/s) in the salamander Dicamptodon tenebrosus was undertaken. Muscles located ventrally on the hind limb become active just before foot placement on the substrate, and maintain activity through the first half of the stance phase. Dorsally located muscles begin activity at or just before the start of the swing phase, and fire through the first half of swing. Several muscles showed a secondary EMG burst during the stride. The second burst in most ventral muscles occurred in late stance. In all dorsal muscles with double bursts, the second burst occurred in the middle of stance. Comparison of electromyographic onset and offset values for Dicamptodon to those for presumed homologues in other tetrapods reveals similarity in activity patterns for all ventral and two dorsal muscles despite anatomical rearrangements, supporting the hypothesis of neuromuscular conservatism for some muscles but not others.Key words Salamander 9 Locomotion 9 Electromyography 9 Tetrapod 9 Neuromuscular conservatism Abbreviations BF biceps femoris muscle 9 CDF caudofemoralis muscle 9 CPIT caudalipuboischiotibialis muscle 9 Dist distal" EDC extensor digitorum communis muscle" EMG electromyogram 9 EXF extensor cruris et tarsi fibularis muscle 9EXT extensor cruris tibialis muscle 9 FMFB femorofibularis muscle 9 FPC flexor primordialis communis muscle 9 Gastroc gastrocnemius muscle 9 ILFB iliofibularis muscle 9 ILFM iliofemoralis muscle 9 ILTA extensor iliotibialis pars anterior muscle 9 ILTP extensor iliotibialis pars posterior muscle 9 ISC ischiocaudalis muscle 9 1SF ischioflexorius muscle 9 1SFM ischiofemoralis muscle 9 ITCR iliotrochantericus cranialis muscle 9 ITM iliotrochantericus medius muscle 9 MG medial gastrocnemius muscle 9 PFM pubifemoralis muscle 9 PIFE puboischiofemoralis externus muscle 9 PIF! puboischiofemoralis internus muscle 9 PIT puboischiotibialis muscle 9 Prox proximal 9 PTB pubotibialis muscle 9 Sol soleus muscle 9 ST semitendinosus muscle 9 SVL snout-vent length

SUMMARYMangrove rivulus (Kryptolebias marmoratus) are small fusiform teleosts (Cyprinodontiformes) with the ability to locomote on land, despite lacking apparent morphological adaptations for terrestrial movement. Rivulus will leave their aquatic habitat for moist, terrestrial environments when water conditions are poor, or, as we show here, to capture terrestrial insects. Specimens were conditioned to eat pinhead crickets on one side of their aquaria. After 2weeks of conditioning, a barrier with a slope of 15deg was partially submerged in the middle of the tank, forcing the fish to transition from water to land and back into water in order to feed. Kinematics during the transition were recorded using Fastec high-speed video cameras (125-250framess -1 ). Videos were analyzed using Didge and ImageJ software programs. Transition behaviors were characterized and analyzed according to their specific type. Body oscillation amplitude and wave duration were quantified for movements along the substrate, along with initial velocity for launching behaviors. Kryptolebias marmoratus used a diverse suite of behaviors to transition from water to land. These behaviors can be categorized as launches, squiggles and pounces. Prey were captured terrestrially and brought underwater for consumption. Kryptolebias marmoratus's suite of behaviors represents a novel solution to non-tetrapodal terrestrial transition, which suggests that fishes may have been able to exploit land habitats transiently, without leaving any apparent evidence in the fossil record. Supplementary material available online at