Body mass and composition responses to short-term low energy intake are seasonally dependent in Steller sea lions (Eumetopias jubatus)

Steller sea lions (Eumetopias jubatus) were fed restricted iso-caloric amounts of Pacific herring (Clupea pallasi) or walleye pollock (Theragra chalcogramma) for 8–9 days, four times over the course of a year to investigate effects of season and prey composition on sea lion physiology. At these levels, the sea lions lost body mass at a significantly higher rate during winter (1.6 ± 0.14 kg day⁻¹), and at a lower rate during summer (1.2 ± 0.32 kg day⁻¹). Decreases in body fat mass and standard metabolic rates during the trials were similar throughout the seasons and for both diet types. The majority of the body mass that was lost when eating pollock derived from decreases in lipid mass, while a greater proportion of the mass lost when eating herring derived from decreases in lean tissue, except in the summer when the pattern was reversed. Metabolic depression was not observed during all trials despite the constant loss of body mass. Our study supports the hypothesis that restricted energy intake may be more critical to Steller sea lions in the winter months, and that the type of prey consumed (e.g., herring or pollock) may have seasonally specific effects on body mass and composition.     

No Evidence of Metabolic Depression in Western Alaskan Juvenile Steller Sea Lions (Eumetopias jubatus)

Steller sea lion (Eumetopias jubatus) populations have undergone precipitous declines through their western Alaskan range over the last four decades with the leading hypothesis to explain this decline centering around changing prey quality, quantity, or availability for this species (i.e., nutritional stress hypothesis). Under chronic conditions of reduced food intake sea lions would conserve energy by limiting energy expenditures through lowering of metabolic rate known as metabolic depression. To examine the potential for nutritional stress, resting metabolic rate (RMR) and body composition were measured in free-ranging juvenile Steller sea lions (N = 91) at three distinct geographical locations (Southeast Alaska, Prince William Sound, Central Aleutian Islands) using open-flow respirometry and deuterium isotope dilution, respectively. Average sea lion RMR ranged from 6.7 to 36.2 MJ d⁻¹ and was influenced by body mass, total body lipid, and to a lesser extent, ambient air temperature and age. Sea lion pups captured in the Aleutian Islands (region of decline) had significantly greater body mass and total body lipid stores when compared to pups from Prince William Sound (region of decline) and Southeast Alaska (stable region). Along with evidence of robust body condition in Aleutian Island pups, no definitive differences were detected in RMR between sea lions sampled between eastern and western populations that could not be accounted for by higher percent total body lipid content, suggesting that that at the time of this study, Steller sea lions were not experiencing metabolic depression in the locations studied.     

Changes in metabolism in response to fasting and food restriction in the Steller sea lion (Eumetopias jubatus)

Many animals lower their resting metabolism (metabolic depression) when fasting or consuming inadequate food. We sought to document this response by subjecting five Steller sea lions to periods of: (1) complete fasting; or (2) restricting them to 50% of their normal herring diet. The sea lions lost an average of 1.5% of their initial body mass per day (2.30 kg/d) during the 9-14-day fast, and their resting metabolic rates decreased 31%, which is typical of a "fasting response". However, metabolic depression did not occur during the 28-day food restriction trials, despite the loss of 0.30% of body mass per day (0.42 kg/d). This difference in response suggests that undernutrition caused by reduced food intake may stimulate a "hunger response", which in turn might lead to increased foraging effort. The progressive changes in metabolism we observed during the fasts were related to, but were not directly caused by, changes in body mass from control levels. Combining these results with data collected from experiments when Steller sea lions were losing mass on low energy squid and pollock diets reveals a strong relationship between relative changes in body mass and relative changes in resting metabolism across experimental conditions. While metabolic depression caused by fasting or consuming large amounts of low energy food reduced the direct costs from resting metabolism, it was insufficient to completely overcome the incurred energy deficit.     

Steller sea lions Eumetopias jubatus and nutritional stress: evidence from captive studies

• 1 Numbers of Steller sea lions Eumetopias jubatus in the North Pacific have declined. According to the nutritional stress hypothesis, this decline is due to reduced food availability. Data from studies conducted on pinnipeds in the laboratory are used here to test if the nutritional stress hypothesis can explain the decline of Steller sea lions.
• 2 Overall, there is strong evidence for biologically meaningful differences in the nutritional quality of major prey species. Steller sea lions can partly compensate for low‐quality prey by increasing their food consumption.
• 3 There appear to be no detrimental effects of low‐lipid prey on sea lion growth or body composition when sea lions can consume sufficient quantities of prey. However, the ability to increase consumption is physiologically limited, particularly in young animals. Overall, it is more difficult to maintain energy intake on a diet of low‐quality prey than on a normal diet.
• 4 Under conditions of inadequate food intake (either due to decreased prey availability or quality, or increased energy requirements) the overall impacts of nutritional stress are complex, and are dependent upon season, prey quality, age and the duration and intensity of the nutritional stress event.
• 5 Studies on pinnipeds in the laboratory have been instrumental in identifying the conditions under which changes in sea lion prey can result in nutritional stress and the nature of the physiological impacts of nutritional stress events.

     

Seasonal variability in otariid energetics: implications for the effects of predators on localized prey resources

Otariids, like other wild mammals, contend with a wide variety of energetic demands across seasons. However, due to the cryptic behaviors of this marine group, few studies have been able to examine longitudinal energetic costs or the potential impact of these costs on seasonal or annual prey requirements. Here we evaluated the changes in energy demand and intake of female California sea lions (Zalophus californianus) during reproductive (n=2 sea lions) and nonreproductive (n=3) periods. Monthly measurements included resting metabolic rate, blood hormone levels, body condition (blubber thickness and body mass), and caloric intake for adult sea lions throughout molting, late pregnancy, lactation, and postweaning. We found that maintenance energy demands decreased from 32.0 to 23.1 MJ d(-1) before pupping, remaining stable at 19.4+/-0.6 MJ d(-1) during lactation and postweaning. Energy intake rates to meet these demands showed marked changes with activity level and the reproductive cycle, reaching a peak intake of 3.6 times baseline levels during lactation. Translating this into prey demands, we find that 20,000 reproductively active females on San Nicolas Island rookeries would maximally require 4,950 metric tons of Pacific whiting during a month of the breeding season. This localized impact is reduced significantly with postbreeding dispersal and demonstrates the importance of considering spatial and temporal factors driving the energetic requirements of predators when designing marine protected areas.     

Metabolic effects of low-energy diet on steller sea lions, Eumetopias jubatus

Diets of six Steller sea lions (Eumetopias jubatus) were switched between a high (herring) and a low (squid) energy density food for 14 d to determine the effects on ingested prey mass, body mass, resting metabolic rate, and the heat increment of feeding. Body mass was measured daily, and resting metabolism was measured weekly by gas respirometry. Ingested food mass did not differ significantly between the squid diet and the control or the recovery herring diet periods. As a result of differences in energy density, gross energy intake was significantly lower during the squid diet phase than during either the control or recovery periods. As a result, sea lions lost an average of 1.1 kg/d, totaling 12.2% of their initial body mass by the end of the experimental period. The heat increment of feeding for a 4-kg squid meal was significantly lower than for a similarly sized meal of herring. Decreases in both absolute (24.0 to 18.0 MJ/d, -24%) and mass-corrected (903 to 697 kJ/d/kg0.67, -20%) metabolism were observed by the end of the squid feedings. This study suggests that sea lions can depress their resting metabolism in response to decreases in energy intake or body mass, regardless of satiation level.     

Impact of changing diet regimes on Steller sea lion body condition

A leading theory for the cause of the decline of Steller sea lions is nutritional stress, which led to chronic high juvenile mortality and possibly episodic adult mortality. Nutritional stress may have resulted from either poor quality or low abundance of prey. The objective of this study was to determine whether we could predict shifts in body condition (i.e., body mass or body fat content) over different seasons associated with a change in diet (i.e., toward lower quality prey). Captive Steller sea lions (n= 3) were fed three different diet regimes, where Diet 1 approximated the diet in the Kodiak area in the 1970s prior to the documented decline in that area, Diet 2 approximated the species composition in the Kodiak area after the decline had begun, and Diet 3 approximated the diet in southeast Alaska where the Steller sea lion population has been increasing for over 25 yr. All the animals used in this study were still growing and gained mass regardless of diet. Body fat (%) varied between 13% and 28%, but was not consistently high or low for any diet regime or season. Mean intake (in kg) of Diet 2 was significantly greater for all sea lions during all seasons. All animals did, however, tend to gain less body mass on Diets 2 and 3, as well as during the breeding and postbreeding seasons. They also tended to gain more mass during the winter and on Diet 1, though these differences were not statistically significant. Thus, changing seasonal physiology of Steller sea lions appears to have more impact on body condition than quality of prey, provided sufficient quantity of prey is available. Steller sea lions are opportunistic predators and are evidently able to thrive on a variety of prey. Our results indicate that Steller sea lions are capable of compensating for prey of low quality.     

Changes in glucocorticoids, IGF-I and thyroid hormones as indicators of nutritional stress and subsequent refeeding in Steller sea lions (Eumetopias jubatus)

Physiological responses to changes in energy balance are tightly regulated by the endocrine system through glucocorticoids, IGF-I and thyroid hormones. Changes in these hormones were studied in eight captive female Steller sea lions that experienced changes in food intake, body mass, body composition, and blood metabolites during summer and winter. During a period of energy restriction, one group of sea lions was fed reduced amounts of Pacific herring and another was fed an isocaloric diet of walleye pollock, after which both groups returned to their pre-experimental diets of herring. Cortisol was negatively and IGF-I was positively associated with changes in body mass during periods of energy restriction (mass loss associated with increase in cortisol and decrease in IGF-I) and refeeding (body mass maintenance associated with stable hormone concentrations in summer and compensatory growth linked to decrease in cortisol and increase in IGF-I in winter). Cortisol and IGF-I were also correlated with changes in lipid and lean mass, respectively. Consequently, these two hormones likely make adequate biomarkers for nutritional stress in sea lions, and when combined provide indication of the energetic strategy (lipid vs lean mass catabolism) animals adopt to cope with changes in nutrient intake. Unlike type of diet fed to the sea lions, age of the animals also impacted hormonal responses, with younger animals showing more intense hormonal changes to nutritional stress. Thyroid hormones, however, were not linked to any physiological changes observed in this study.     

The decline of Steller sea lions Eumetopias jubatus in Alaska: a review of the nutritional stress hypothesis

1. The decline of Steller sea lions Eumetopias jubatus in the Gulf of Alaska and Aleutian Islands between the late 1970s and 1990s may have been related to reduced availability of suitable prey. Many studies have shown that pinnipeds and other mammals suffering from nutritional stress typically exhibit reduced body size, reduced productivity, high mortality of pups and juveniles, altered blood chemistry and specific behavioural modifications. 2. Morphometric measurements of Steller sea lions through the 1970s and 1980s in Alaska indicate reduced body size. Reduced numbers of pups born and an apparent increase in juvenile mortality rates also appear to be nutritionally based. Blood chemistry analyses have further shown that Steller sea lions in the Gulf of Alaska and Aleutian Islands area exhibited signs of an acute phase reaction, or immune reaction, in response to unidentified physical and/or environmental stress. Behavioural studies during the 1990s have not noted any changes that are indicative of an overall shortage in the quantity of prey available to lactating female sea lions. 3. The data collected in Alaska are consistent with the hypothesis that Steller sea lions in the declining regions were nutritionally compromised because of the relative quality of prey available to them (chronic nutritional stress), rather than because of the overall quantity of fish per se (acute nutritional stress). This is further supported by captive studies that indicate the overall quality of prey that has been available to Steller sea lions in the declining population could compromise the health of Steller sea lions and hinder their recovery.     

The energetic consequences of behavioral variation in a marine carnivore

Intraspecific variability in foraging behavior has been documented across a range of taxonomic groups, yet the energetic consequences of this variation are not well understood for many species. Understanding the effect of behavioral variation on energy expenditure and acquisition is particularly crucial for mammalian carnivores because they have high energy requirements that place considerable pressure on prey populations. To determine the influence of behavior on energy expenditure and balance, we combined simultaneous measurements of at‐sea field metabolic rate (FMR) and foraging behavior in a marine carnivore that exhibits intraspecific behavioral variation, the California sea lion (Zalophus californianus). Sea lions exhibited variability in at‐sea FMR, with some individuals expending energy at a maximum of twice the rate of others. This variation was in part attributable to differences in diving behavior that may have been reflective of diet; however, this was only true for sea lions using a foraging strategy consisting of epipelagic (<200 m within the water column) and benthic dives. In contrast, sea lions that used a deep‐diving foraging strategy all had similar values of at‐sea FMR that were unrelated to diving behavior. Energy intake did not differ between foraging strategies and was unrelated to energy expenditure. Our findings suggest that energy expenditure in California sea lions may be influenced by interactions between diet and oxygen conservation strategies. There were no apparent energetic trade‐offs between foraging strategies, although there was preliminary evidence that foraging strategies may differ in their variability in energy balance. The energetic consequences of behavioral variation may influence the reproductive success of female sea lions and result in differential impacts of individuals on prey populations. These findings highlight the importance of quantifying the relationships between energy expenditure and foraging behavior in other carnivores for studies addressing fundamental and applied physiological and ecological questions.     

The influence of depth on a breath-hold diver: Predicting the diving metabolism of Steller sea lions (Eumetopias jubatus)

Diving animals must endeavor to increase their dive depths and prolong the time they spend exploiting resources at depth. Results from captive and wild studies suggest that many diving animals extend their foraging bouts by decreasing their metabolisms while submerged. We measured metabolic rates of Steller sea lions (Eumetopias jubatus) trained to dive to depth in the open ocean to investigate the relationships between diving behaviour and the energetic costs of diving. We also constructed a general linear model to predict the oxygen consumption of sea lions diving in the wild. The resultant model suggests that swimming distance and depth of dives significantly influence the oxygen consumption of diving Steller sea lions. The predictive power of the model was tested using a cross-validation approach, whereby models reconstructed using data from pairs of sea lions were found to accurately predict the oxygen consumption of the third diving animal. Predicted oxygen consumption during dives to depth ranged from 3.37 L min^− 1 at 10 m, to 1.40 L min^− 1 at 300 m over a standardized swimming distance of 600 m. This equated to an estimated metabolic rate of 97.54 and 40.52 MJ day^− 1, and an estimated daily feeding requirement of 18.92 and 7.96 kg day^− 1 for dives between 10 and 300 m, respectively. The model thereby provides information on the potential energetic consequences that alterations in foraging strategies due to changes in prey availability could have on wild populations of sea lions.     

Dietary restriction causes chronic elevation of corticosterone and enhances stress response in red-legged kittiwake chicks

Release of corticosterone in hungry kittiwake chicks facilitates begging and allows them to restore depleted energy reserves by increasing parental food provisioning. However, in order to avoid detrimental effects of chronic elevation of corticosterone, chicks might suppress adrenocortical activity in response to prolonged food shortages. In this study we examined temporal dynamics of corticosterone release in red-legged kittiwake (Rissa brevirostris) chicks exposed to prolonged restrictions in energy content and/or nutritional quality (low versus high lipid content) of their food. Starting at the age of 15 days, chicks were fed either high- or low-lipid fish at 40%, 65%, and 100% of ad libitum energy intake. Body mass measurements and baseline plasma samples were taken on a weekly basis after beginning of the treatment. After 3 weeks of treatment, chicks were exposed to a standardized acute handling and restraint stress protocol, where in addition to a baseline sample, three plasma samples were taken at intervals up to 50 min. We found that food-restricted chicks had lower body mass, chronically (during 2-3 weeks) elevated baseline and higher acute stress-induced levels of corticosterone compared to chicks fed ad libitum. Low lipid content of food further exacerbated these effects. An increase in baseline levels of corticosterone was observed within a week after energy requirements of food-restricted chicks exceeded their daily energy intake. A tendency for suppression of adrenocortical activity was observed in treatments fed low-lipid diets only at the end of the experiment. We suggest that nest-bound chicks, if food-stressed, might suffer deleterious effects of chronic elevation of corticosterone.     

Comparing total body lipid content of young‐of‐the‐year Steller sea lions among regions of contrasting population trends

Steller sea lion (Eumetopias jubatus) young‐of‐the‐year (YOY) are nutritionally dependent upon their dam through the majority of their first year. Several indices of body condition were measured in YOY 1.5–9 mo of age captured in Southeast Alaska (n = 122), the Gulf of Alaska (n = 182), and the Aleutian Islands (n = 38) to test the hypothesis that nutritional stress impacted the ability of adult female Steller sea lions to adequately nourish their late gestation YOY in the central Aleutian Islands in the early 2000s. Body mass (kg) and percent total body lipid content (%TBL) increased with age in all three regions of Alaska that were sampled (P < 0.05). Young‐of‐the‐year 7–9 mo of age were leaner in Southeast Alaska (27.6% ± 1.0%) and Gulf of Alaska (29.5% ± 0.8%) than in the Aleutian Islands (35.7% ± 1.2%, P < 0.001). Condition indices calculated from morphometric measures did not strongly predict the %TBL measured by isotope dilution. The trend for Aleutian Island YOY to have larger body mass and larger body fat reserves are counter to what would be expected if dams were unable to adequately provision their late lactation YOY due to inadequate food availability in the central Aleutian Islands.     

Hormone changes indicate that winter is a critical period for food shortages in Steller sea lions

Given that many marine mammals display seasonal energetic priorities, it is important to investigate whether the impact of unexpected food restriction differs during the year. Steller sea lions (Eumetopias jubatus) fed restricted diets for up to 9 days during spring, summer, fall, and winter lost an average of 10% of their initial body mass. We tracked changes in the levels of three hormones (cortisol, total thyroxine-TT4, total triiodothyronine-TT3) and one blood metabolite (blood urea nitrogen-BUN) following a food restriction in relation to season, body mass, body composition, and metabolism. Degree of changes in cortisol, TT3, and BUN after food restriction was significantly affected by season. The greatest changes in cortisol (+231%), BUN (+11.4%), TT4 (-23.3%), and TT3 (-35.6%) occurred in the winter (November/December) when rates of body mass loss were also greatest. Changes in cortisol levels were positively related to total body mass loss, while changes in TT3 levels were negatively related. While greater increases in BUN were related to greater rates of mass loss, the use of BUN levels as an indicator of metabolic state is complicated by the type and level of food intake. The observed changes in hormone levels support morphological data suggesting Steller sea lions may be more strongly impacted by short-term, reduced energy intake during winter than at other times of the year.     

Weaning mass affects changes in body composition and food intake in harbour seal pups during the first month of independence

In phocid seals, the transition to nutritional independence is abrupt, with females abandoning their offspring after weaning and returning to sea. We hypothesized that body size at weaning may play an important role in the nature of this transition. We studied the changes in body composition and water flux of newly weaning harbour seals over the first 4-6 wk postweaning. Thirty-three pups were dosed with deuterium oxide to estimate total body water (TBW) and a subset of 24 was dosed twice to estimate changes in body composition and water flux. All pups lost body mass over the study period, but TBW increased during the period of mass loss, indicating continued lean tissue growth. Combined data from this and our early study indicated that heavy (>median mass) pups were relatively fatter (41.0% vs. 37.1%) and had significantly greater total body energy at weaning than did light (< or = median mass) pups. Percentage TBW declined linearly over time in light pups but was constant in heavy pups for the first 19 d postweaning and then declined linearly. Both the temporal pattern and composition of mass loss differed between light and heavy pups. Estimated food intake increased in the second 2 wk of study compared to the first 2 wk, in both light and heavy pups, reflecting increased foraging success but at levels still insufficient to meet daily expenditures of most individuals.     

Beneath the surface: Profiling blubber depth in pinnipeds with infrared imaging

Infrared thermography (IRT) was assessed as a non-invasive tool to evaluate body condition in juvenile female harbor seals (Phoca vitulina), (n=6) and adult female Steller sea lions (Eumetopias jubatus), (n=2). Surface temperature determined by IRT and blubber depth assessed with portable imaging ultrasound were monitored concurrently at eight body sites over the course of a year in long-term captive individuals under controlled conditions. Site-specific differences in surface temperature were noted between winter and summer in both species. Overall, surface temperature was slightly higher and more variable in harbor seals (9.8±0.6 °C) than Steller sea lions (9.1±0.5 °C). Limited site-specific relationships were found between surface temperature and blubber thickness, however, insulation level alone explained a very small portion of the variance. Therefore, while validated IRT data collection can potentially provide valuable information on the health, condition and metabolic state of an animal, it cannot provide a generalized proxy for blubber depth.     

Predator-prey size relationships in an African large-mammal food web

1. Size relationships are central in structuring trophic linkages within food webs, leading to suggestions that the dietary niche of smaller carnivores is nested within that of larger species. However, past analyses have not taken into account the differing selection shown by carnivores for specific size ranges of prey, nor the extent to which the greater carcass mass of larger prey outweighs the greater numerical representation of smaller prey species in the predator diet. Furthermore, the top-down impact that predation has on prey abundance cannot be assessed simply in terms of the number of predator species involved. 2. Records of found carcasses and cause of death assembled over 46 years in the Kruger National Park, South Africa, corrected for under-recording of smaller species, enabled a definitive assessment of size relationships between large mammalian carnivores and their ungulate prey. Five carnivore species were considered, including lion (Panthera leo), leopard (Panthera pardus), cheetah (Acinonyx jubatus), African wild dog (Lycaon pictus) and spotted hyena (Crocuta crocuta), and 22 herbivore prey species larger than 10 kg in adult body mass. 3. These carnivores selectively favoured prey species approximately half to twice their mass, within a total prey size range from an order of magnitude below to an order of magnitude above the body mass of the predator. The three smallest carnivores, i.e. leopard, cheetah and wild dog, showed high similarity in prey species favoured. Despite overlap in prey size range, each carnivore showed a distinct dietary preference. 4. Almost all mortality was through the agency of a predator for ungulate species up to the size of a giraffe (800-1200 kg). Ungulates larger than twice the mass of the predator contributed substantially to the dietary intake of lions, despite the low proportional mortality inflicted by predation on these species. Only for megaherbivores substantially exceeding 1000 kg in adult body mass did predation become a negligible cause of mortality. 5. Hence, the relative size of predators and prey had a pervasive structuring influence on biomass fluxes within this large-mammal food web. Nevertheless, the large carnivore assemblage was dominated overwhelmingly by the largest predator, which contributed the major share of animals killed across a wide size range.     

MASS ESTIMATES USING BODY MORPHOLOGY IN STELLER SEA LIONS

Abstract: Analytical procedures developed from studies on phocids for relating body morphometrics (length and girth) to mass were applied to 390 Steller sea lions from the Gulf of Alaska and the Bering Sea. The equations relating body volume to mass were significant and had very high correlation coefficients for animals from about 60 to over 900 kg with a standard error of ±0.6% in fitted mass. There was a significant difference in these relationships for female sea lions studied in the 1970s and those examined in the 1980s. The latter sea lions were smaller and/or leaner than those collected in the 1970s.     

Milk intake,growth and energy consumption in pups of ice-breeding grey seals (Halichoerus grypus) from the Gulf of St. Lawrence,Canada

In this study we document growth, milk intake and energy consumption in nursing pups of icebreeding grey seals (Halichoerus grypus). Change in body composition of the pups, change in milk composition as lactation progresses, and mass transfer efficiency between nursing mothers and pups are also measured. Mass transfer efficiency between mother-pup pairs (n=8) was 42.5±8.4%. Pups were gaining a daily average of 2.0±0.7 kg (n=12), of which 75% was fat, 3% protein and 22% water. The total water influx was measured to be 43.23±8.07 ml·kg^-1·day^-1. Average CO₂ production was 0.85±0.20 ml·g^-1·h^-1, which corresponds to a field metabolic rate of 0.55±0.13 MJ·kg^-1·day^-1, or 4.5±0.9 times the predicted basal metabolic rate based on body size (Kleiber 1975). Water and fat content in the milk changed dramatically as lacation progressed. At day 2 of nursing, fat and water content were 39.5±1.9% and 47.3±1.5%, respectively, while the corresponding figures for day 15 were 59.6±3.6% fat and 28.4±2.6% water. Protein content of the milk remained relatively stable during the lactation period with a value of 11.0±0.8% at day 2 and 10.4±0.3% at day 15. Pups drank an average of 3.5±0.9 kg of milk daily, corresponding to a milk intake of 1.75 kg per kg body mass gained. The average daily energy intake of pups was 82.58±19.80 MJ, while the energy built up daily in the tissue averaged 61.72±22.22 MJ. Thus, pups assimilated 74.7% of the energy they received via milk into body tissue. The lactation energetics of ice-breeding grey seals is very similar to that of their land-breeding counterparts.Abbreviations bm body mass - BMR basal metabolic rate - FMR field metabolic rate - IU international unit - RQ respiration quotient - HTO tritiated water - HT¹⁸O doubly labeled water - TBW total body water - VHF very high frequency     

The costs of carnivory

Mammalian carnivores fall into two broad dietary groups: smaller carnivores (<20 kg) that feed on very small prey (invertebrates and small vertebrates) and larger carnivores (>20 kg) that specialize in feeding on large vertebrates. We develop a model that predicts the mass-related energy budgets and limits of carnivore size within these groups. We show that the transition from small to large prey can be predicted by the maximization of net energy gain; larger carnivores achieve a higher net gain rate by concentrating on large prey. However, because it requires more energy to pursue and subdue large prey, this leads to a 2-fold step increase in energy expenditure, as well as increased intake. Across all species, energy expenditure and intake both follow a three-fourths scaling with body mass. However, when each dietary group is considered individually they both display a shallower scaling. This suggests that carnivores at the upper limits of each group are constrained by intake and adopt energy conserving strategies to counter this. Given predictions of expenditure and estimates of intake, we predict a maximum carnivore mass of approximately a ton, consistent with the largest extinct species. Our approach provides a framework for understanding carnivore energetics, size, and extinction dynamics.