Body mass and the energy efficiency of locomotion: lessons from incline running

Rates of oxygen consumption were measured for two bipedal runners (two species of quail) and two quadrupedal runners (two small species of rodents), with average body masses that ranged from 0.035 to 0.217 kg, trained to run on a treadmill set to horizontal and then to a 10 degrees incline. Rates of oxygen consumption increased linearly with speed for all four species and the rates of increase were significantly higher (P < or = 0.05) for all four species when the animals were run on an incline than when they were run on a horizontal. The estimated metabolic energy cost to lift 1 kg mass 1 m vertically was similar for bipeds and quadrupeds of similar body mass and inversely related to body mass for both running styles. When the data for the animals used in the present study are combined with similar data for adult animals from the literature, the results show that the metabolic energy efficiencies of locomotion, estimated from the cost of vertical work, are the same for bipedal and quadrupedal runners. In both groups, the metabolic energy efficiency of locomotion is directly related to body mass for animals smaller than 1 kg body mass.     

Body mass and the energy efficiency of locomotion: lessons from incline running.

Rates of oxygen consumption were measured for two bipedal runners (two species of quail) and two quadrupedal runners (two small species of rodents), with average body masses that ranged from 0.035 to 0.217 kg, trained to run on a treadmill set to horizontal and then to a 10 degrees incline. Rates of oxygen consumption increased linearly with speed for all four species and the rates of increase were significantly higher (P < or = 0.05) for all four species when the animals were run on an incline than when they were run on a horizontal. The estimated metabolic energy cost to lift 1 kg mass 1 m vertically was similar for bipeds and quadrupeds of similar body mass and inversely related to body mass for both running styles. When the data for the animals used in the present study are combined with similar data for adult animals from the literature, the results show that the metabolic energy efficiencies of locomotion, estimated from the cost of vertical work, are the same for bipedal and quadrupedal runners. In both groups, the metabolic energy efficiency of locomotion is directly related to body mass for animals smaller than 1 kg body mass.     

Quadrupedal locomotor performance in two species of arboreal squirrels: predicting energy savings of gliding

Gliding allows mammals to exploit canopy habitats of old-growth forests possibly as a means to save energy. To assess costs of quadrupedal locomotion for a gliding arboreal mammal, we used open-flow respirometry and a variable-speed treadmill to measure oxygen consumption and to calculate cost of transport, excess exercise oxygen consumption, and excess post-exercise oxygen consumption for nine northern flying squirrels (Glaucomys sabrinus) and four fox squirrels (Sciurus niger). Our results indicate that oxygen consumption during exercise by flying squirrels was 1.26–1.65 times higher than predicted based on body mass, and exponentially increased with velocity (from 0.84 ± 0.03 ml O₂ kg⁻¹ s⁻¹ at 0.40 m s⁻¹ to 1.55 ± 0.03 ml O₂ kg⁻¹ s⁻¹ at 0.67 m s⁻¹). Also, cost of transport in flying squirrels increased with velocity, although excess exercise oxygen consumption and excess post-exercise oxygen consumption did not. In contrast, oxygen consumption during exercise for fox squirrels was similar to predicted, varying from 0.51 (±0.02) ml O₂ kg⁻¹ s⁻¹ at 0.63 m s⁻¹ to 0.54 (±0.03) ml O₂ kg⁻¹ s⁻¹ at 1.25 m s⁻¹. In addition, the cost of transport for fox squirrels decreased with velocity, while excess exercise oxygen consumption and excess post-exercise oxygen consumption did not. Collectively, these observations suggest that unlike fox squirrels, flying squirrels are poorly adapted to prolonged bouts of quadrupedal locomotion. The evolution of skeletal adaptations to climbing, leaping, and landing and the development of a gliding membrane likely has increased the cost of quadrupedal locomotion by >50% while resulting in energy savings during gliding and reduction in travel time between foraging patches.     

The energetics of mimicry: the cost of pedestrian transport in a formicine ant and its mimic, a clubionid spider

ABSTRACT. Little information exists on the energetics of locomotion in small insects, and none in small spiders. We examined standard rate of oxygen consumption (SO₂) and net cost of transport (NCOT) in Camponotus sericeiventris (Guerin), a formicine ant, Myrmecotypus rettenmeyeri (Unzicker), its clubionid spider mimic (mean masses 43 and 24 mg, respectively), and an unrelated clubionid, Clubiona barroana (Chickering), mean mass 37mg. All species are from Barro Colorado Island, Panama. NCOT in the species studied was 12, 21 and 27 ml O₂g^-1 km^-1, respectively; NCOT in the two spiders was significantly higher than in the ant. The minimum cost of transport of M. rettenmeyeri (8.6ml O₂g^-1 km^-1), though low, did not differ significantly from the value predicted for an insect of its body mass, while its Y intercept elevation (estimated O₂ at zero running speed/SO₂) was extremely high. Y intercept elevation may be high in spiders as a group, leading to high NCOT at low running speeds, and hence to a tendency to adopt ambush predation. It is also predicted that, to minimize locomotion cost per unit distance, spiders will travel faster than insects of equivalent body mass.     

Walking performance and aerobic and anaerobic metabolism of Carcinus maenas (L.) in sea water at 15°C

Walking performance of the shore crab Carcinus maenas (L.) in sea water at 15 °C was assessed. In large crabs there was an inverse relationship between fatigue time and speed; crabs ran for $\text{\$}\text{?}$10 min at 3.2 m·min^?1 and for only 2 min at 14 m·min^?1. There were linear relationships between oxygen consumption and walking speeds for small and large animals walking at up to 4 m·min^?1 Estimates of maximum oxygen consumption were proportional to W^0.13 whereas inactive consumption is proportional to W^0.44 this resulted in aerobic scope (i.e. the difference between inactive and maximal rates of oxygen consumption) remaining almost constant across a weight range of animals whereas the aerobic expansibility (maximal rates/inactive rates) declined from 7- to 4-fold with increasing size. After a 12-h period without handling (settled animals) the animals could immediately become active and reach maximal rates of oxygen consumption similar to those of animals handled 1 h before the experiment. The aerobic expansibility of these settled animals could range from 21 to 8 times their inactive rates of oxygen consumption in small and large animals respectively. After 10 min of exercise oxygen consumption and whole body lactate levels returned to pre-exercise values within 5 to 25 min. The net oxygen debts range from 16 to 64% of the net oxygen consumption increase during exercise in small and large animals respectively.Calculations of the energy gained from lactate accumulation indicated that the net aerobic energy production during walking was supplemented from 4 to 71 % by anaerobic metabolism in small and large animals respectively. With increasing animal size the decline in aerobic expansibility was offset by an increased capacity for lactate production so that the overall maximum energy production during sustained activity remained almost constant at around seven times the inactive rate. The cost of transport (the net increase in oxygen consumption per g per m) falls with increased walking speed and increased animal size.     

Locomotor energetics in primates: gait mechanics and their relationship to the energetics of vertical and horizontal locomotion

All primates regularly move within three-dimensional arboreal environments and must often climb, but little is known about the energetic costs of this critical activity. Limited previous work on the energetics of incline locomotion suggests that there may be differential selective pressures for large compared to small primates in choosing to exploit a complex arboreal environment. Necessary metabolic and gait data have never been collected to examine this possibility and biomechanical mechanisms that might explain size-based differences in the cost of arboreal movement. Energetics and kinematics were collected for five species of primate during climbing and horizontal locomotion. Subjects moved on a treadmill with a narrow vertical substrate and one with a narrow horizontal substrate at their maximum sustainable speed for 10–20 min while oxygen consumption was monitored. Data during climbing were compared to those during horizontal locomotion and across size. Results show that climbing energetic costs were similar to horizontal costs for small primates (<0.5 kg) but were nearly double for larger species. Spatio-temporal gait characteristics suggest that the relationship between the cost of locomotion and the rate of force production changes between the two locomotor modes. Thus, the main determinants of climbing costs are fundamentally different from those during horizontal locomotion. These new results combining spatiotemporal and energetic data confirm and expand on our previous argument (Hanna et al.: Science 320 (2008) 898) that similar costs of horizontal and vertical locomotion in small primates facilitated the successful occupation of a fine-branch arboreal milieu by the earliest primates.     

Comparison of leucine kinetics in endurance-trained and sedentary humans

Whole body leucine kinetics was compared inendurance-trained athletes and sedentary controls matched for age,gender, and body weight. Kinetic studies were performed during 3 h ofrest, 1 h of exercise (50% maximal oxygen consumption), and 2 h ofrecovery. When leucine kinetics were expressed both per unit of bodyweight and per unit of fat-free mass, both groups demonstrated anincrease in leucine oxidation during exercise(P < 0.01). Trained athletes had agreater leucine rate of appearance during exercise and recovery compared with their sedentary counterparts(P < 0.05) and an increased leucineoxidation at all times on the basis of body weight(P < 0.05). However, all of thesebetween-group differences were eliminated when leucine kinetics werecorrected for fat-free tissue mass. Therefore, correctionof leucine kinetics for fat-free mass may be important whencross-sectional investigations on humans are performed. Furthermore,leucine oxidation, when expressed relative to whole-body oxygenconsumption during exercise, was similar between groups. It isconcluded that there was no difference between endurance-trained andsedentary humans in whole body leucine kinetics during rest, exercise,or recovery when expressed per unit of fat-free tissue mass.

     

The energy cost of embryonic development in fishes and amphibians,with emphasis on new data from the Australian lungfish, <Emphasis Type="Italic">Neoceratodus forsteri</Emphasis>

The rate of oxygen consumption throughout embryonic development is used to indirectly determine the ‘cost’ of development, which includes both differentiation and growth. This cost is affected by temperature and the duration of incubation in anamniote fish and amphibian embryos. The influences of temperature on embryonic development rate, respiration rate and energetics were investigated in the Australian lungfish, Neoceratodus forsteri, and compared with published data. Developmental stage and oxygen consumption rate were measured until hatching, upon which wet and dry gut-free masses were determined. A measure of the cost of development, the total oxygen required to produce 1 mg of embryonic dry tissue, increased as temperature decreased. The relationship between the oxygen cost of development (C, ml mg⁻¹) and dry hatchling mass (M, mg) in fishes and amphibians is described by C = 0.30 M^{0.22 ± 0.13 (95% CI)}, r ² = 0.52. The scaling exponent indicates that the cost of embryonic development increases disproportionally with increasing hatchling mass. At 15 and 20°C, N. forsteri cost of development is significantly lower than the regression mean for all species, and at 25°C is lower than the allometrically scaled data set. Unexpectedly, incubation of N. forsteri is long, despite natural development under relatively warm conditions, and may be related to a large genome size. The low cost of development may be associated with construction of a rather sluggish fish with a low capacity for aerobic metabolism. The metabolic rate is lower in N. forsteri hatchlings than in any other fishes or amphibians at the same temperature, which matches the extremely low aerobic metabolic scope of the juveniles.     

Head‐turning morphologies: Evolution of shape diversity in the mammalian atlas–axis complex

Mammals flex, extend, and rotate their spines as they perform behaviors critical for survival, such as foraging, consuming prey, locomoting, and interacting with conspecifics or predators. The atlas–axis complex is a mammalian innovation that allows precise head movements during these behaviors. Although morphological variation in other vertebral regions has been linked to ecological differences in mammals, less is known about morphological specialization in the cervical vertebrae, which are developmentally constrained in number but highly variable in size and shape. Here, we present the first phylogenetic comparative study of the atlas–axis complex across mammals. We used spherical harmonics to quantify 3D shape variation of the atlas and axis across a diverse sample of species, and performed phylogenetic analyses to investigate if vertebral shape is associated with body size, locomotion, and diet. We found that differences in atlas and axis shape are partly explained by phylogeny, and that mammalian subclades differ in morphological disparity. Atlas and axis shape diversity is associated with differences in body size and locomotion; large terrestrial mammals have craniocaudally elongated vertebrae, whereas smaller mammals and aquatic mammals have more compressed vertebrae. These results provide a foundation for investigating functional hypotheses underlying the evolution of neck morphologies across mammals.     

Capacity for sustained terrestrial locomotion in an insect: Energetics,thermal dependence,and kinematics

Summary The capacity for sustained, terrestrial locomotion in the cockroach. Blaberus discoidalis, was determined in relation to running speed, metabolic cost, aerobic capacity, and ambient temperature (T _a=15, 23, and 34°C; acclimation temperature=24°C). Steady-state thoracic temperature (T _tss) increased linearly with speed at each T _a.The difference between T _tss and T _awas similar at each experimental temperature with a maximum increase of 7°C. Steady-state oxygen consumption (VO_2ss) increased linearly with speed at each T _aand had a low thermal dependence (Q₁₀=1.0-1.4). The minimum cost of locomotion (the slope of the VO_2ss versus speed function) was independent of T _a.Cockroaches attained a maximal oxygen consumption (VO_2max). increased with T _afrom 2.1 ml O₂·g^-1·h^-1 at 15°C to 4.9 ml O₂·g^-1·h^-1 at 23°C, but showed no further increase at 34°C, VO_2max increased 23-fold over resting VO₂ at 23°C, 10-fold at 34°C, and 15-fold at 15°C. Endurance correlated with the speed at which VO_2max was attained (MAS, maximal aerobic speed). Temperature affected the kinematics of locomotion. compared to cockroaches running at the same speed, but higher temperatures (23–34°C), low temperature (15°C) increased protraction time, reduced stride frequency, and reduced stability by increasing body pitching. The thermal independence of the minimum cost of locomotion (C_min), the low thermal dependence of VO_2ss (i.e., y-intercept of the VO_2ss versus speed function), and a typical Q₁₀ of 2.0 for VO_2max combined to increase MAS and endurance in B. discoidalis when T _awas increased from 15 to 23°C. Exerciserelated endothermy enabled running cockroaches to attain a greater VO_2max, metabolic scope, and endurance capacity at 23°C than would be possible if T _tss remained equal to T _a. The MAS of B. discoidalis was similar to that of other arthropods that use trachea, but was 2-fold greater than ectotherms, such as salamanders, frogs, and crabs of a comparable body mass.Abbreviations T _a ambient temperature - T _t thoracic temperature - T _tss steady state thoracic temperature during exercise - T _trest thoracic temperature during rest - VO₂ oxygen consumption - VO_2rest oxygen consumption during rest - VO_2ss steady-state oxygen consumption during exercise - VO_2max maximal oxygen consumption; MAS maximum aerobic speed - C _min minimum cost of locomotion - t _end endurance time     

Integrating the Physiology, Mechanics and Behavior of Rapid Running Ghost Crabs: Slow and Steady Doesn't Always Win the Race

In 1979 Bliss predicted that, "land crabs are and will undoubtedlycontinue to be promising objects of scientific research." Studiesof rapid running ghost crabs support her contention and haveresulted in several general findings relating to locomotionand activity. 1) Energy exchange mechanisms during walking aregeneral and not restricted to quadrupedal and bipedal morphologies.2) "Equivalent gaits," such as trots and gallops, may existin 4-, 6- and 8-legged animals that differ greatly in leg andskeletal (i.e., exo- vs. endoskeletal) design. These findingssupport the hypothesis that terrestrial locomotion in many speciescan modeled by an inverted pendulum or spring-mass system. 3)An open circulatory system and chitin-covered gills do not necessarilylimit the rate at which oxygen consumption can be increasedor the factorial increase oxygen consumption over resting rates.4) Interspecific and intraspecific i.e., ontogenetic) scalingof sub-maximal oxygen consumption and maximal aerobic speedcan differ significantly. 5) Locomotion at speeds above themaximal aerobic speed requiring non-aerobic contributions maybe far more costly than can be predicted from aerobic costsalone. The cost transport may attain a minimum at less thanmaximum speed. 6) The speed which elicits maximal oxygen consumptionduring continuous exercise is attained at moderate walking speedsin crabs and probably other ectotherms. Speeds 15- to 20-foldfaster are possible, but cannot be sustained. 7) The low enduranceassociated with the low maximal oxygen consumption and maximalaerobic speed of ectotherms moving continuously can be increasedor decreased by altering locomotor behavior and moving intermittently.Ectotherms can locomote at high speeds and travel for considerabledistances or remain active for long periods by including restpauses. Alternatively, intense activity with extended exerciseperiods with short pause periods may actually reduce behavioralcapacity or work accomplished relative to continuous activityduring which the behavior is carried out at a lower intensitylevel without pauses.     

Physiological costs of reproduction in the Sydney rock oyster<Emphasis Type="Italic"> Saccostrea glomerata</Emphasis>

In this study, triploid Sydney rock oysters Saccostrea glomerata, which do not reproduce and have only limited gonadal development, were used to calculate the cost of producing and maintaining somatic tissues. The consumption of oxygen was measured and converted to units of energy expended. The consumption of oxygen of diploid oysters, in different stages of the reproductive cycle, was also measured. Knowing the costs of producing and maintaining somatic tissues (obtained from the triploid oysters), it was possible to calculate the energy demand of somatic and reproductive tissues of diploid oysters. The focus of this study was to test whether this method would work, to investigate if this method would give results in accordance with modern life-history theory and to test hypotheses about costs of reproduction in oysters. It was found that in diploid oysters, 27% of the consumed oxygen was needed for reproductive processes. It was also found that the costs of production and maintenance of reproductive tissues were on average 84% of those of somatic tissues. Costs for the production and maintenance of somatic tissues decreased over time. Costs for reproduction also decreased, but were dependent on the stage of gonadal development. If the relative mass of gametes in the gonads was large, the costs were relatively small; if the mass was relatively small, the costs were large. Differences between traits of males and females were never significant, suggesting that reproductive effort and costs were similar in males and females. It was estimated that if diploid oysters did not reproduce, they could gain 64% more somatic ash-free dry mass. Thus, in terms of growth, reproduction is an expensive activity.     

Metabolic costs of capital energy storage in a small‐bodied ectotherm

Reproduction is energetically financed using strategies that fall along a continuum from animals that rely on stored energy acquired prior to reproduction (i.e., capital breeders) to those that rely on energy acquired during reproduction (i.e., income breeders). Energy storage incurs a metabolic cost. However, previous studies suggest that this cost may be minimal for small‐bodied ectotherms. Here I test this assumption. I use a laboratory feeding experiment with the European green crab Carcinus maenas to establish individuals with different amounts of energy storage. I then demonstrate that differences in energy storage account for 26% of the variation in basal metabolic costs. The magnitudes of these costs for any individual crab vary through time depending on the amount of energy it has stored, as well as on temperature‐dependent metabolism. I use previously established relationships between temperature‐ and mass‐dependent metabolic rates, combined with a feasible annual pattern of energy storage in the Gulf of Maine and annual sea surface temperature patterns in this region, to estimate potential annual metabolic costs expected for mature female green crabs. Results indicate that energy storage should incur an ~8% increase in metabolic costs for female crabs, relative to a hypothetical crab that did not store any energy. Translated into feeding, for a medium‐sized mature female (45 mm carapace width), this requires the consumption of an additional ~156 mussels annually to support the metabolic cost of energy storage. These results indicate, contrary to previous assumptions, that the cost of energy storage for small‐bodied ectotherms may represent a considerable portion of their basic operating energy budget. An inability to meet these additional costs of energy storage may help explain the recent decline of green crabs in the Gulf of Maine where reduced prey availability and increased consumer competition have combined to hamper green crab foraging success in recent years.     

Feeding habits of ocellate spot skate,Okamejei kenojei (Müller & Henle, 1841), in coastal waters of Taean,Korea

The feeding habits of Okamejei kenojei were studied using 592 specimens collected in the coastal waters of Taean, Korea from April 2008 to March 2009. O. kenojei is a bottom‐feeding carnivore that consumes mainly shrimp, fishes, and crabs. Its diet also includes small quantities of amphipods, mysids, cephalopods, euphausiids, copepods, isopods, and polychaetes. The total length (TL) of individuals in this study ranged from 8.2 to 49.0 cm. Cluster analysis based on %IRI (index of relative importance) identified three size classes. Group A (< 20 cm TL) ate primarily caridean shrimp and amphipods; group B (20–30 cm TL) ate exclusively shrimp; and group C (> 30 cm TL) ate penaeoidean shrimp, fishes, and crabs. O. kenojei showed ontogenetic changes in feeding habits. Although shrimps were the primary food consumed by all size groups, the proportion of shrimp in the total diet decreased and the consumption of fishes and crabs gradually increased with the body size of O. kenojei. Size of the prey organisms also increased. Smaller individuals fed mainly on small prey, such as amphipods, mysids, and small shrimp, whereas larger individuals preferred larger prey, such as larger shrimp, fishes, and crabs. The size‐related diet breadth and the percentage of empty stomachs were significant; the diet breadth gradually increased with body size, whereas the percentage of empty stomachs decreased. Seasonal changes in the O. kenojei diet were not significant, but shrimp constituted 97.3% of the summer diet by %IRI. Seasonal changes in diet breadth and the percentage of empty stomachs were not significant.     

Caterpillars selected for large body size and short development time are more susceptible to oxygen‐related stress

Recent studies suggest that higher growth rates may be associated with reduced capacities for stress tolerance and increased accumulated damage due to reactive oxygen species. We tested the response of Manduca sexta (Sphingidae) lines selected for large or small body size and short development time to hypoxia (10 kPa) and hyperoxia (25, 33, and 40 kPa); both hypoxia and hyperoxia reduce reproduction and oxygen levels over 33 kPa have been shown to increase oxidative damage in insects. Under normoxic (21 kPa) conditions, individuals from the large‐selected (big‐fast) line were larger and had faster growth rates, slightly longer developmental times, and reduced survival rates compared to individuals from a line selected for small size (small‐fast) or an unselected control line. Individuals from the big‐fast line exhibited greater negative responses to hyperoxia with greater reductions in juvenile and adult mass, growth rate, and survival than the other two lines. Hypoxia generally negatively affected survival and growth/size, but the lines responded similarly. These results are mostly consistent with the hypothesis that simultaneous acquisition of large body sizes and short development times leads to reduced capacities for coping with stressful conditions including oxidative damage. This result is of particular importance in that natural selection tends to decrease development time and increase body size.     

Cost reduction in the cold: heat generated by terrestrial locomotion partly substitutes for thermoregulation costs in Knot Calidris canutus

To test whether heat generated during locomotion substitutes for the thermoregulation cost, oxygen consumption of four post-absorptive temperate-wintering Knot Calidris canutus was measured at air temperatures of 25̀C (thermoneutral) and 10̀C (c. 10̀ below the lower critical temperature) when the birds were at rest at night and during running on a treadmill. After allowing for body mass, the thermoregulation cost at 10̀C was significantly lower in active birds compared with birds at rest. At rest, the birds spent, on average, 0.50 watt (W; range, 0.47-0.57 W) on thermoregulation. During exercise, this cost factor averaged 0.33 W (range, 0.25-0.42 W). The average difference in thermoregulation cost was 35% (ranging from 26% to 49% between individuals) and provides an estimate of the amount of substituted heat. A review of nine studies, all restricted to small birds, showed that substitution is a widespread phenomenon. The consequences of such partial substitution for the annual energetics of Knot wintering in the temperate Wadden Sea v tropical west Africa are examined. Compared with a previous additive model, the model which includes substitution (i.e. the use of heat produced during activity) reduces the differences in maintenance metabolism between the two wintering strategies by 17%, from 1.19 W to 0.99 W.     

Behaviour of juvenile mud crabs Scylla serrata in aquaculture: Response to odours of moulting or injured crabs

Behaviour of juvenile mud crabs, Scylla serrata (70–90 mm carapace width, CW) were observed in response to odours of moulting and injured conspecifics and food (pilchard) under controlled flow conditions using bioassay technique. This study was undertaken to better understand the role that chemical cues have in mediating the attraction of cannibal crabs to moulting crabs in an aquaculture growout facility and thus aid in more successful production. In response to moult odour juvenile S. serrata spent 5.6 ± 1.9% of the time in locomotion but this did not differ significantly (P > 0.05) from that of control odours (seawater, 2.6 ± 1.2%), however a tactile response was observed for moult odour not seen in controls. Juveniles exposed to the odour of injured conspecifics (12.2 ± 2.3% of time in locomotion) and to food (22.7 ± 3.1%) differed significantly from the seawater control (P < 0.05). The possibility that the active agent in moult water is relatively dilute was considered, however varying concentrations of food and crushed conspecific odour failed to demonstrate concentration dependant behavioural results. Variation in the form of size (carapace width) and sex was revealed, so too was preliminary evidence of behavioural differences between crab hatchings. For example, larger crabs (around 80 mm CW) increased tactile investigation of the odour inlet pipe in response to the crab-based odours of moulting (3.5 ± 1.7% of the time) and injured conspecifics (2.6 ± 1.0%), though response to food remained constant over all sizes (10.0% of the time). In another experiment, larger female crabs maintained a high frequency of tactile response to the odour of injured conspecifics. In males, the response was attenuated in larger individuals. Despite no differences being found between moult odour and controls some evidence exists to suggest that a small proportion of crabs were responding to the moult odour and that this still has the potential to cause a dramatic cumulative reduction in growout survival. Variation in size and subsequent behaviour of individuals poses the question of whether behaviour contributes to growth rate and might be a focus of genetic selection for more uniform growth.     

Adult aquatic insects: Potential contributors to riparian food webs in Australia's wet–dry tropics

Abstract Changes in the abundance and biomass of aquatic and terrestrial aerial insects with distance (mid‐stream, 0, 10–15 and 160 m) from lowland streams were examined across the dry season landscape in Kakadu National Park, northern Australia. Malaise traps and sticky intercept traps were used to sample the insects at four streams, spaced over an area of 1650 km². Malaise and intercept catches were dominated by Diptera (flies and midges), both numerically and by biomass. Chironomid midges were the most abundant taxon, making up 43.4 and 51.0% of the malaise and intercept trap catches, respectively. However, most chironomids were small (less than 3 mm body length), contributing 34.9% to intercept trap biomass, but only 5.2% in malaise traps. Ceratopogonid midges and caddisflies (Trichoptera) accounted for most of the remaining adult aquatic insects. Major terrestrial components were Diptera and Hymenoptera in malaise traps and Coleoptera and Diptera in intercept traps. The total abundance and biomass of insects were much greater over streams and along the water's edge than in riparian (10–15 m) and savanna (160 m) habitats primarily because of the presence of large numbers of adult aquatic insects. The abundance and biomass of terrestrial insects in malaise traps showed no relationship with distance, but intercept trap catches suggested slightly greater abundances over the water and at the water's edge. The great abundance of aquatic insects relative to terrestrial insects close to streams suggests that they have the potential to be an important component of the diets of riparian insectivores, and predation may be an important pathway by which aquatic nutrients and energy are moved into terrestrial food webs.     

Effects of Temperature and Water Loss on Terrestrial Locomotor Performance in Land Crabs: Integrating Laboratory and Field Studies

SYNOPSIS. Terrestrial and semi-terrestrial crustaceans are exposedto fluctuations in ambient temperature and conditions that favorevaporative water loss. These environmental stresses alter performancelimits in the laboratory and behavior in the field. The maximalrate of oxygen consumption, maximum aerobic speed, and endurancecapacity are greater at a body temperature (T_b) of 24°Cthan at 15°C or 30°C in the ghost crab, Ocypode quadrata.The total metabolic cost to move at the same relative speedis greater at a T_b of 24°C than at 15°C. Slower aerobickinetics at 15°C result in a smaller relative contributionof oxidative metabolism to total metabolic cost. However, therelative contributions from accelerated glycolysis are similarat both temperatures. When locomotion is intermittent, the totaldistance traveled before fatigue can be similar at Tbs of 15and 24°C but result from different movement and pause durationsat these temperatures. Performance limits of the ghost crabare negatively affected by dehydration and are sensitive torates of water loss. In the laboratory, endurance capacity ofthe fiddler crab, Uca pugilator, is greater at a T_b of 30°Cthan at 25°C. In the field, freely moving fiddler crabswith a T_b of 30°C travel at faster mean preferred speeds,as determined by motion analysis, than crabs at 25°C. Datafor land crabs support and advance general ectothermic modelsfor the effects of temperature and dehydration on locomotorperformance.