Body oxygen stores, aerobic dive limits, and the diving abilities of juvenile and adult muskrats (Ondatra zibethicus)

Intraspecific variability in body oxygen reserves, muscle buffering capacity, diving metabolic rate, and diving behavior were examined in recently captured juvenile and adult muskrats. Allometric scaling exponents for lung (b=1.04), blood (b=0.91), and total body oxygen storage capacity (b=1.09) did not differ from unity. The concentration of skeletal muscle myoglobin scaled positively with mass in 254-600-g juveniles (b=1.63) but was mass-independent in larger individuals. Scaling exponents for diving metabolic rate and calculated aerobic dive limit (ADL) were 0.74 and 0.37, respectively. Contrary to allometric predictions, we found no evidence that the diving abilities of muskrats increased with age or body size. Juveniles aged 1-2 mo exhibited similar dive times but dove more frequently than summer-caught adults. Average and cumulative dive times and dive&rcolon;surface ratios were highest for fall- and winter-caught muskrats. Total body oxygen reserves were greatest in winter, mainly due to an increase in blood oxygen storage capacity. The buffering capacity of the hind limb swimming muscles also was highest in winter-caught animals. Several behavioral indicators of dive performance, including average and maximum duration of voluntary dives, varied positively with blood hemoglobin and muscle myoglobin concentration of muskrats. However, none of the behavioral measures were strongly correlated with the total body oxygen reserves or ADLs derived for these same individuals.     

Regional heterothermy and conservation of core temperature in emperor penguins diving under sea ice

Temperatures were recorded at several body sites in emperor penguins (Aptenodytes forsteri) diving at an isolated dive hole in order to document temperature profiles during diving and to evaluate the role of hypothermia in this well-studied model of penguin diving physiology. Grand mean temperatures (+/-S.E.) in central body sites during dives were: stomach: 37.1+/-0.2 degrees C (n=101 dives in five birds), pectoral muscle: 37.8+/-0.1 degrees C (n=71 dives in three birds) and axillary/brachial veins: 37.9+/-0.1 degrees C (n=97 dives in three birds). Mean diving temperature and duration correlated negatively at only one site in one bird (femoral vein, r=-0.59, P<0.05; range <1 degrees C). In contrast, grand mean temperatures in the wing vein, foot vein and lumbar subcutaneous tissue during dives were 7.6+/-0.7 degrees C (n=157 dives in three birds), 20.2+/-1.2 degrees C (n=69 in three birds) and 35.2+/-0.2 degrees C (n=261 in six birds), respectively. Mean limb temperature during dives negatively correlated with diving duration in all six birds (r=-0.29 to -0.60, P<0.05). In two of six birds, mean diving subcutaneous temperature negatively correlated with diving duration (r=-0.49 and -0.78, P<0.05). Sub-feather temperatures decreased from 31 to 35 degrees C during rest periods to a grand mean of 15.0+/-0.7 degrees C during 68 dives of three birds; mean diving temperature and duration correlated negatively in one bird (r=-0.42, P<0.05). In general, pectoral, deep venous and even stomach temperatures during diving reflected previously measured vena caval temperatures of 37-39 degrees C more closely than the anterior abdominal temperatures (19-30 degrees C) recently recorded in diving emperors. Although prey ingestion can result in cooling in the stomach, these findings and the lack of negative correlations between internal temperatures and diving duration do not support a role for hypothermia-induced metabolic suppression of the abdominal organs as a mechanism of extension of aerobic dive time in emperor penguins diving at the isolated dive hole. Such high temperatures within the body and the observed decreases in limb, anterior abdomen, subcutaneous and sub-feather temperatures are consistent with preservation of core temperature and cooling of an outer body shell secondary to peripheral vasoconstriction, decreased insulation of the feather layer, and conductive/convective heat loss to the water environment during the diving of these emperor penguins.     

Seasonal changes in the oxygen storage capacity and aerobic dive limits of the muskrat (Ondatra zibethicus)

Summary The oxygen storage capacity and partitioning of body oxygen reserves were compared in summer-and winter-acclimatized muskrats (Ondatra zibethicus). Blood volume, blood oxygen capacity, and skeletal muscle myoglobin content were higher in December than in July (P<0.02). Total lung capacity increased only slightly in winter (P>0.05). The oxygen storage capacity of a diving muskrat was calculated at 25.2 ml O₂ STPD · kg^-1 in July, compared to 35.7 ml O₂ STPD · kg^-1 in December. Blood comprised the major storage compartment in both seasons, accounting for 57% and 65% of the total oxygen stores in summer and winter, respectively. Based on available oxygen stores and previous estimates of the cost of diving, the aerobic dive limit (ADL) increased from 40.9 s in July to 57.9 s in December. Concurrent behavioral studies suggested that most voluntary diving by muskrats is aerobic. However, the proportion of dives exceeding the calculated ADL of these animals was shown to vary with the context of the dive. Only 3.5% of all dives initiated by muskrats floating in the water exceeded their estimated ADL. Provision of a dry resting site and access to a submerged food source increased this proportion to 18–61%, depending on the underwater distance that foraging muskrats were required to swim. Serial dives exceeding the estimated ADL were not accompanied by extended postdive recovery periods.Abbreviations ADL acrobic dive limit - Hb hemoglobin - Hct hematocrit - Mb myoglobin - PaO₂ arterial O₂ tension - STPD standard temperature and pressure, dry     

Temperature regulation in emperor penguins foraging under sea ice

Inferior vena caval (IVC) and anterior abdominal (AA) temperatures were recorded in seven emperor penguins (Aptenodytes forsteri) foraging under sea ice in order to evaluate the hypothesis that hypothermia-induced metabolic suppression might extend aerobic diving time. Diving durations ranged from 1 to 12.5 min, with 39% of dives greater than the measured aerobic dive limit of 5.6 min. Anterior abdominal temperature decreased progressively throughout dives, and partially returned to pre-dive values during surface intervals. The lowest AA temperature was 19 degrees C. However, mean AA temperatures during dives did not correlate with diving durations. In six of seven penguins, only minor fluctuations in IVC temperatures occurred during diving. These changes were often elevations in temperature. In the one exception, although IVC temperatures decreased, the reductions were less than those in the anterior abdomen and did not correlate with diving durations. Because of these findings, we consider it unlikely that regional hypothermia in emperor penguins leads to a significant reduction in oxygen consumption of the major organs within the abdominal core. Rather, temperature profiles during dives are consistent with a model of regional heterothermy with conservation of core temperature, peripheral vasoconstriction, and cooling of an outer body shell.     

Recording the free-living behaviour of small-bodied, shallow-diving animals with data loggers

1. Time-depth data recorders (TDRs) have been widely used to explore the behaviour of relatively large, deep divers. However, little is known about the dive behaviour of small, shallow divers such as semi-aquatic mammals. 2. We used high-resolution TDRs to record the diving behaviour of American mink Mustela vison (weight of individuals 580-1275 g) in rivers in Oxfordshire (UK) between December 2005 and March 2006. 3. Dives to > 0.2 m were measured in all individuals (n = 6). Modal dive depth and duration were 0.3 m and 10 s, respectively, although dives up to 3 m and 60 s in duration were recorded. Dive duration increased with dive depth. 4. Temperature data recorded by TDRs covaried with diving behaviour: they were relatively cold (modal temperature 4-6 degrees C across individuals) when mink were diving and relatively warm (modal temperature 24-36 degrees C across individuals) when mink were not diving. 5. Individuals differed hugely in their use of rivers, reflecting foraging plasticity across both terrestrial and aquatic environments. For some individuals there was < 1 dive per day while for others there was > 100 dives per day. 6. We have shown it is now possible to record the diving behaviour of small free-living animals that only dive a few tens of centimetres, opening up the way for a new range of TDR studies on shallow diving species.     

Diving heart rate development in postnatal harbour seals, Phoca vitulina

Harbour seals, Phoca vitulina, dive from birth, providing a means of mapping the development of the diving response, and so our objective was to investigate the postpartum development of diving bradycardia. The study was conducted May-July 2000 and 2001 in the St. Lawrence River Estuary (48 degrees 41'N, 68 degrees 01'W). Both depth and heart rate (HR) were remotely recorded during 86,931 dives (ages 2-42 d, n = 15) and only depth for an additional 20,300 dives (combined data covered newborn to 60 d, n = 20). The mean dive depth and mean dive durations were conservative during nursing (2.1 +/- 0.1 m and 0.57 +/- 0.01 min, range = 0-30.9 m and 0-5.9 min, respectively). The HR of neonatal pups during submersion was bimodal, but as days passed, the milder of the two diving HRs disappeared from their diving HR record. By 15 d of age, most of the dive time was spent at the lower diving bradycardia rate. Additionally, this study shows that pups are born with the ability to maintain the lower, more fully developed dive bradycardia during focused diving but do not do so during shorter routine dives.     

Dive bout organization in the Chinstrap penguin at seal island, antarctica

Diving behavior of 2 breeding Chinstrap penguins (Pygoscelis antarctica) was studied focusing first and primarily on dive bouts rather than dives themselves. Analysis of dive bout organization revealed (1) though there are differences between solitary dives and dive bouts in dive duration and dive depth, the first dives of dive bouts do not differ from solitary dives in the dive parameters, (2) mean dive duration during bout correlates positively to both mean dive depth during bout and mean surface interval during bout, while number of dives during bout negatively correlates to both cost (consumed energy) and duration of a dive cycle during bout. These findings suggest the following possibilities on foraging behavior of penguins: (1) their decision to repeat diving depends on the result of the first dive at a site, and the first dives of bouts would tend to be searching or evaluating dives though they would be also successful foraging dives, (2) they repeat diving at a foraging patch until foraging efficiency decrease to a threshold of diminishing returns.     

Diving pattern and performance in relation to foraging ecology in the gentoo penguin, Pygoscelis papua

We present data on diving pattern and performance (dive depth, duration, frequency and organization during the foraging trip) in gentoo penguins Pygoscelis papua , obtained using time-depth recorders ( n = 9 birds, 99 foraging trips). These data are used to estimate various parameters of foraging activity, e.g. foraging range, prey capture rates, and are compared in relation to breeding chronology. Foraging trip duration was 6 h and 10 h, and trip frequency 1.0/day and 0.96/day, during the brooding and creche periods, respectively. Birds spent on average 52%of each foraging trip diving. Dive depth and duration were highly bimodal: shallow dives (< 21 m) averaged 4 m and 0.23 min, and deep dives (> 30 m) 80 m and 2.5 min, respectively. Birds spent on average 71%and 25%of total diving time in deep and shallow dives, respectively. For deep dives, dive duration exceeded the subsequent surface interval, but shallow dives were followed by surface intervals 2–3 times dive duration. We suggest that most shallow dives are searching/exploratory dives and most deep dives are feeding dives. Deep dives showed clear diel patterns averaging 40 m at dawn and dusk and 80–90 m at midday. Estimated foraging ranges were 2.3 km and 4.1 km during the brood and creche period, respectively. Foraging trip duration increased by 4 h between the brood and creche periods but total time spent in deep dives (i.e. time spent feeding) was the same (3 h). Of 99 foraging trips, 56%consisted of only one dive bout and 44%of 2–4 bouts delimited by extended surface intervals > 10 min. We suggest that this pattern of diving activity reflects variation in spatial distribution of prey rather than the effect of physiological constraints on diving ability.     

Hemoglobin concentrations and blood gas tensions of free-diving Weddell seals

Arterial blood gas tensions, pH, and hemoglobin concentrations were measured in four free-diving Weddell seals Leptonychotes weddelli. A microprocessor-controlled sampling system enabled us to obtain 24 single and 31 serial aortic blood samples. The arterial O2 tension (PaO2) at rest [78 +/- 13 (SD) Torr] increased with diving compression to a maximum measured value of 232 Torr and then rapidly decreased to 25-35 Torr. The lowest diving PaO2 we measured was 18 Torr just before the seal surfaced from a 27-min dive. A consistent increase of arterial hemoglobin concentrations from 15.1 +/- 1.10 to 22.4 +/- 1.41 g/100 ml (dives less than 17 min) and to 25.4 +/- 0.79 g/100 ml (dives greater than 17 min) occurred during each dive. We suggest that an extension of the sympathetic outflow of the diving reflex possibly caused profound contraction of the Weddell seal's very large spleen (0.89% of body wt at autopsy), although we have no direct evidence. This contraction may have injected large quantities of red blood cells (2/3 of the total) into the seal's central circulation during diving and allowed arterial O2 content to remain constant for the first 15-18 min of long dives. The increase of arterial CO2 tensions during the dive and the compression increase of arterial N2 tensions were also moderated by injecting red blood cells sequestered at ambient pressure. After each dive circulating red blood cells are oxygenated and rapidly sequestered, possibly in the spleen during the first 15 min of recovery.     

Diving in shallow water: the foraging ecology of darters (Aves: Anhingidae)

Diving birds have to overcome buoyancy, especially when diving in shallow water. Darters and anhingas (Anhingidae) are specialist shallow-water divers, with adaptations for reducing their buoyancy. Compared to closely-related cormorants (Phalacrocoracidae), darters have fully wettable plumage, smaller air sacs and denser bones. A previous study of darter diving behaviour reported no relationship between dive duration and water depth, contrary to optimal dive models. In this study I provide more extensive observations of African darters Anhinga melanogaster rufa diving in water<5 m deep at two sites. Dive duration increases with water depth at both sites, but the relationship is weak. Dives were longer than dives by cormorants in water of similar depth (max 108 s in water 2.5 m deep), with dives of up to 68 s observed in water<0.5 m deep. Initial dives in a bout were shorter than expected, possibly because their plumage was not fully saturated. Dive efficiency (dive:rest ratio) was 5–6, greater than cormorants (2.7±0.4 for 18 species) and other families of diving birds (average 0.2–4.3). Post-dive recovery periods increased with dive duration, but only slowly, resulting in a strong increase in efficiency with dive duration. All dives are likely to fall within the theoretical anaerobic dive limit. Foraging bouts were short (17.8±4.3 min) compared to cormorants, with birds spending 80±5% of time underwater. Darters take advantage of their low buoyancy to forage efficiently in shallow water, and their slow, stealthy dives are qualitatively different from those of other diving birds. However, they are forced to limit the duration of foraging bouts by increased thermoregulatory costs associated with wettable plumage.     

Diving behaviour of guillemot Uria aalge, puffin Fratercula arctica and razorbill Alca tor da as shown by radio-telemetry

Information on dive and pause times and the numbers of dives in a sequence were obtained for six guillemots and single razorbill and puffin. There were marked differences in diving performance between the species with the order of ranking, in descending order of dive and pause duration, being guillemot, razorbill and puffin. For guillemots, 80% of dives were of 20–119 sec duration and 80% of pauses were 0–59 sec; the maximum dive lasted 202 sec. Puffin dives and pauses were much shorter, with 81% of dives lasting 0–39 sec and 95% of pauses being less than 20 sec, the longest dive was 115 sec. Comparisons of diving sequences made by the same individual indicated some flexibility in all aspects of the sequence but there were broad interspecific differences in the organization of the sequence. The puffin generally made a large number of relatively short dives separated by very short pauses which resulted in a high diving rate (1–5 dives/min) and the bird spending 78% of its time underwater. In contrast, guillemots had much shorter sequences with a few long dives and pauses and lower rates of diving (0–5-0-6 dives/min) and proportions of time underwater (61–65%). Guillemots and puffins may forage at different depths and have different foraging strategies.     

Diving and haulout behavior of crabeater seals in the Weddell Sea,Antarctica, during March 1986

Summary Time-depth recorders were used to study the diving and haulout behavior of six crabeater seals in the marginal. ice edge zone of the Weddell Sea during March 1986. Haulout patterns revealed the seals' clear preference for diving during darkness and hauling out onto sea ice during daylight. Seals did not necessarily haul out every day; individual seals hauled out on 80–100% of days during the study period. Four general dive types were identified: 1) traveling dives, 2) foraging dives, 3) crepuscular foraging dives, and 4) exploratory dives. Nearly continual diving occurred for extended periods (about 16 h) nightly, with one individual diving up to 44 h without interruption. Foraging dives occurring during crepuscular periods were deeper than those made during the darkest hours. The authors suggest that the distinct diel pattern of dive timing and depth may be related to possible predator avoidance behavior by the seals' principal prey, Antarctic Krill.     

Diving through the thermal window: implications for a warming world

Population decline and a shift in the geographical distribution of some ectothermic animals have been attributed to climatic warming. Here, we show that rises in water temperature of a few degrees, while within the thermal window for locomotor performance, may be detrimental to diving behaviour in air-breathing ectotherms (turtles, crocodilians, marine iguanas, amphibians, snakes and lizards). Submergence times and internal and external body temperature were remotely recorded from freshwater crocodiles (Crocodylus johnstoni) while they free-ranged throughout their natural habitat in summer and winter. During summer, the crocodiles'' mean body temperature was 5.2 ± 0.1°C higher than in winter and the largest proportion of total dive time was composed of dive durations approximately 15 min less than in winter. Diving beyond 40 min during summer required the crocodiles to exponentially increase the time they spent on the surface after the dive, presumably to clear anaerobic debt. The relationship was not as significant in winter, even though a greater proportion of dives were of a longer duration, suggesting that diving lactate threshold (DLT) was reduced in summer compared with winter. Additional evidence for a reduced DLT in summer was derived from the stronger influence body mass exerted upon dive duration, compared to winter. The results demonstrate that the higher summer body temperature increased oxygen demand during the dive, implying that thermal acclimatization of the diving metabolic rate was inadequate. If the study findings are common among air-breathing diving ectotherms, then long-term warming of the aquatic environment may be detrimental to behavioural function and survivorship.     

Diving and haul-out patterns of walruses Odobenus rosmarus on Svalbard

Nine male walruses were equipped with dive recording devices in Svalbard to investigate walrus diving and haul-out behaviour in late summer. Dive information on 6,018 dives was collected by 3 satellite linked dive recorders. Additional dive information on 7,769 dives was obtained from 3 time depth recorders. The deepest dive recorded was 67 m, but mean depth of foraging dives was 22.5 m. The longest-lasting dive recorded was 24 min, but mean duration of foraging dives was 6 min. The walruses, on average, spent 56 h in the water followed by 20 h hauled out on land.     

Diving behaviour and diet of the blue-eyed shag at South Georgia

Summary This paper describes a concurrent investigation of individual variation in diet, diving patterns and performance of blue-eyed shags Phalacrocorax atriceps breeding at South Georgia. Within one day individual shags exhibited one of three foraging strategies: short diving (4 birds, all dives 120 s) and mixed diving (15 birds, predominantly long but with a few short dives). The mean number of dives per day was significantly higher in shags that only made short dives (mean=172.0, SE=43.2) than birds with a mixed diving strategy (mean=40.5, SE=4.7) and birds that made only long dives (mean=30.8, SE=1.8). Diet was assessed using hard remains recovered from pellets regurgitated by the shags. Small nototheniid fish (c. 10 kJ per item) were by far the commonest prey but most pellets contained additional items. The frequency of pellets with additional items of higher energy value than nototheniid fish (10.c. 900 kJ per item), lower energy value (>1–10 kJ per item) and both higher and lower energy items was strikingly similar to the frequency of shags making long, short and both long and short dives respectively. Predicted aerobic dive limits suggested that during long dives, blue-eyed shags were probably sustained by anaerobic metabolism. Models of prey capture rates demonstrated that for both long and short diving, many items must be caught per dive when birds are feeding on prey at the lower end of the energy range. Predicted capture rates for the commonest recorded prey (small fish) differ markedly between the two diving strategies.     

Role of adenosine in the hypoxia-induced hypothermia of toads

The concept that hypoxia elicits a drop in body temperature (T(b)) in a wide variety of animals is not new, but the mechanisms remain unclear. We tested the hypothesis that adenosine mediates hypoxia-induced hypothermia in toads. Measurements of selected T(b) were performed using a thermal gradient. Animals were injected (into the lymph sac or intracerebroventricularly) with aminophylline (an adenosine receptor antagonist) followed by an 11-h period of hypoxia (7% O(2)) or normoxia exposure. Control animals received saline injections. Hypoxia elicited a drop in T(b) from 24.8 +/- 0.3 to 19. 5 +/- 1.1 degrees C (P < 0.05). Systemically applied aminophylline (25 mg/kg) did not change T(b) during normoxia, indicating that adenosine does not alter normal thermoregulatory function. However, aminophylline (25 mg/kg) significantly blunted hypoxia-induced hypothermia (P < 0.05). To assess the role of central thermoregulatory mechanisms, a smaller dose of aminophylline (0.25 mg/kg), which did not alter hypoxia-induced hypothermia systemically, was injected into the fourth cerebral ventricle. Intracerebroventricular injection of aminophylline (0.25 mg/kg) caused no significant change in T(b) under normoxia, but it abolished hypoxia-induced hypothermia. The present data indicate that adenosine is a central and possibly peripheral mediator of hypoxia-induced hypothermia.     

The influence of oxygen and carbon dioxide on diving behaviour of tufted ducks, Aythya fuligula

While optimal diving models focus on the diver's oxygen (O(2)) stores as the predominant factor influencing diving behaviour, many vertebrate species surface from a dive before these stores are exhausted and may commence another dive well after their O(2) stores have been resaturated. This study investigates the influence of hypoxia and also hypercapnia on the dive cycle of tufted ducks, Aythya fuligula, in terms of surface duration and dive duration. The birds were trained to surface into a respirometer box after each dive to a feeding tray so that rates of O(2) uptake (VO2) and carbon dioxide output (VCO2) at the surface could be measured. Although Vco2 initially lagged behind Vo2, both respiratory gas stores were close to full adjustment after the average surface duration, indicating that they probably had a similar degree of influence on surface duration. Chemoreceptors, which are known to influence diving behaviour, detect changes in O(2) and CO(2) partial pressures in the arterial blood. Thus, the need to restore blood gas levels appears to be a strong stimulus to continue ventilation. Mean surface duration coincided with peak instantaneous respiratory exchange ratio due to predive anticipatory hyperventilation causing hypocapnia. For comparison, the relationship between surface duration and O(2) uptake in reanalysed data for two grey seals indicated that one animal tended to dive well after fully restocking its O(2) stores, while the other dived at the point of full restocking. More CO(2) is exchanged than O(2) in tufted ducks during the last few breaths before the first dive of a bout, serving to reduce CO(2) stores and suggesting that hypercapnia rather than hypoxia is more often the limiting factor on asphyxia tolerance during dives. Indeed, according to calculations of O(2) stores and O(2) consumption rates over modal diving durations, a lack of O(2) does not seem to be associated with the termination of a dive in tufted ducks. However, factors other than CO(2) are also likely to be important, and perhaps more so, such as food density and rate of food ingestion. Because some predictive success has been demonstrated for optimal diving models, they should continue to incorporate O(2) stores as a variable, but their validity is likely to be improved by also focusing on CO(2) stores.     

Sensitivity to hypercapnia and elimination of CO2 following diving in Steller sea lions (Eumetopias jubatus)

The diving ability of marine mammals is a function of how they use and store oxygen and the physiological control of ventilation, which is in turn dependent on the accumulation of CO₂. To assess the influence of CO₂ on physiological control of dive behaviour, we tested how increasing levels of inspired CO₂ (hypercarbia) and decreasing inspired O₂ (hypoxia) affected the diving metabolic rate, submergence times, and dive recovery times (time to replenish O₂ stores and eliminate CO₂) of freely diving Steller sea lions. We also measured changes in breathing frequency of diving and non-diving individuals. Our findings show that hypercarbia increased breathing frequency (as low as 2 % CO₂), but did not affect metabolic rate, or the duration of dives or surface intervals (up to 3 % CO₂). Changes in breathing rates indicated respiratory drive was altered by hypercarbia at rest, but blood CO₂ levels remained below the threshold that would alter normal dive behaviour. It took the sea lions longer to remove accumulated CO₂ than it did for them to replenish their O₂ stores following dives (whether breathing ambient air, hypercarbia, or hypoxia). This difference between O₂ and CO₂ recovery times grew with increasing dive durations, increasing hypercarbia, and was greater for bout dives, suggesting there could be a build-up of CO₂ load with repeated dives. Although we saw no evidence of CO₂ limiting dive behaviour, the longer time required to remove CO₂ may eventually exhibit control over the overall time they can spend in apnoea and overall foraging duration.     

Diving pattern and performance in the macaroni penguin Eudptes chrysolophus

The pattern and characteristics of diving in two female macaroni penguins Eudyptes chrysolophus was studied, during the brooding period, using continuous-recording time-depth recorders, for a total of I8 days (15 consecutive days) during which the depth, duration and timing of 4876 dives were recorded. Diving in the first 11 days was exclusively diurnal, averaging 244 dives on trips lasting 12 hours. Near the end of the brooding period trips were longer and included diving at night. About half of all trips (except those involving continuous night-time diving) was spent in diving and dive rate averaged 14–25 dives per hour (42 per hour at night). The duration of day time dives varied between trips, and averaged 1.4–1.7 min, with a subsequent surface interval of 0.5–0.9 min. Dive duration was significantly directly related to depth, the latter accounting for 53% of the variation. The average depths of daytime dives were 20–35 m (maximum depth 11 5 m). Dives at night were shorter (average duration 0.9 min) and much shallower (maximum 11 m); depth accounted for only 6% of the variation in duration. Estimates of potential prey capture rates (3–5 krill per dive; one krill every 17–20 s) are made. Daily weight changes in chicks were directly related to number of dives, but not to foraging trip duration nor time spent diving. Of the other species at the same site which live by diving to catch krill, gentoo penguins forage exclusively diurnally, making longer. deeper dives; Antarctic fur seals, which dive to similar depths as macaroni penguins, do so mainly at night.     

Heart rates and gas exchange in the Amazonian Manatee (Trichechus inunguis) in relation to diving

Unrestrained Amazonian manatees (Trichechus inunguis) maintained a constant heart rate during diving and exhibited a slight tachycardia during breathing. 'Forcing' the manatees to dive caused a marked bradycardia. They exhibited a more pronounced tachycardia during breathing after 'forced' dives and hyperventilated during recovery dives. Manatees are capable of dives exceeding 10 min duration without having to resport to anaerobic metabolism, and even after 10 min dives recover within 3-4 short dives. The ability of manatees to make long dives, in spite of relatively poor O2 stores, is due to their low metabolic rate, while the rapid recovery is aided by their high CO2 stores which minimizes CO2 storage in the body. In manatees the changes in alveolar O2 and CO2 pressure (PAO2 and PACO2) in relation to dive time are slower and more variable than in other marine mammals. The lower rate of change is probably due to the manatees' reduced metabolic rate, while the greater variability is due to their breathing pattern, in which both ventilation and body gas stores influence alveolar gases.