Differential design for hopping in two species of wallabies.

Hindlimb musculoskeletal anatomy and steady speed over ground hopping mechanics were compared in two species of macropod marsupials, tammar wallabies and yellow-footed rock wallabies (YFRW). These two species are relatively closely related and are of similar size and general body plan, yet they inhabit different environments with presumably different musculoskeletal demands. Tammar wallabies live in relatively flat, open habitat whereas yellow-footed rock wallabies inhabit steep cliff faces. The goal of this study was to explore musculoskeletal differences between tammar wallabies and yellow-footed rock wallabies and determine how these differences influence each species' hopping mechanics. We found the cross-sectional area of the combined ankle extensor tendons of yellow-footed rock wallabies was 13% greater than that of tammar wallabies. Both species experienced similar ankle joint moments during steady-speed hopping, however due to a lower mechanical advantage at this joint, tammar wallabies produced 26% more muscle force. Thus, during moderate speed hopping, yellow-footed rock wallabies operated with 38% higher tendon safety factors, while tammar wallabies were able to store 73% more elastic strain energy (2.18 J per leg vs. 1.26 J in YFRW). This likely reflects the differing demands of the environments inhabited by these two species, where selection for non-steady locomotor performance in rocky terrain likely requires trade-offs in locomotor economy.

Routine metabolic rate of southern bluefin tuna (Thunnus maccoyii).

Routine metabolic rate (RMR) was measured in fasting southern bluefin tuna, Thunnus maccoyii, the largest tuna species studied so far (body mass=19.6 kg (+/-1.9 SE)). Mean mass-specific RMR was 460 mg kg(-1) h(-1) (+/-34.9) at a mean water temperature of 19 degrees C. When evaluated southern bluefin tuna standard metabolic rate (SMR) is added to published values of other tuna species, there is a strong allometeric relationship with body mass (423 M(0.86), R(2)=0.97). This demonstrates that tuna interspecific SMR scale with respect to body mass similar to that of other active teleosts, but is approximately 4-fold higher. However, RMR (not SMR) is most appropriate in ram-ventilating species that are physiologically unable to achieve complete rest. Respiration was measured in a large (250,000 l) flexible polypropylene respirometer (mesocosm respirometer) that was deployed within a marine-farm sea cage for 29 days. Fasted fish were maintained within the respirometer up to 42 h while dissolved oxygen dropped by 0.056 (+/-0.004) mg l(-1) h(-1). Fish showed no obvious signs of stress. They swam at 1.1 (+/-0.1) fork lengths per second and several fed within the respirometer immediately after measurements.

Control of jaw movements in two species of macropodines (Macropus eugenii and Macropus rufus).

The masticatory motor patterns of three tammar wallabies and two red kangaroos were determined by analyzing the pattern of electromyographic (EMG) activity of the jaw adductors and correlating it with lower jaw movements, as recorded by digital video and videoradiography. Transverse jaw movements were limited by the width of the upper incisal arcade. Molars engaged in food breakdown during two distinct occlusal phases characterized by abrupt changes in the direction of working-side hemimandible movement. Separate orthal (Phase I) and transverse (Phase II) trajectories were observed. The working-side lower jaw initially was drawn laterally by the balancing-side medial pterygoid and then orthally by overlapping activity in the balancing- and working-side temporalis and the balancing-side superficial masseter and medial pterygoid. Transverse movement occurred principally via the working-side medial pterygoid and superficial masseter. This pattern contrasted to that of placental herbivores, which are known to break down food when they move the working-side lower jaw transversely along a relatively longer linear path without changing direction during the power stroke. The placental trajectory results from overlapping activity in the working- and balancing-side adductor muscles, suggesting that macropods and placental herbivores have modified the primitive masticatory motor pattern in different ways.

Adaptive changes in trabecular architecture in relation to functional strain patterns and disuse.

Principal strains and their orientation, determined from in vivo and in situ strains recorded from the lateral cortical surface of the calcaneus of potoroos (a small marsupial) during treadmill exercise and tension applied via the Achilles tendon, were compared with the underlying trabecular architecture and its alignment to test Wolff's "trajectorial theory" of trabecular alignment. In vivo and in situ principal compressive strains (-800 to -2000 mu e) were found to be aligned (mean 161 +/- 7 degrees) close to the preferred alignment (160 degrees) of underlying trabeculae within the calcaneal metaphysis [a second trabecular arcade was closely aligned (70 degrees) with the direction (71 degrees) of principal tensile strain]. This finding represents quantitative verification of Wolff's trajectorial theory of trabecular alignment. These trabecular alignments, as measured by trabecular anisotropy (TbAn, the ratio of horizontal: vertical intercepts), remained unchanged (p > 0.05) after 8 weeks of disuse. However, trabecular bone volume fraction (BV/TV, -35%), trabecular thickness (TbTh, -25%), and trabecular number (TbN, -16%) were reduced for the tenotomized calcaneii relative to their contralateral controls (p < 0.001 to < 0.003). The reduction in trabecular number was associated with a corresponding increase in trabecular spacing (TbSp, +30%). Together, these results suggest that once trabecular alignment is established during growth (along the directions of principal strain during locomotion), it is not altered when functional strains are removed.

Metabolic rates associated with distress and begging calls in birds.

Rates of oxygen consumption during begging behaviour in nestlings of seven species and distress call behaviour in adults of eight species of Australian birds were measured. A transparent mask coupled to an open-flow respirometry system was used, and calling was elicited by the presentation of food or by the perceived threat of a predator. Distress calling significantly increased oxygen consumption above the measured resting levels in six of the species of birds tested. The factorial increase in oxygen consumption during distress calling was independent of body mass. In most cases, begging behaviour in juvenile birds caused a significant increase in metabolic rate, with some individuals showing factorial increases over fourfold. There was a significant negative correlation between body mass and the factorial cost of begging behaviour.

Aquatic and terrestrial locomotory energetics in a toad and a turtle: a search for generalisations among ectotherms.

Murray short-necked turtles were trained to walk on a motorised treadmill and to swim in a recirculating flume. Through filmed records, the frequency of limb movement and the time that thrust was directed against the substrate were measured. The animals wore masks when walking and accessed air when swimming from a ventilated capsule placed on top of the water surface. Measurement of the exhalant O(2) and CO(2) levels from these devices enabled the measurement of metabolic rates. Equivalent data were obtained from swimming and hopping cane toads, although repeatable measures of limb frequency and contact times were not obtained due to the intermittent form of locomotion in this species. Comparing the cost of transport, the energy required to transport a mass of animal over a unit distance, with other animals showed that toads do not have a cheap form of terrestrial locomotion, but turtles do; turtles use half the cost predicted from their body mass. This economy of locomotion is consistent with what is known about turtle muscle, the mechanics of their gait, and the extremely long contact time for a limb with the substrate. Swimming in toads is energetically expensive, whereas turtles, on the basis of mass, use about the same energy to transport a unit mass as an equivalent-size fish. The data were compared with the predictions of the Kram-Taylor hypothesis for locomotory scaling, and walking turtles were found to provide a numerical fit. The data show that both terrestrial and aquatic locomotory energetics in toads are generally higher than predictions on the basis of mass, whereas in turtles they are lower.

Ventilation and metabolism in a large semifossorial marsupial: the effect of graded hypoxia and hypercapnia.

Metabolic and ventilatory variables were measured in a large semifossorial marsupial, the hairy-nosed wombat (Lasiorhinus latifrons, 21.9 kg). In normoxia, the rate of oxygen consumption was 63% of that predicted for a similar-sized marsupial, and the level of ventilation (V(E)) was such that the convective requirement (V(E)/VO2) was similar to other mammals. Exposure to hypercapnia (5% CO(2)) evoked a hyperventilatory response (3.55 x normoxia) that was no different to that observed for epigeal (surface-dwelling) marsupials; the increase in V(E) was primarily achieved with an increase in tidal volume. Exposure to hypoxia (15% to 8% O(2)) resulted in a hyperventilation (principally through an increase in frequency), although the response was blunted (in 8% O(2), 1.85 x normoxia) and only at the severest levels did hypometabolism contribute. The attenuated response to hypoxia in the wombat is presumably a reflection of a semifossorial lifestyle and a tolerance to this respiratory stimulant.

Partitioning of respiration between the gills and air-breathing organ in response to aquatic hypoxia and exercise in the pacific tarpon, Megalops cyprinoides.

The evolution of air-breathing organs (ABOs) is associated not only with hypoxic environments but also with activity. This investigation examines the effects of hypoxia and exercise on the partitioning of aquatic and aerial oxygen uptake in the Pacific tarpon. The two-species cosmopolitan genus Megalops is unique among teleosts in using swim bladder ABOs in the pelagic marine environment. Small fish (58-620 g) were swum at two sustainable speeds in a circulating flume respirometer in which dissolved oxygen was controlled. For fish swimming at 0.11 m s(-1) in normoxia (Po2 = 21 kPa), there was practically no air breathing, and gill oxygen uptake was 1.53 mL kg(-0.67) min(-1). Air breathing occurred at 0.5 breaths min(-1) in hypoxia (8 kPa) at this speed, when the gills and ABOs accounted for 0.71 and 0.57 mL kg(-0.67) min(-1), respectively. At 0.22 m s(-1) in normoxia, breathing occurred at 0.1 breaths min(-1), and gill and ABO oxygen uptake were 2.08 and 0.08 mL kg(-0.67) min(-1), respectively. In hypoxia and 0.22 m s(-1), breathing increased to 0.6 breaths min(-1), and gill and ABO oxygen uptake were 1.39 and 1.28 mL kg(-0.67) min(-1), respectively. Aquatic hypoxia was therefore the primary stimulus for air breathing under the limited conditions of this study, but exercise augmented oxygen uptake by the ABOs, particularly in hypoxic water.

Modulation of proximal muscle function during level versus incline hopping in tammar wallabies (Macropus eugenii).

We examined the functional role of two major proximal leg extensor muscles of tammar wallabies during level and inclined hopping (12 degrees, 21.3% grade). Previous in vivo studies of hopping wallabies have revealed that, unlike certain avian bipeds, distal hindlimb muscles do not alter their force-length behavior to contribute positive work during incline hopping. This suggests that proximal muscles produce the increased mechanical work associated with moving up an incline. Based on relative size and architectural anatomy, we hypothesized that the biceps femoris (BF), primarily a hip extensor, and the vastus lateralis (VL), the main knee extensor, would exhibit changes in muscle strain and activation patterns consistent with increased work production during incline versus level hopping. Our results clearly support this hypothesis. The BF experienced similar activation patterns during level and incline hopping but net fascicle shortening increased (-0.5% for level hopping versus -4.2% for incline hopping) during stance when the muscle likely generated force. Unlike the BF, the VL experienced active net lengthening during stance, indicating that it absorbs energy during both level and incline hopping. However, during incline hopping, net lengthening was reduced (8.3% for level hopping versus 3.9% for incline hopping), suggesting that the amount of energy absorbed by the VL was reduced. Consequently, the changes in contractile behavior of these two muscles are consistent with a net production of work by the whole limb. A subsidiary aim of our study was to explore possible regional variation within the VL. Although there was slightly higher fascicle strain in the proximal VL compared with the distal VL, regional differences in strain were not significant, suggesting that the overall pattern of in vivo strain is fairly uniform throughout the muscle. Estimates of muscle work based on inverse dynamics calculations support the conclusion that both the BF and VL contribute to the additional work required for incline hopping. However, on a muscle mass-specific basis, these two muscles appear to contribute less than their share. This indicates that other hindlimb muscles, or possibly trunk and back muscles, must contribute substantial work during incline hopping.

Joint work and power associated with acceleration and deceleration in tammar wallabies (Macropus eugenii).

Measurements of joint work and power were determined using inverse dynamics analysis based on ground reaction force and high-speed video recordings of tammar wallabies as they decelerated and accelerated while hopping over a force platform on level ground. Measurements were obtained over a range of accelerations ranging from -6 m s(-2) to 8 m s(-2). The goal of our study was to determine which joints are used to modulate mechanical power when tammar wallabies change speed. From these measurements, we also sought to determine which hind limb muscle groups are the most important for producing changes in mechanical work. Because our previous in vivo analyses of wallaby distal muscle function indicated that these muscle-tendon units favor elastic energy savings and perform little work during steady level and incline hopping, we hypothesized that proximal muscle groups operating at the hip and knee joint are most important for the modulation of mechanical work and power. Of the four hind limb joints examined, the ankle joint had the greatest influence on the total limb work, accounting for 89% of the variation observed with changing speed. The hip and metatarsophalageal (MP) joints also contributed to modulating whole limb work, but to a lesser degree than the ankle, accounting for 28% (energy production) and -24% (energy absorption) of the change in whole limb work versus acceleration, respectively. In contrast, the work produced at the knee joint was independent of acceleration. Based on the results of our previous in vivo studies and given that the magnitude of power produced at the ankle exceeds that which these muscles alone could produce, we conclude that the majority of power produced at the ankle joint is likely transferred from the hip and knee joints via proximal bi-articular muscles, operating in tandem with bi-articular ankle extensors, to power changes in hopping speed of tammar wallabies. Additionally, over the observed range of performance, peak joint moments at the ankle (and resulting tendon strains) did not increase significantly with acceleration, indicating that having thin tendons favoring elastic energy storage does not necessarily limit a tammar wallaby's ability to accelerate or decelerate.

The mechanics of jumping versus steady hopping in yellow-footed rock wallabies.

The goal of our study was to explore the mechanical power requirements associated with jumping in yellow-footed rock wallabies and to determine how these requirements are achieved relative to steady-speed hopping mechanics. Whole body power output and limb mechanics were measured in yellow-footed rock wallabies during steady-speed hopping and moving jumps up to a landing ledge 1.0 m high (approximately 3 times the animals' hip height). High-speed video recordings and ground reaction force measurements from a runway-mounted force platform were used to calculate whole body power output and to construct a limb stiffness model to determine whole limb mechanics. The combined mass of the hind limb extensor muscles was used to estimate muscle mass-specific power output. Previous work suggested that a musculoskeletal design that favors elastic energy recovery, like that found in tammar wallabies and kangaroos, may impose constraints on mechanical power generation. Yet rock wallabies regularly make large jumps while maneuvering through their environment. As jumping often requires high power, we hypothesized that yellow-footed rock wallabies would be able to generate substantial amounts of mechanical power. This was confirmed, as we found net extensor muscle power outputs averaged 155 W kg(-1) during steady hopping and 495 W kg(-1) during jumping. The highest net power measured reached nearly 640 W kg(-1). As these values exceed the maximum power-producing capability of vertebrate skeletal muscle, we suggest that back, trunk and tail musculature likely play a substantial role in contributing power during jumping. Inclusion of this musculature yields a maximum power output estimate of 452 W kg(-1) muscle. Similar to human high-jumpers, rock wallabies use a moderate approach speed and relatively shallow leg angle of attack (45-55 degrees) during jumps. Additionally, initial leg stiffness increases nearly twofold from steady hopping to jumping, facilitating the transfer of horizontal kinetic energy into vertical kinetic energy. Time of contact is maintained during jumping by a substantial extension of the leg, which keeps the foot in contact with the ground.

An allometric study of lung morphology during development in the Australian pelican, Pelicanus conspicillatus, from embryo to adult.

Pelicans produce altricial chicks that develop into some of the largest birds capable of sustained flight. We traced pulmonary morphological development in the Australian pelican, Pelicanus conspicillatus, from third trimester embryos to adults. We described growth and development with allometric relationships between lung components and body mass or lung volume, according to the equation y = ax(b). Pelican lung volume increased faster than body mass (b = 1.07). Relative to lung volume, the airways and vascular spaces increased allometrically (b > 1) in embryos, but isometrically (b approximately 1) after hatching. Parabronchial mantle volume decreased (b < 1) prior to hatching and increased isometrically thereafter. Surface area of air capillaries, blood capillaries and the blood-gas barrier increased relative to lung volume (b > 0.67) before and after hatching. Barrier thickness decreased before hatching, remained constant in juveniles and decreased by adulthood. The anatomical diffusing capacity significantly increased before hatching (b = 4.44) and after hatching (b = 1.26). Although altricial pelicans developed pulmonary complexity later than precocial turkeys, the volume-specific characteristics were similar. However, lungs of volant adult pelicans became significantly larger, with a greater capacity for gas exchange, than lungs of terrestrial turkeys. Exchange characteristics of growing pelican lungs were inferior to those of adult birds of 26 other species, but converged with them at maturity.

The energetics and cardiorespiratory correlates of mammalian terrestrial locomotion.

Energy costs of locomotion in mammals can be predicted from running speed and body mass, with the minimum cost decreasing regularly with increasing mass (Mb-0.30). The predictive value of this model is surprising, given the differences in gait and limb structure among mammals. The decrease in mass-specific cost cannot be explained by the work done in moving the limbs and the centre of mass, as animals of different sizes do the same amount of work to move a unit mass a unit distance. The magnitude of the muscle forces involved and the shortening velocity are more likely causes. Terrestrial mammals use a variety of gaits to minimise locomotory energy costs with a 'preferred speed' within each of those gaits correlating with the point of greatest economy. The maximum mass-specific energy cost during locomotion is about 10 times the resting level, but there is marked variation among species, especially between wild and domestic forms. The total cost for locomotion in mammals lies between 1 and 6% of the daily energy budget. Hopping is an energetically cheap way of moving in large animals and correlates with phase-locking of respiratory and limb frequencies. This form of coupling is also seen in most other mammals, especially at higher running speeds. Comparison of the relative costs of running, flying and swimming for a given body mass shows a respective decrease, but each of these costs scales similarly with body size.

Scaling of respiratory variables and the breathing pattern in adult marsupials.

Oxygen consumption (VO2) and a number of components of the breathing pattern were measured in 14 species of non-fossorial marsupials ranging in mass from 0.008 to 30 kg. All the exponents of the allometric relationships for VO2, ventilation (VE) and breathing pattern scaled as previously determined for eutherians. However, compared to eutherians, marsupials had significantly lower VO2 and the breathing pattern was deeper (+23%) and slower (-31%). While VE was not significantly below that reported for eutherians it matched VO2 such that VE/VO2 remained mass-independent and at a level similar to that observed in the other infraclasses of mammals. Thus, it would appear that the increase in metabolic rate that occurred during the evolution of mammalian homeothermy was accompanied by parallel changes in VE. It is suggested that these changes in VE were mediated by a shortening of the respiratory cycle, facilitated by the eventual abolition of the end-expiratory pause (TP), and an increase in respiratory drive (VT/TI). In response to 5% CO2 all animals in this study increased their VE by increasing both tidal volume (VT) and frequency (f), predominantly through the removal of TP. The increase in VE was less than previously reported for eutherians, suggesting a reduction in chemosensitivity in marsupials. Furthermore, the similarity in slopes for VE, VT and f between air and 5% CO2 suggests that the gain of the respiratory system is independent of species size within marsupials.

Energetics of locomotion by the Australian water rat (Hydromys chrysogaster): a comparison of swimming and running in a semi-aquatic mammal.

Semi-aquatic mammals occupy a precarious evolutionary position, having to function in both aquatic and terrestrial environments without specializing in locomotor performance in either environment. To examine possible energetic constraints on semi-aquatic mammals, we compared rates of oxygen consumption for the Australian water rat (Hydromys chrysogaster) using different locomotor behaviors: swimming and running. Aquatic locomotion was investigated as animals swam in a water flume at several speeds, whereas water rats were run on a treadmill to measure metabolic effort during terrestrial locomotion. Water rats swam at the surface using alternate pelvic paddling and locomoted on the treadmill using gaits that included walk, trot and half-bound. Water rats were able to run at twice their maximum swimming velocity. Swimming metabolic rate increased with velocity in a pattern similar to the 'humps' and 'hollows' for wave drag experienced by bodies moving at the water surface. Metabolic rate increased linearly during running. Over equivalent velocities, the metabolic rate for running was 13-40 % greater than for swimming. The minimum cost of transport for swimming (2.61 J N-1 m-1) was equivalent to values for other semi-aquatic mammals. The lowest cost for running (2.08 J N-1 m-1) was 20 % lower than for swimming. When compared with specialists at the extremes of the terrestrial-aquatic continuum, the energetic costs of locomoting either in water or on land were high for the semi-aquatic Hydromys chrysogaster. However, the relative costs for H. chrysogaster were lower than when an aquatic specialist attempts to move on land or a terrestrial specialist attempts to swim.

Energetics of locomotion in a monotreme, the echidna Tachyglossus aculeatus.

The steady state oxygen consumption of two echidnas was measured during locomotion on a treadmill. The change in power input with change in velocity is similar to that found in other mammals, but the total energy requirement for locomotion is less. The significance of these findings in an animal with low basal metabolism and distally heavy limbs is discussed.

The relationship between body mass and rate of rewarming from hibernation and daily torpor in mammals.

1. Rewarming rate from torpor and body mass were inversely related in 86 mammals ranging in body mass between 2 and 8500 g. 2. Most of the mammalian taxa investigated showed a similar change of rewarming rate with body mass. Only the insectivores showed a more pronounced increase in rewarming with a decrease in body mass than did the other taxa. The rates of rewarming of marsupials were similar to those of placentals. 3. At low air temperature (Ta), the rate of rewarming of marsupials was not related to body mass, although a strong relationship between the two variables was observed in the same species at high Ta. 4. The slopes relating rewarming rates and body mass of the mammalian groups and taxa analysed here were similar to those obtained earlier for mass-specific basal metabolic rate (BMR) and body mass in mammals, suggesting that the rate of rewarming and BMR are physiologically linked.

Energetic cost of locomotion as a function of ambient temperature and during growth in the marsupial Potorous tridactylus.

In the marsupial, the potoroo, multiple regression analysis shows that ambient temperature makes a minor (2%) contribution towards variation in oxygen consumption with speed. This suggests that the heat generated during running is substituted for heat which would otherwise have to be generated for temperature regulation. Maximum levels of oxygen consumption are also temperature-independent over the range 5-25 degrees C, but plasma lactate concentrations at the conclusion of exercise significantly increase with ambient temperature. Adult potoroos show a linear increase in oxygen consumption with speed, and multiple regression indicates that the most significant factor affecting energy use during running is stride length. Juvenile potoroos have an incremental cost of locomotion about 40% lower than that predicted on the basis of body mass. The smaller animals meet the demands of increasing speed by increasing stride length rather than stride frequency, as would be expected in a smaller species. Our results show that juvenile potoroos diverge significantly from models based only on adult animals in incremental changes in stride frequency, length and the cost of transport, suggesting that they are not simply scaled-down adults.

Central cardiovascular shunts in the perinatal marsupial.

BACKGROUND: Marsupials are born at an early stage of development after a short period of gestation. In this study the nature and timing of closure of the central cardiovascular shunts was investigated. METHODS: Light and scanning electron microscopy were used to determine changes in central cardiovascular shunts in eight marsupial species with gestation periods of between 12.5 and 36.5 days and birth weights ranging from 12.5 mg to 740 mg. Laboratory mice with a birth weight of about 1,000 mg and a gestation period of 21 days were included for comparison. RESULTS: Marsupials have a ductus arteriosus and an interatrial communication. The former closes rapidly after birth in the marsupial; however the interatrial communication is in the form of a fenestrated septum, which closes as a result of tissue proliferation over a period of days after birth. An additional central shunt, an interventricular foramen, was found to persist in three species for a short time after birth. In one species, the eastern native cat, Dasyurus viverrinus, which has a gestation period of about 19 days and low birth weight of about 12.5 mg, in addition to the two common shunts there was a large interventricular communication and septation of the outflow tract was incomplete. CONCLUSION: In adapting from intra-uterine life, it seems that marsupials have adopted different, but equally effective strategies, with regard to the circulatory system.

Oxygen transport and acid-base balance during exercise in the tammar wallaby.

Rates of oxygen consumption (V(O)2), body temperatures and pulmonary blood temperatures, blood gases and blood pH were measured for seven 4.9+/-0.8 (SE) kg tammar wallabies (Macropus eugenii) during rest and during treadmill hopping. For animals resting on the treadmill V(O)2 averaged 0.030+/-0.003 L min(-1). During hopping V(O)2 increased linearly with speed up to 2.5 m sec(-1). Above 2.5 m sec(-1) V(O)2 was independent of hopping speed and averaged 0.340+/-0.004 L min(-1). At rest, rectal temperatures and pulmonary blood temperatures averaged 36 degrees C. During treadmill hopping, rectal temperatures and pulmonary blood temperatures increased similarly, to 39 degrees C. The Pv(CO)2 increased and pHv decreased in proportion to the increased V(O)2. The Pa(CO)2 and pHa were not significantly changed from values for animals resting on the treadmill. Cardiac output (Vb) averaged 0.97+/-0.04 L min(-1) when the wallabies were at rest on the treadmill and increased linearly with treadmill speeds up to 2.5 m sec(-1). Above 2.5 m sec(-1) Vb was independent of hopping speed and averaged 2.9+/-0.04 L min(-1). When data for all speeds were combined, Vb increased linearly with V(O)2. Thus, in spite of their unique mode of locomotion wallabies have maintained relationships between pulmonary ventilation and V(O)2 and between Vb and V(O)2 that are similar to those reported for eutherian mammals.