SUMMARYMudskippers use pectoral fins for their primary mode of locomotion on land and pectoral fins in conjunction with the axial musculature and caudal fin to move in water. We hypothesized that distinct pectoral fin movements enable effective locomotion in each environment. Additionally, we made three functional predictions about fin movements during locomotion on land versus water: the pectoral fin is depressed more on land than in water; the pectoral fin will have greater changes in fin area between propulsive and recovery phases in water versus land; anterior and posterior excursions will be greater on land than in water. Locomotion was recorded in each environment using a high-speed digital-imaging system and kinematic variables were calculated from digitized landmark points. Variables were analyzed using principal components analysis and matched pairs ttests. Mudskippers produce distinct kinematic patterns across environments (P<0.003), although only some of our predictions were supported. The magnitude of fin depression is the same across habitats. However, depression occurs during the propulsive phase on land (by -0.60 cm), whereas during the propulsive phase in water the fin is elevated (by +0.13 cm). We were unable to support the hypothesis that fin orientation differs between environments. Lastly, anterior extension of the fin is greater on land (1.8 cm, versus 1.3 cm in water), creating a larger stride length in this environment. We posit that the mudskipper pectoral fin may facilitate stability in water and thrust production on land, and suggest that the robust fin morphology of the goby lineage may predispose species within this group to terrestrial locomotion.

SUMMARYMany amphibious organisms undergo repeated aquatic to terrestrial transitions during their lifetime; limbless, elongate organisms that make such transitions must rely on axial-based locomotion in both habitats. How is the same anatomical structure employed to produce an effective behavior across such disparate habitats? Here, we examine an elongate amphibious fish, the ropefish (Erpetoichthys calabaricus), and ask: (1) how do locomotor movements change during the transition between aquatic and terrestrial environments and (2) do distantly related amphibious fishes demonstrate similar modes of terrestrial locomotion? Ropefish were examined moving in four experimental treatments (in which the water level was to lowered mimic the transition between environments) that varied from fully aquatic to fully terrestrial. Kinematic parameters (lateral excursion, wavelength, amplitude and frequency) were calculated for points along the midline of the body and compared across treatments. Terrestrial locomotion in the ropefish is characterized by long, slow, large-amplitude undulations down the length of the body; in contrast, aquatic locomotion is characterized by short-wavelength, small-amplitude, high-frequency undulations that gradually increase in an anterior to posterior direction. Experimental treatments with intermediate water levels were more similar to aquatic locomotion in that they demonstrated an anterior to posterior pattern of increasing lateral excursion and wave amplitude, but were more similar to terrestrial locomotion with regard to wavelength, which did not change in an anterior to posterior direction. Finally, the ropefish and another elongate amphibious fish, the eel, consistently exhibit movements characterized by 'path following' when moving on land, which suggests that elongate fishes exhibit functional convergence during terrestrial locomotion. Supplementary material available online at

While emergent behaviours have long been reported for air-breathing osteichthyians, only recently have researchers undertaken quantitative analyses of terrestrial locomotion. This review summarizes studies of sustained periodic terrestrial movements by air-breathing fishes and quantifies the contributions of the paired appendages and the axial body to forward propulsion. Elongate fishes with axial-based locomotion, e.g. the ropefish Erpetoichthys calabaricus, generate an anterior-to-posterior wave of undulation that travels down the axial musculoskeletal system and pushes the body against the substratum at multiple points. In contrast, appendage-based locomotors, e.g. the barred mudskipper Periophthalmus argentilineatus, produce no axial bending during sustained locomotion, but instead use repeated protraction-retraction cycles of the pectoral fins to elevate the centre of mass and propel the entire body anteriorly. Fishes that use an axial-appendage-based mechanism, e.g. walking catfishes Clarias spp., produce side-to-side, whole-body bending in co-ordination with protraction-retraction cycles of the pectoral fins. Once the body is maximally bent to one side, the tail is pressed against the substratum and drawn back through the mid-sagittal plane, which elevates the centre of mass and rotates it about a fulcrum formed by the pectoral fin and the ground. Although appendage-based terrestrial locomotion appears to be rare in osteichthyians, many different species appear to have converged upon functionally similar axial-based and axial-appendage-based movements. Based on common forms observed across divergent taxa, it appears that dorsoventral compression of the body, elongation of the axial skeleton or the presence of robust pectoral fins can facilitate effective terrestrial movement by air-breathing fishes.

Flow diversion and invasive species are two major threats to freshwater ecosystems, threats that restoration efforts attempt to redress. Yet, few restoration projects monitor whether removal of these threats improve target characteristics of the ecosystem. Fewer still have an appropriate experimental design from which causal inferences can be drawn as to the relative merits of removing exotic fish, restoring flow, or both. We used a dam decommissioning in Fossil Creek, Arizona, to compare responses of native fish to exotic fish removal and flow restoration, using a beforeafter-control-impact design with three impact treatments: flow restoration alone where exotics had not been present, flow restoration and exotic fish removal, and flow restoration where exotics remain and a control reach that was unaffected by restoration actions. We show that removal of exotic fish dramatically increased native fish abundance.Flow restoration also increased native fish abundance, but the effect was smaller than that from removing exotics. Flow restoration had no effect where exotic fish remained, although it may have had other benefits to the ecosystem. The cost to restore flow ($12 million) was considerably higher than that to eradicate exotics ($1.1 million). The long-term influence of flow restoration could increase, as travertine dams grow and re-shape the creek increasing habitat for native fish. But in the 2-year period considered here, the return on investment for extirpating exotics far exceeded that from flow restoration. Projects aimed to restore native fish by restoring flow should also consider the additional investment required to eradicate exotic fish.

Titin has long been known to contribute to muscle passive tension. Recently, it was also demonstrated that titin-based stiffness increases upon Ca 2+ activation of wild-type mouse psoas myofibrils stretched beyond overlap of the thick and thin filaments. In addition, this increase in titin-based stiffness was impaired in single psoas myofibrils from mdm mice, characterized by a deletion in the N2A region of the Ttn gene. Here, we investigated the effects of activation on elastic properties of intact soleus muscles from wild-type and mdm mice to determine whether titin contributes to active muscle stiffness. Using load-clamp experiments, we compared the stress-strain relationships of elastic elements in active and passive muscles during unloading, and quantified the change in stiffness upon activation. Results from wild-type muscles show that upon activation, the elastic modulus increases, elastic elements develop force at 15% shorter lengths, and there was a 2.9-fold increase in the slope of the stress-strain relationship. These results are qualitatively and quantitatively similar to results from single wild-type psoas myofibrils. In contrast, mdm soleus showed no effect of activation on the slope or intercept of the stress-strain relationship, which is consistent with impaired titin activation observed in single mdm psoas myofibrils. Therefore, it is likely that titin plays a role in the increase of active muscle stiffness during rapid unloading. These results are consistent with the idea that, in addition to the thin filaments, titin is activated upon Ca 2+ influx in skeletal muscle.

Many teleosts that live at the water's edge will voluntarily strand themselves to evade predators or escape poor conditions-this behavior has been repeatedly observed in the field for killifishes (Cyprinodontiformes). Although most killifishes are considered fully aquatic and possess no obvious morphological specializations to facilitate terrestrial locomotion, individuals from several different species have been observed moving across land via a "tail flip" behavior that generates a terrestrial jump. Like aquatic fast starts, terrestrial jumps are produced by high-curvature lateral flexion of the body (stage one), followed by contralateral flexion of the posterior body (stage two). Here, terrestrial jumps and aquatic fast starts are quantified for two littoral teleosts: Gambusia affinis (a killifish, Cyprinodontiformes) and Danio rerio (a small carp, Cypriniformes) to determine if the tail flip is produced by other (non-killifish) teleosts and to test the null hypothesis that the tail flip is a fast start behavior, performed on land. Both Danio and Gambusia produce tail flip-driven terrestrial jumps, which are kinematically distinct from aquatic escapes and characterized by (1) a prolonged stage one, during which the fish bends, lifting and rolling the center of mass over the caudal peduncle, and (2) a relatively brief stage two, wherein the caudal peduncle pushes against the substrate to launch the fish into the aerial phase. The ability of these fully aquatic fishes to employ the same structure to produce distinct kinematic patterns in disparate environments suggests that a new behavior has evolved to facilitate movement on land and that anatomical novelty is not a prerequisite for effective terrestrial locomotion.

Shivering frequency scales predictably with body mass and is 10 times higher in a mouse than a moose. The link between shivering frequency and body mass may lie in the tuning of muscle elastic properties. Titin functions as a muscle 'spring', so shivering frequency may be linked to titin's structure. The muscular dystrophy with myositis (mdm) mouse is characterized by a deletion in titin's N2A region. Mice that are homozygous for the mdm mutation have a lower body mass, stiffer gait and reduced lifespan compared with their wild-type and heterozygous siblings. We characterized thermoregulation in these mice by measuring metabolic rate and tremor frequency during shivering. Mutants were heterothermic at ambient temperatures of 20-37°C while wild-type and heterozygous mice were homeothermic. Metabolic rate increased at smaller temperature differentials (i.e. the difference between body and ambient temperatures) in mutants than in nonmutants. The difference between observed tremor frequencies and shivering frequencies predicted by body mass was significantly larger for mutant mice than for wild-type or heterozygous mice, even after accounting for differences in body temperature. Together, the heterothermy in mutants, the increase in metabolic rate at low temperature differentials and the decreased tremor frequency demonstrate the thermoregulatory challenges faced by mice with the mdm mutation. Oscillatory frequency is proportional to the square root of stiffness, and we observed that mutants had lower active muscle stiffness in vitro. The lower tremor frequencies in mutants are consistent with reduced active muscle stiffness and suggest that titin affects the tuning of shivering frequency.

Aquatic vertebrates that emerge onto land to spawn, feed, or evade aquatic predators must return to the water to avoid dehydration or asphyxiation. How do such aquatic organisms determine their location on land? Do particular behaviors facilitate a safe return to the aquatic realm? In this study, we asked: will fully-aquatic mosquitofish (Gambusia affinis) stranded on a slope modulate locomotor behavior according to body position to facilitate movement back into the water? To address this question, mosquitofish (n = 53) were placed in four positions relative to an artificial slope (30° inclination) and their responses to stranding were recorded, categorized, and quantified. We found that mosquitofish may remain immobile for up to three minutes after being stranded and then initiate either a “roll” or a “leap”. During a roll, mass is destabilized to trigger a downslope tumble; during a leap, the fish jumps up, above the substrate. When mosquitofish are oriented with the long axis of the body at 90° to the slope, they almost always (97%) initiate a roll. A roll is an energetically inexpensive way to move back into the water from a cross-slope body orientation because potential energy is converted back into kinetic energy. When placed with their heads toward the apex of the slope, most mosquitofish (>50%) produce a tail-flip jump to leap into ballistic flight. Because a tail-flip generates a caudually-oriented flight trajectory, this locomotor movement will effectively propel a fish downhill when the head is oriented up-slope. However, because the mass of the body is elevated against gravity, leaps require more mechanical work than rolls. We suggest that mosquitofish use the otolith-vestibular system to sense body position and generate a behavior that is “matched” to their orientation on a slope, thereby increasing the probability of a safe return to the water, relative to the energy expended.