SPERM WHALE SURFACE ACTIVITY FROM TRACKING BY RADIO AND SATELLITE TAGS1

Three 12-m sperm whales (Physeter catodon) were tagged and tracked west of Dominica in the southeast Caribbean to follow the surfacing patterns and movements of these presumed subadult males. Whale N was tagged in April 1993 with a 30-MHz radio tag and tracked for two days. Whale H was tagged in April 1995 with a 30-MHz radio tag and tracked for 4.6 d. Whale A was tagged in April 1995 with a satellite-monitored tag tracked by ARGOS for 21.5 d, the first four of which were concurrent with the tracking of Whale H, an associate. The tagged whales remained west of Dominica for at least 2, 5, and 13 d, respectively. Whales N and A then moved southward to waters off Martinique. There were no apparent effects on the whales by tagging or the presence of the tags. The whales averaged speeds of 2.6-3.5 km/h. Surfacings, indicated by tag signals, were of two types: short surfacings apparently primarily for respiration, averaging 7-10.5 min between repeated longer dives, occurring day and night; and extended surfacings seemingly for rest and social interactions with conspecifics, occurring mostly in daylight. Whales were near the surface for 20.4%–22.6% of the total time (26.6%–27.1% during the day and 14.9%–17.1% at night). Delayed blowing was observed as Whale N surfaced for 8.3 min between 47- and 45-min dives but delayed the first of its 31 blows for 1.5 min after surfacing.     

Antarctic killer whales make rapid,round-trip movements to subtropical waters: evidence for physiological maintenance migrations?

Killer whales (Orcinus orca) are important predators in high latitudes, where their ecological impact is mediated through their movements. We used satellite telemetry to provide the first evidence of migration for killer whales, characterized by fast (more than 12 km h(-1), 6.5 knots) and direct movements away from Antarctic waters by six of 12 type B killer whales tagged when foraging near the Antarctic Peninsula, including all tags transmitting for more than three weeks. Tags on five of these whales revealed consistent movements to subtropical waters (30-37° S) off Uruguay and Brazil, in surface water temperatures ranging from -1.9°C to 24.2°C; one 109 day track documented a non-stop round trip of almost 9400 km (5075 nmi) in just 42 days. Although whales travelled slower in the warmest waters, there was no obvious interruption in swim speed or direction to indicate calving or prolonged feeding. Furthermore, these movements were aseasonal, initiating over 80 days between February and April; one whale returned to within 40 km of the tagging site at the onset of the austral winter in June. We suggest that these movements may represent periodic maintenance migrations, with warmer waters allowing skin regeneration without the high cost of heat loss: a physiological constraint that may also affect other whales.     

MOVEMENTS OF NORTH PACIFIC BLUE WHALES DURING THE FEEDING SEASON OFF SOUTHERN CALIFORNIA AND THEIR SOUTHERN FALL MIGRATION1

The satellite-acquired locations of 10 blue whales (Balaenoptera musculus) tagged off southern California with Argos radio tags were used to identify (1) their movements during the late summer feeding season; (2) the routes and rate of travel for individuals on their southern fall migration; and (3) a possible winter calving/breeding area. Whales were tracked from 5.1 to 78.1 d and from 393 to 8,668 km. While in the Southern California Bight, most of the locations for individual whales were either clumped or zigzagged in pattern, suggesting feeding or foraging (searching for prey). Average speeds ranged from 2.4 to 7.2 km/h. One whale moved north to Cape Mendocino, and four migrated south along the Baja California, Mexico coast, two passing south of Cabo San Lucas on the same day. One of the latter whales traveled an additional 2,959 km south in 30.5 d to within 450 km of the Costa Rican Dome (CRD), an upwelling feature. The timing of this migration suggests the CRD may be a calving/breeding area for North Pacific blue whales. Although blue whales have previously been sighted in the Eastern Tropical Pacific (ETP), this is the first evidence that whales from the feeding aggregation off California range that far south. The productivity of the CRD may allow blue whales to feed during their winter calving/breeding season, unlike gray whales (Eschrichtius robustus) and humpbacks (Megaptera novaeangliae) which fast during that period.     

Seasonal and Geographic Variation of Southern Blue Whale Subspecies in the Indian Ocean

Understanding the seasonal movements and distribution patterns of migratory species over ocean basin scales is vital for appropriate conservation and management measures. However, assessing populations over remote regions is challenging, particularly if they are rare. Blue whales (Balaenoptera musculus spp) are an endangered species found in the Southern and Indian Oceans. Here two recognized subspecies of blue whales and, based on passive acoustic monitoring, four “acoustic populations” occur. Three of these are pygmy blue whale (B.m. brevicauda) populations while the fourth is the Antarctic blue whale (B.m. intermedia). Past whaling catches have dramatically reduced their numbers but recent acoustic recordings show that these oceans are still important habitat for blue whales. Presently little is known about the seasonal movements and degree of overlap of these four populations, particularly in the central Indian Ocean. We examined the geographic and seasonal occurrence of different blue whale acoustic populations using one year of passive acoustic recording from three sites located at different latitudes in the Indian Ocean. The vocalizations of the different blue whale subspecies and acoustic populations were recorded seasonally in different regions. For some call types and locations, there was spatial and temporal overlap, particularly between Antarctic and different pygmy blue whale acoustic populations. Except on the southernmost hydrophone, all three pygmy blue whale acoustic populations were found at different sites or during different seasons, which further suggests that these populations are generally geographically distinct. This unusual blue whale diversity in sub-Antarctic and sub-tropical waters indicates the importance of the area for blue whales in these former whaling grounds.     

Hybridization of Southern Hemisphere blue whale subspecies and a sympatric area off Antarctica: impacts of whaling or climate change?

Understanding the degree of genetic exchange between subspecies and populations is vital for the appropriate management of endangered species. Blue whales (Balaenoptera musculus) have two recognized Southern Hemisphere subspecies that show differences in geographic distribution, morphology, vocalizations and genetics. During the austral summer feeding season, the Antarctic blue whale (B. m. intermedia) is found in polar waters and the pygmy blue whale (B. m. brevicauda) in temperate waters. Here, we genetically analyzed samples collected during the feeding season to report on several cases of hybridization between the two recognized blue whale Southern Hemisphere subspecies in a previously unconfirmed sympatric area off Antarctica. This means the pygmy blue whales using waters off Antarctica may migrate and then breed during the austral winter with the Antarctic subspecies. Alternatively, the subspecies may interbreed off Antarctica outside the expected austral winter breeding season. The genetically estimated recent migration rates from the pygmy to Antarctic subspecies were greater than estimates of evolutionary migration rates and previous estimates based on morphology of whaling catches. This discrepancy may be due to differences in the methods or an increase in the proportion of pygmy blue whales off Antarctica within the last four decades. Potential causes for the latter are whaling, anthropogenic climate change or a combination of these and may have led to hybridization between the subspecies. Our findings challenge the current knowledge about the breeding behaviour of the world's largest animal and provide key information that can be incorporated into management and conservation practices for this endangered species.     

Movements of satellite-monitored humpback whales on their feeding ground along the Antarctic Peninsula

Humpback whales were instrumented with satellite transmitters off the western Antarctic Peninsula in January of 2004–2006 to examine their movement patterns and habitat use. Whales were tracked from 4 to 80 days (mean = 36.5 days). Distance and travel rate estimates for nine individuals ranged from 223 to 4,356 km and from 17 to 75 km/day, respectively. Considerable individual variation was observed in direction, speed and range of movements. The overall pattern was characterized by short- and long-distance movements between presumed foraging areas with relatively short residency times. Travel rates were lower at these sites, characterized by erratic movements, than during traveling between them. Area usage for six individuals based on the 95% fixed kernel home range with least squares cross-validation ranged from 2,771 to 172,356 km². The management boundary between the feeding grounds associated with Breeding Stocks G and A needs revision, as current available data suggest it should be located to the east of 50°W. This study is the first to present detailed information on the movements of humpback whales in the Southern Ocean.     

First direct measurements of behavioural responses by Cuvier's beaked whales to mid-frequency active sonar

Most marine mammal­ strandings coincident with naval sonar exercises have involved Cuvier''s beaked whales (Ziphius cavirostris). We recorded animal movement and acoustic data on two tagged Ziphius and obtained the first direct measurements of behavioural responses of this species to mid-frequency active (MFA) sonar signals. Each recording included a 30-min playback (one 1.6-s simulated MFA sonar signal repeated every 25 s); one whale was also incidentally exposed to MFA sonar from distant naval exercises. Whales responded strongly to playbacks at low received levels (RLs; 89–127 dB re 1 µPa): after ceasing normal fluking and echolocation, they swam rapidly, silently away, extending both dive duration and subsequent non-foraging interval. Distant sonar exercises (78–106 dB re 1 µPa) did not elicit such responses, suggesting that context may moderate reactions. The observed responses to playback occurred at RLs well below current regulatory thresholds; equivalent responses to operational sonars could elevate stranding risk and reduce foraging efficiency.     

Fine‐scale movements and behaviors of coyotes (Canis latrans) during their reproductive period

In canids, resident breeders hold territories but require different resources than transient individuals (i.e., dispersers), which may result in differential use of space, land cover, and food by residents and transients. In the southeastern United States, coyote (Canis latrans) reproduction occurs during spring and is energetically demanding for residents, but transients do not reproduce and therefore can exhibit feeding behaviors with lower energetic rewards. Hence, how coyotes behave in their environment likely differs between resident and transient coyotes. We captured and monitored 36 coyotes in Georgia during 2018–2019 and used data from 11 resident breeders, 12 predispersing residents (i.e., offspring of resident breeders), and 11 transients to determine space use, movements, and relationships between these behaviors and landcover characteristics. Average home range size for resident breeders and predispersing offspring was 20.7 ± 2.5 km² and 50.7 ± 10.0 km², respectively. Average size of transient ranges was 241.4 ± 114.5 km². Daily distance moved was 6.3 ± 3.0 km for resident males, 5.5 ± 2.7 km for resident females, and 6.9 ± 4.2 km for transients. We estimated first‐passage time values to assess the scale at which coyotes respond to their environment, and used behavioral change‐point analysis to determine that coyotes exhibited three behavioral states. We found notable differences between resident and transient coyotes in regard to how landcover characteristics influenced their behavioral states. Resident coyotes tended to select for areas with denser vegetation while resting and foraging, but for areas with less dense vegetation and canopy cover when walking. Transient coyotes selected areas closer to roads and with lower canopy cover while resting, but for areas farther from roads when foraging and walking. Our findings suggest that behaviors of both resident and transient coyotes are influenced by varying landcover characteristics, which could have implications for prey.     

High Source Levels and Small Active Space of High-Pitched Song in Bowhead Whales (Balaena mysticetus)

The low-frequency, powerful vocalizations of blue and fin whales may potentially be detected by conspecifics across entire ocean basins. In contrast, humpback and bowhead whales produce equally powerful, but more complex broadband vocalizations composed of higher frequencies that suffer from higher attenuation. Here we evaluate the active space of high frequency song notes of bowhead whales (Balaena mysticetus) in Western Greenland using measurements of song source levels and ambient noise. Four independent, GPS-synchronized hydrophones were deployed through holes in the ice to localize vocalizing bowhead whales, estimate source levels and measure ambient noise. The song had a mean apparent source level of 185±2 dB rms re 1 µPa @ 1 m and a high mean centroid frequency of 444±48 Hz. Using measured ambient noise levels in the area and Arctic sound spreading models, the estimated active space of these song notes is between 40 and 130 km, an order of magnitude smaller than the estimated active space of low frequency blue and fin whale songs produced at similar source levels and for similar noise conditions. We propose that bowhead whales spatially compensate for their smaller communication range through mating aggregations that co-evolved with broadband song to form a complex and dynamic acoustically mediated sexual display.     

Aerial photogrammetry and tag-derived tissue density reveal patterns of lipid-store body condition of humpback whales on their feeding grounds

Monitoring the body condition of free-ranging marine mammals at different life-history stages is essential to understand their ecology as they must accumulate sufficient energy reserves for survival and reproduction. However, assessing body condition in free-ranging marine mammals is challenging. We cross-validated two independent approaches to estimate the body condition of humpback whales (Megaptera novaeangliae) at two feeding grounds in Canada and Norway: animal-borne tags (n = 59) and aerial photogrammetry (n = 55). Whales that had a large length-standardized projected area in overhead images (i.e. whales looked fatter) had lower estimated tissue body density (TBD) (greater lipid stores) from tag data. Linking both measurements in a Bayesian hierarchical model to estimate the true underlying (hidden) tissue body density (uTBD), we found uTBD was lower (−3.5 kg m⁻³) in pregnant females compared to adult males and resting females, while in lactating females it was higher (+6.0 kg m⁻³). Whales were more negatively buoyant (+5.0 kg m⁻³) in Norway than Canada during the early feeding season, possibly owing to a longer migration from breeding areas. While uTBD decreased over the feeding season across life-history traits, whale tissues remained negatively buoyant (1035.3 ± 3.8 kg m⁻³) in the late feeding season. This study adds confidence to the effectiveness of these independent methods to estimate the body condition of free-ranging whales.     

A bowhead whale in the eastern North Pacific

Bowhead whales occur in the Arctic year‐round. Their movements are largely correlated with seasonal expansions and reductions of sea ice, but a few recent extralimital sightings have occurred in the eastern and western North Atlantic and one was also documented in the western North Pacific over 50 years ago. Here we present details of a juvenile bowhead whale that was photographed and filmed from above and below the water while it was skim‐feeding in Caamaño Sound, BC, Canada on May 31, 2016. This sighting occurred over 2000 km southeast from the nearest known range for this species in the Bering Sea at a time that most bowhead whales in that region would have been migrating northeast. This sighting represents the first and only documentation of a bowhead whale in the eastern North Pacific to date.     

Assessment of wound healing of tagged gray (Eschrichtius robustus) and blue (Balaenoptera musculus) whales in the eastern North Pacific using long‐term series of photographs

Tags have been used to examine migration routes and habitat use of large whales for >40 yr, however, evaluation of tag wound healing has largely been short‐term, anecdotal or generalized. This study developed methods for systematic photographic assessment of long‐term external consequences of tag placement, to determine potential differences in wound healing between species and tag types and thus advise future tagging efforts to possibly minimize undesirable side effects. Tag site appearance and healing characteristics were evaluated by two reviewers and a time series evaluated by five veterinarians from photographs during 995 postdeployment encounters with 34 gray and 63 blue whales tagged in the North Pacific. Blue whale resightings were less frequent, but spanned a longer time period due to earlier tag deployments than the more frequent gray whale follow‐up observations. Swelling occurred in 74% of reencountered gray whales, with the highest frequency 6 mo postdeployment. Swellings were common in blue whales with early tag designs but rare with current models. Depressions occurred in 82% of gray and 71% of blue whales. This study demonstrates the value of follow‐up studies of tagged animals and systematic scoring of photographs to quantitatively compare tag response.     

Crossing Latitudes—Long-Distance Tracking of an Apex Predator

Tiger sharks (Galeocerdo cuvier) are apex predators occurring in most tropical and warm temperate marine ecosystems, but we know relatively little of their patterns of residency and movement over large spatial and temporal scales. We deployed satellite tags on eleven tiger sharks off the north-western coast of Western Australia and used the Brownian Bridge kernel method to calculate home ranges and analyse movement behaviour. One individual recorded one of the largest geographical ranges of movement ever reported for the species, travelling over 4000 km during 517 days of monitoring. Tags on the remainder of the sharks reported for shorter periods (7-191 days). Most of these sharks had restricted movements and long-term (30-188 days) residency in coastal waters in the vicinity of the area where they were tagged. Core home range areas of sharks varied greatly from 1166.9 to 634,944 km². Tiger sharks spent most of their time in water temperatures between 23°-26°C but experienced temperatures ranging from 6°C to 33°C. One shark displayed seasonal movements among three distinct home range cores spread along most of the coast of Western Australia and generalized linear models showed that this individual had different patterns of temperature and depth occupancy in each region of the coast, with the highest probability of residency occurring in the shallowest areas of the coast with water temperatures above 23°C. These results suggest that tiger sharks can migrate over very large distances and across latitudes ranging from tropical to the cool temperate waters. Such extensive long-term movements may be a key element influencing the connectivity of populations within and among ocean basins.     

Separating pygmy and Antarctic blue whales using long-forgotten ovarian data

Pygmy blue whales ( Balaenoptera musculus brevicauda ) are ≤24.1 m and are generally found north of 52°S in summer, whereas the more southerly Antarctic blue whales ( B. m. intermedia ) may exceed 30 m. Previous assessments have assumed that catches and recent surveys south of 60°S recorded Antarctic blue whales, but these may have included pygmy blue whales. Here, we use ovarian corpora, which accumulate with ovulations and hence with length, to separate these subspecies. The resulting Bayesian mixture model, applied to 1,380 Northern Region (north of 52°S and 35°–180°E) and 3,844 Southern Ocean (south of 52°S) blue whales, estimated that only 0.1% (95% credibility intervals 0.0%–0.4%) of the Antarctic region blue whales were pygmy blue whales and, unexpectedly, found significantly lower lifetime ovulation counts for pygmy blue whales than for Antarctic blue whales (7.6 vs . 13.6). Over four decades, despite substantial depletion of Antarctic blue whales, there was no trend in the estimated proportion of pygmy blue whales in the Antarctic. Several lines of investigation found no evidence for sizeable numbers of pygmy blue whales in ovarian corpora data collected in the 1930s, as was previously hypothesized.     

SPERM WHALES TAGGED WITH TRANSPONDERS AND TRACKED UNDERWATER BY SONAR

Abstract: Two sperm whales tagged with acoustic transponder tags were tracked by sonar during a cruise from 16 to 30 October 1991 in the southeast Caribbean west of Dominica Island. The whales dove to depths of 400–600 m and more, including a dive to 1,185 m and one possibly to 2,000 m. They were tracked for periods of 3–14 h, over distances of 8.5–40 km. The tagged whales were found together four and eight days after tagging, and were tracked simultaneously for 13 h, over 31 km. Whale movements on different days at the surface averaged from 0.68 to 0.82 m/set, with dive descent rates from 0.82 to 1.13 m/set, ascent rates from 0.74 to 1.16 m/set, and horizontal movement during dives from 0.76 to 1.29 m/set. Dives lasted from 18 min to 1 h and 13 min, averaging 33 and 41 min on different days. Every track ended when tag signals became obscured at night by dense biological scatterers concentrated in offshore areas where the whales were diving. Both tagged whales appear to have been males of 15 and 11m, each dominant in different groups; but when together the larger whale was dominant, as evidenced by chases and agonistic vocalizations. The whales did not appear to react to the tags or to the sounds associated with tracking (30, 32, and 36 kHz).     

Morphometric analysis of Chilean blue whales and implications for their taxonomy

In the Southern Hemisphere, blue whales are currently divided into two subspecies, Antarctic blue whales (Balaenoptera musculus intermedia) and pygmy blue whales (B. m. brevicauda), but there is some debate about whether Chilean blue whales should also be considered as a separate subspecies. Here, we provide novel morphometric data to directly address this taxonomic question from a biological survey of 60 blue whales taken during the 1965/1966 Chilean whaling season. The data show that maximum body length and mean body length of both sexually mature females and males for Chilean blue whales are intermediate between pygmy and Antarctic blue whales; and that fluke-anus lengths of Chilean blue whales are significantly different from pygmy blue whales, but not necessarily from Antarctic blue whales. There is also some support from the data that snout-eye measurements are different among all three groups. These data provide further confirmation that Chilean blue whales are a distinct population requiring separate management from other blue whale populations, and are also consistent with suggestions that Chilean blue whales are not the same subspecies as pygmy blue whales.     

Aerobic Capacity,Activity Levels and Daily Energy Expenditure in Male and Female Adolescents of the Kenyan Nandi Sub-Group

The relative importance of genetic and socio-cultural influences contributing to the success of east Africans in endurance athletics remains unknown in part because the pre-training phenotype of this population remains incompletely assessed. Here cardiopulmonary fitness, physical activity levels, distance travelled to school and daily energy expenditure in 15 habitually active male (13.9±1.6 years) and 15 habitually active female (13.9±1.2) adolescents from a rural Nandi primary school are assessed. Aerobic capacity () was evaluated during two maximal discontinuous incremental exercise tests; physical activity using accelerometry combined with a global positioning system; and energy expenditure using the doubly labelled water method. The of the male and female adolescents were 73.9±5.7 ml^. kg^−1. min⁻¹ and 61.5±6.3 ml^. kg^−1. min⁻¹, respectively. Total time spent in sedentary, light, moderate and vigorous physical activities per day was 406±63 min (50% of total monitored time), 244±56 min (30%), 75±18 min (9%) and 82±30 min (10%). Average total daily distance travelled to and from school was 7.5±3.0 km (0.8–13.4 km). Mean daily energy expenditure, activity-induced energy expenditure and physical activity level was 12.2±3.4 MJ^. day⁻¹, 5.4±3.0 MJ^. day⁻¹ and 2.2±0.6. 70.6% of the variation in was explained by sex (partial R² = 54.7%) and body mass index (partial R² = 15.9%). Energy expenditure and physical activity variables did not predict variation in once sex had been accounted for. The highly active and energy-demanding lifestyle of rural Kenyan adolescents may account for their exceptional aerobic fitness and collectively prime them for later training and athletic success.     

New insights into the northward migration route of gray whales between Vancouver Island,British Columbia,and southeastern Alaska

The route taken by northward migrating gray whales during spring between Vancouver Island and southeastern Alaska, a distance of about 575 km, has long been uncertain. It is generally believed that the whales closely follow the western, outer coastline of Haida Gwaii (formerly the Queen Charlotte Islands), an archipelago lying between Vancouver Island and southeastern Alaska, consistent with their pattern of migrating close to shore over the majority of their northward migratory corridor. By tracking satellite‐tagged individuals and surveying whales from shore bases, we provide evidence that this is not the primary migratory corridor, but instead that most whales migrate through Hecate Strait and Dixon Entrance, broad waterways that lie to the east and north of Haida Gwaii. By using this route, northbound gray whales potentially face a wider range of industrial activities and developments than they would by migrating along the outer coast.     

Running demands and activity profile of men’s rugby sevens: a tournament scenario

This study profiled the changes in running performances and collisions within a Rugby sevens tournament. Sixteen male players were equipped with global positioning system units while competing at the 2015 and 2016 Asia Rugby Sevens series held in Colombo and Hong Kong, respectively. Both tournaments consisted of 4 matches each, and were played over 2 days (i.e., 2 matches/day). Total distance (TD) covered increased in match 3 compared with matches 1 (19 ± 19%; p < 0.001) and 2 (16 ± 11%; p = 0.001), whilst a decrease in TD in match 4 compared with match 3 (8 ± 9%; p = 0.019) was observed. Distances covered within 6.1–12 km·h^-1 and 12.1–14 km·h^-1 speed bands were generally higher in matches 3 and/or 4 when compared with match 1 and/or 2 (p < 0.05). Frequency of entries into 14.1–18 km·h^-1 speed zone was decreased in match 4 compared with match 3 (45 ± 41%; p = 0.009), whilst incidences of heavy, very heavy and severe collisions were generally higher in matches 3 or 4 compared with matches 1 or 2 (p < 0.05). In conclusion, while some decrements in the final match were evident, running performance were generally maintained throughout despite the competitive and congested nature of Rugby Sevens tournaments     

Economical Speed and Energetically Optimal Transition Speed Evaluated by Gross and Net Oxygen Cost of Transport at Different Gradients

The oxygen cost of transport per unit distance (CoT; mL·kg^-1·km^-1) shows a U-shaped curve as a function of walking speed (v), which includes a particular walking speed minimizing the CoT, so called economical speed (ES). The CoT-v relationship in running is approximately linear. These distinctive walking and running CoT-v relationships give an intersection between U-shaped and linear CoT relationships, termed the energetically optimal transition speed (EOTS). This study investigated the effects of subtracting the standing oxygen cost for calculating the CoT and its relevant effects on the ES and EOTS at the level and gradient slopes (±5%) in eleven male trained athletes. The percent effects of subtracting the standing oxygen cost (4.8 ± 0.4 mL·kg^-1·min^-1) on the CoT were significantly greater as the walking speed was slower, but it was not significant at faster running speeds over 9.4 km·h^-1. The percent effect was significantly dependent on the gradient (downhill > level > uphill, P < 0.001). The net ES (level 4.09 ± 0.31, uphill 4.22 ± 0.37, and downhill 4.16 ± 0.44 km·h^-1) was approximately 20% slower than the gross ES (level 5.15 ± 0.18, uphill 5.27 ± 0.20, and downhill 5.37 ± 0.22 km·h^-1, P < 0.001). Both net and gross ES were not significantly dependent on the gradient. In contrast, the gross EOTS was slower than the net EOTS at the level (7.49 ± 0.32 vs. 7.63 ± 0.36 km·h^-1, P = 0.003) and downhill gradients (7.78 ± 0.33 vs. 8.01 ± 0.41 km·h^-1, P < 0.001), but not at the uphill gradient (7.55 ± 0.37 vs. 7.63 ± 0.51 km·h^-1, P = 0.080). Note that those percent differences were less than 2.9%. Given these results, a subtraction of the standing oxygen cost should be carefully considered depending on the purpose of each study.