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Natural history collections provide an unparalleled resource for documenting population responses to past anthropogenic change. However, in many cases, traits measured on specimens may vary temporally in response to a number of different anthropogenic pressures or demographic processes. While teasing apart these different drivers is challenging, approaches that integrate analyses of spatial and temporal series of specimens can provide a robust framework for examining whether traits exhibit common responses to ecological variation in space and time. We applied this approach to analyze bill morphology variation in California Savannah Sparrows (Passerculus sandwichensis). We found that bill surface area increased in birds from higher salinity tidal marshes that are hotter and drier. Only the coastal subspecies, alaudinus, exhibited a significant increase in bill size through time. As with patterns of spatial variation, alaudinus populations occupying higher salinity tidal marshes that have become warmer and drier over the past century exhibited the greatest increases in bill surface area. We also found a significant negative correlation between bill surface area and total evaporative water loss (TEWL) and estimated that observed increases in bill size could result in a reduction of up to 16.2% in daily water losses. Together, these patterns of spatial and temporal variation in bill size were consistent with the hypothesis that larger bills are favored in freshwater‐limited environments as a mechanism of dissipating heat, reducing reliance on evaporative cooling, and increasing water conservation. With museum collections increasingly being leveraged to understand past responses to global change, this work highlights the importance of considering the influence of many different axes of anthropogenic change and of integrating spatial and temporal analyses to better understand the influence of specific human impacts on population change over time.

The present study attempts to assess changes in soccer players’ physical and technical activity profiles due to thermal stress, measured with the Universal Thermal Climate Index (UTCI), in training centres and during matches of the 2018 FIFA World Cup in Russia. The study also verifies the theoretical models of soccer players’ physiological parameters. The study sample consisted of 945 observations of 340 players of national teams taking part in the World Cup in Russia. The measured variables included physical activities: total distance covered, distances covered with an intensity of 20–25 km/h, number of sprints; technical activities: number of shots, number of passes, pass accuracy and physiological indicators: evaporative water loss and heart rate. In addition, the final ranking places of each national team were also used in the study. The UTCI was calculated based on meteorological data recorded at training centres and during matches. The UTCI records were then classified into two ranges: NTS—no thermal stress (UTCI 9–26 °C) and TS—thermal stress (UTCI > 26 °C). Climatic conditions at soccer training centres assessed as involving “no thermal stress” are found to be more beneficial for increasing the total distance covered and the number of sprints performed by players during a match. The theoretical models for determining soccer players’ physiological parameters used in the study reduce the players’ heart rate effort and evaporative water loss, which is in line with findings in studies by other authors. The climatic conditions at soccer training centres and during tournament matches should be taken into account in planning preparations for future World Cup tournaments, especially those in hotter countries.

The Arctic is warming at approximately twice the global rate, with well-documented indirect effects on wildlife. However, few studies have examined the direct effects of warming temperatures on Arctic wildlife, leaving the importance of heat stress unclear. Here, we assessed the direct effects of increasing air temperatures on the physiology of thick-billed murres (Uria lomvia), an Arctic seabird with reported mortalities due to heat stress while nesting on sun-exposed cliffs. We used flow-through respirometry to measure the response of body temperature, resting metabolic rate, evaporative water loss and evaporative cooling efficiency (the ratio of evaporative heat loss to metabolic heat production) in murres while experimentally increasing air temperature. Murres had limited heat tolerance, exhibiting: (1) a low maximum body temperature (43.3°C); (2) a moderate increase in resting metabolic rate relative that within their thermoneutral zone (1.57 times); (3) a small increase in evaporative water loss rate relative that within their thermoneutral zone (1.26 times); and (4) a low maximum evaporative cooling efficiency (0.33). Moreover, evaporative cooling efficiency decreased with increasing air temperature, suggesting murres were producing heat at a faster rate than they were dissipating it. Larger murres also had a higher rate of increase in resting metabolic rate and a lower rate of increase in evaporative water loss than smaller murres; therefore, evaporative cooling efficiency declined with increasing body mass. As a cold-adapted bird, murres' limited heat tolerance likely explains their mortality on warm days. Direct effects of overheating on Arctic wildlife may be an important but under-reported impact of climate change., Summary: Thick-billed murres have limited heat tolerance, displaying low evaporative cooling efficiencies and increases in physiological traits associated with heat stress at low air temperatures relative to desert birds.

Made available in DSpace on 2019-10-04T12:33:36Z (GMT). No. of bitstreams: 0 Previous issue date: 2019-01-01 Serrapilheira Institute Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) Geographical gradients of body size express climate-driven constraints on animals, but whether they exist and what causes them in ectotherms remains contentious. For amphibians, the water conservation hypothesis posits that larger bodies reduce evaporative water loss (EWL) along dehydrating gradients. To address this hypothesis mechanistically, we build on well-established biophysical equations of water exchange in anurans to propose a state-transition model that predicts an increase of either body size or resistance to EWL as alternative specialization along dehydrating gradients. The model predicts that species whose water economy is more sensitive to variation in body size than to variation in resistance to EWL should increase in size in response to increasing potential evapotranspiration (PET). To evaluate the model predictions, we combine physiological measurements of resistance to EWL with geographic data of body size for four different anuran species. Only one species, Dendropsophus minutus, was predicted to exhibit a positive body size-PET relationship. Results were as predicted for all cases, with one species-Boana faber-showing a negative relationship. Based on an empirically verified mathematical model, we show that clines of body size among anurans depend on the current values of those traits and emerge as an advantage for water conservation. Our model offers a mechanistic and compelling explanation for the cause and variation of gradients of body size in anurans. Univ Fed Sergipe, Dept Ecol, BR-49100000 Sao Cristovao, Sergipe, Brazil Univ Sao Paulo, Inst Biosci, Dept Physiol, BR-05508090 Sao Paulo, Brazil Rey Juan Carlos Univ, Dept Biol & Geol, Phys & Inorgan Chem, Madrid 28933, Spain Univ Fed Sao Carlos, Dept Environm Sci, BR-18052780 Sorocaba, SP, Brazil Univ Fed Goias, Dept Ecol, BR-74001970 Goiania, Go, Brazil Sao Paulo State Univ, Dept Zool, BR-13506900 Rio Claro, SP, Brazil Univ Fed Sergipe, Dept Biol, BR-49100000 Sao Cristovao, Sergipe, Brazil Sao Paulo State Univ, Dept Zool, BR-13506900 Rio Claro, SP, Brazil Serrapilheira Institute: G-1709-18372 CNPq: 303180/2016-1 CNPq: 402469/2016-0 CAPES: 88881.157451/2017-01 FAPESP: 10/20061-6 FAPESP: 14/05624-5 FAPESP: 17/10338-0 CNPq: 166109/2015-0 CNPq: 302045/2012-0 CNPq: 306811/2015-4 CNPq: 472397/2013-3 CNPq: 201210767000812 FAPESP: 2012/07356-2 FAPESP: 2013/50714-0 FAPESP: 13/04190-9

The maintenance of optimal body temperatures has profound consequences on all aspects of ectotherms life history, like fitness and performance, and has been the subject of research for decades. In contrast, for reptiles, comparatively less is known on the potential effects of water balance on physiology, ecology and behaviour. In recent years, several recent studies are pointing out the importance of studying thermoregulation and water balance within the same framework. Here, we used a Mediterranean lizard, Psammodromus algirus, to investigate how field body temperature and water loss rates may vary among populations, between sexes and along an altitudinal gradient. We found little variation in field body temperatures among populations, while within each population, field body temperature may change daily, seasonally and differ between sexes. On the contrary, water loss rates decreased with elevation and showed no seasonal trend or difference between sexes. Microclimatic data indicated that highland lizards experienced more fluctuating conditions, a factor that may explain their lower water loss rates. Other factors, like inter-population differences in ectoparasite intensity, might also contribute in explaining the observed patterns. We present here the first data for a lacertid lizard on the inter-population variability and sexual difference in water loss rates and point out that water balance may play a fundamental role in regulating lizard activity during the hottest and driest period of the year., Permits to capture and handle lizards were granted by the government of Madrid region (permit number: 10/165952.9/18). MS was supported by a doctoral grant (SFRH/BD/103549/ 2014) from Fundacao para a Ciencia e a Tecnologia (FCT, Portugal). EC was supported by a postdoctoral grant (SFRH/BPD/109182/2015) from FCT. JM is supported by the MINECO CGL2014-53523-P project. MAC is supported by project PTDC/BIA-CBI/28014/2017 (FCT).

Marsupials have relatively low body temperatures and metabolic rates, and are therefore considered to be maladapted for life in cold habitats such as alpine environments. We compared body temperature, energetics and water loss as a function of ambient temperature for 4 Antechinus species, 2 from alpine habitats and 2 from low altitude habitats. Our results show that body temperature, metabolic rate, evaporative water loss, thermal conductance and relative water economy are markedly influenced by ambient temperature for each species, as expected for endothermic mammals. However, despite some species and individual differences, habitat (alpine vs non-alpine) does not affect any of these physiological variables, which are consistent with those for other marsupials. Our study suggests that at least under the environmental conditions experienced on the Australian continent, life in an alpine habitat does not require major physiological adjustments by small marsupials and that they are physiologically equipped to deal with sub-zero temperatures and winter snow cover.

In terrestrial endotherms, evaporation is a significant mechanism of water loss in hot environments. Although water is passively lost by evaporation, individuals can regulate it at different levels. Inhabiting a relatively stable environment characterized by mild ambient temperature (Ta) and high humidity can ensure a balanced water budget. Many fossorial rodents are well adapted to live in such conditions. In this study, evaporative water loss (EWL) of fossorial rodent species with different degree of adaptations to underground life (from strictly subterranean to those with regular surface activity) was evaluated. By measuring EWL, the specific contribution of either evaporative or non-evaporative components of heat loss can be determined. With the exception of the silvery mole-rat (Heliophobius argenteocinereus), in all tested rodents EWL is relatively stable below and within the thermoneutral zone (TNZ). As Tas increase above TNZ, EWL increases as does total thermal conductance, but conductance increases several times more than EWL. In addition, non-evaporative routes seem to be more important than evaporative heat loss in the analyzed species. No clear pattern of EWL in relation to a species degree of fossoriality or sociality was detected. In this context, atmosphere of burrows could affect EWL, since the high humidity found inside tunnels can establish limits on evaporation to favor water rather than thermal balance. Fil: Luna, Facundo. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina Fil: Šumbera, Radim. University of South Bohemia; República Checa Fil: Okrouhlík, Jan. University of South Bohemia; República Checa Fil: Mladěnková,Nella. University of South Bohemia; República Checa Fil: Antenucci, Carlos Daniel. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina

Brushtail possums have a wide geographical distribution. Possums from warmer, drier habitats have more frugal energy and water use and increased heat loss capacity at high temperatures. Consideration of geographical patterns in physiological attributes of source populations will likely improve the success and welfare outcomes of translocations., Identifying spatial patterns in the variation of physiological traits that occur within and between species is a fundamental goal of comparative physiology. There has been a focus on identifying and explaining this variation at broad taxonomic scales, but more recently attention has shifted to examining patterns of intra-specific physiological variation. Here we examine geographic variation in the physiology of brushtail possums (Trichosurus), widely distributed Australian marsupials, and discuss how pertinent intra-specific variation may be to conservation physiology. We found significant geographical patterns in metabolism, body temperature, evaporative water loss and relative water economy. These patterns suggest that possums from warmer, drier habitats have more frugal energy and water use and increased capacity for heat loss at high ambient temperatures. Our results are consistent with environmental correlates for broad-scale macro-physiological studies, and most intra-generic and intra-specific studies of marsupials and other mammals. Most translocations of brushtail possums occur into Australia’s arid zone, where the distribution and abundance of possums and other native mammals have declined since European settlement, leading to reintroduction programmes aiming to re-establish functional mammal communities. We suggest that the sub-species T. vulpecula hypoleucus from Western Australia would be the most physiologically appropriate for translocation to these arid habitats, having physiological traits most favourable for the extreme Ta, low and variable water availability and low productivity that characterize arid environments. Our findings demonstrate that geographically widespread populations can differ physiologically, and as a consequence some populations are more suitable for translocation to particular habitats than others. Consideration of these differences will likely improve the success and welfare outcomes of translocation, reintroduction and management programmes.

Made available in DSpace on 2018-11-26T20:04:44Z (GMT). No. of bitstreams: 0 Previous issue date: 2017-12-15 Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) Many adult mammals and birds respond to high surrounding temperatures with thermal tachypnea - an increase in breathing frequency accompanied by shallow tidal volume, with minimal increase in oxygen consumption (VO2). This pattern favors heat dissipation by evaporative water loss (EWL) through the respiratory tract. We asked to what extent this response was apparent at the earliest stages of development, when pulmonary ventilation initiates. Measurements of pulmonary ventilation (V. E; barometric technique), VO2 (open-flow methodology) and EWL (water scrubbers) were performed on chicken embryos at the earliest appearance of pulmonary ventilation, during the internal pipping stage. Data were collected, first, at the normal incubation temperature (37.5 degrees C); then, ambient and egg temperatures were increased to approximately 44 degrees C over a 2 h period. Other embryos of the same developmental stage (controls) were maintained in normothermia for the whole duration of the experiment. During heat exposure, the embryo's VO2 and carbon dioxide production increased little. In contrast, V. E more than doubled (similar to 128% increase), entirely because of the large rise in breathing frequency (similar to 132% increase), with no change in tidal volume. EWLdid not change significantly, probably because, within the egg, the thermal and water vapor gradients are almost nonexistent. We conclude that chicken embryos respond to a major heat load with tachypnea, like many adult mammals and birds do. Its appearance so early in development, although ineffective for heat loss, signifies that thermal tachypnea represents an important breathing response necessary to be functional from hatching. Sa Paulo State Univ, Dept Anim Morphol & Physiol, Coll Agr & Vet Sci, BR-14884900 Jaboticabal, SP, Brazil INCT Fisiol Comparada, Natl Inst Sci & Technol Comparat Physiol, BR-14884900 Jaboticabal, SP, Brazil McGill Univ, Dept Physiol, Montreal, PQ H3G 1Y6, Canada

Made available in DSpace on 2018-11-26T17:24:24Z (GMT). No. of bitstreams: 0 Previous issue date: 2017-05-01 Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) Because of their permeable skin, terrestrial amphibians are constantly challenged by the potential risk of dehydration. However, some of the physiological consequences associated with dehydration may affect aspects that are themselves relevant to the regulation of water balance. Accordingly, we examined the effects of graded levels of dehydration on the rates of evaporative water loss and water absorption through the skin in the terrestrial Neotropical toad, Rhinella schneideri. Concomitantly, we monitored the effects of dehydration on the mass of visceral organs; hematocrit and hemoglobin content; plasma osmolality; and plasma concentration of urea, sodium, chloride, and potassium. We found that dehydration caused an increase in the concentration of body fluids, as indicated by virtually all the parameters examined. There was a proportional change in the relative masses of visceral organs, except for the liver and kidneys, which exhibited a decrease in their relative masses greater than the whole-body level of dehydration. Changesor the preservationof relative organ masses during dehydration may be explained by organ-specific physiological adjustments in response to the functional stress introduced by the dehydration itself. As dehydration progressed, evaporative water loss diminished and water reabsorption increased. In both cases, the increase in body fluid concentration associated with the dehydration provided the osmotic driver for these changes in water flux. Additionally, dehydration-induced alterations on the cutaneous barrier may also have contributed to the decrease in water flux. Dehydration, therefore, while posing a considerable challenge on the water balance regulation of anurans, paradoxically facilitates water conservation and absorption. Univ Estadual Paulista, Inst Biociencias, Dept Zool, Sao Paulo, Brazil Univ Estadual Paulista, Ctr Estudos Ambientais, Sao Paulo, Brazil Univ Estadual Paulista, Inst Biociencias, Dept Zool, Sao Paulo, Brazil Univ Estadual Paulista, Ctr Estudos Ambientais, Sao Paulo, Brazil FAPESP: 10/20061-6 FAPESP: 13/04190-9 CNPq: 302045/2012-0 CNPq: 306811/2015-4 CNPq: 13/14-25780

International audience; Amphibians are the vertebrate group most affected by global change. Their highly permeable skin is involved in maintaining homeostasis (e.g. water and electrolyte equilibrium), which makes them particularly vulnerable to climate warming and skin pathogens. This study focused on the impacts of both desiccation (as a potential consequence of climate warming) and exposure to Batrachochytrium dendrobatidis (Bd), an emergent skin pathogen of amphibians. Bd causes chytridiomycosis, a lethal skin disease of amphibians, and is responsible for mass mortality events in several regions of the world. Because Bd colonizes the superficial layers of the epider- mis, it is assumed to affect water transfer across the skin. We investigated the behavioural pos- tures of the palmate newt Lissotriton helveticus expressed in response to desiccation and their influence on transepidermal water loss (TEWL) rate. We also investigated the effects of repeated 24 h exposure to Bd (i.e. every 4 d for 16 d) on the TEWL and ventral water absorption (VWA) rates of these newts. Our results suggest an efficient behavioural water-conserving mechanism, i.e. an 'S'-shaped posture associated with a restricted activity rate, not affected by repeated exposure to Bd. Similarly, TEWL was not significantly affected in exposed newts. VWA was significantly reduced after just 24 h exposure to Bd without modification until the end of the experiments. Our results suggest that Bd could rapidly inhibit rehydration of L. helveticus through fungal toxins and disrupt an essential function for survival.

We investigated the applicability of 2H to measure the amount of body water (TBW) and water fluxes in relation to diet type and level of food intake in a mollusk-eating shorebird, the Red Knot ( Calidris canutus). Six birds were exposed to eight experimental indoor conditions. Average fractional 2H turnover rates ranged between 0.182 day−1 (SD = 0.0219) for fasting birds and 7.759 day−1 (SD = 0.4535) for birds feeding on cockles ( Cerastoderma edule). Average TBW estimates obtained with the plateau method were within the narrow range of 75.9–85.4 g (or between 64.6 and 70.1% of the body mass). Those obtained with the extrapolation method showed strong day-to-day variations (range 55.7–83.7 g, or between 49.7 and 65.5%). Average difference between the two calculation methods ranged between 0.6% and 36.3%, and this difference was strongly negatively correlated with water flux rate. Average water influx rates ranged between 15.5 g/day (fasting) and 624.5 g/day (feeding on cockles). The latter value is at 26.6 times the allometrically predicted value and is the highest reported to date. Differences in 2H concentrations between the blood and feces (i.e., biological fractionation) were small but significant (−3.4% when fed a pellet diet, and −1.1% for all the other diets), and did not relate to the rate of water flux (χ2 1 = 0.058, P < 0.81). We conclude that the ingested water equilibrated rapidly with the body water pool even in an avian species that shows record water flux rates when living on ingested marine bivalves.

A common perception is that desert birds experience greater extremes of heat and aridity than their mammalian counterparts, in part, because birds do not use burrows as a refuge from the desert environment. We report observations of Dunn's Larks (Eremalauda dunni), Bar-tailed Desert Larks (Ammomanes cincturus), Black-crowned Finch Larks (Eremoprerix nigriceps), and Hoopoe Larks (Alaemon alaudipes) using burrows of the large herbivorous lizard Uromastyx aegypticus as thermal refugia during hot summer days in the Arabian Desert. Continuous recordings of shade air temperature (T-a), soil surface temperature (T-surface), burrow air temperature (Ta-burrow), and burrow substrate temperature (T-substrate) showed that T-surface exceeded 60 degrees C on most days. T-a typically exceeded 45 degrees C, whereas Ta-burrow was around 41 degrees C during midday. Calculations of total evaporative water loss at different temperatures indicated that Hoopoe Larks can potentially reduce their water loss by as much as 81% by sheltering in Uromastyx burrows during the hottest periods of the summer day.

We integrated population, climate, and physiological data to investigate Fynbos birds' vulnerability to recent climate change. Species occupying cool ranges with warming trends showed greatest declines. However, physiological thermal thresholds varied and, with the exception of Cape Rockjumper, were not clearly the cause of decline in most species., We investigated whether observed avian range contractions and population declines in the Fynbos biome of South Africa were mechanistically linked to recent climate warming. We aimed to determine whether there were correlations between preferred temperature envelope, or changes in temperature within species' ranges, and recent changes in range and population size, for 12 Fynbos-resident bird species, including six that are endemic to the biome. We then measured the physiological responses of each species at air temperatures ranging from 24 to 42°C to determine whether physiological thermal thresholds could provide a mechanistic explanation for observed population trends. Our data show that Fynbos-endemic species occupying the coolest regions experienced the greatest recent reductions in range and population size (>30% range reduction between 1991 and the present). In addition, species experiencing the largest increases in air temperature within their ranges showed the greatest declines. However, evidence for a physiological mechanistic link between warming and population declines was equivocal, with only the larger species showing low thermal thresholds for their body mass, compared with other birds globally. In addition, some species appear more vulnerable than others to air temperatures in their ranges above physiological thermal thresholds. Of these, the high-altitude specialist Cape rockjumper (Chaetops frenatus) seems most at risk from climate warming. This species showed: (i) the lowest threshold for increasing evaporative water loss at high temperatures; and (ii) population declines specifically in those regions of its range recording significant warming trends. Our findings suggest that caution must be taken when attributing causality explicitly to thermal stress, even when population trends are clearly correlated with rates of warming. Studies explicitly investigating the mechanisms underlying such correlations will be key to appropriate conservation planning.

Ostrich chick mortality was studied in 2522 chicks that were hatched artificially during the 1999/2000 breeding season. High levels of mortality were observed, with 1978 (78.4 %) of these chicks dying before 90 days after hatching. Atotal of 46.7 %(1177) of these chicks died before 28 days of age, and a further 30.7 %(801) died between 28 and 90 days post-hatching. Chick mortality to 28 days of age could not be conclusively related to sex, day of external pipping or breeder diet. Mortality rates were higher (P 0.05) at the beginning and end of the breeding season than in the middle months. Differences in mortality levels of chicks incubated in different incubators could be related to the time of the breeding season during which the incubator was mostly used. The regression of chick mortality to 28 days of age on day-old chick mass followed a 2nd-degree polynomial. Chicks with day-old masses below 762.5 g were particularly at risk of dying before 28 days after hatching. Chicks hatching from eggs where excessive water loss to 35 days of incubation (18 %) was recorded were also at risk of succumbing before 28 days of age. Chick mortality percentages for the period from 28 to 90 days of age exceeded 80 %in chicks weighing an average of 1050 g at 28 days. Mortaliy percentages declined sharply at higher live masses, to between 20 and 30 % in chicks weighing ?1950 g. This 'core' level of mortality remained throughout, even in the heaviest chicks. It was concluded that the high levels of chick mortality could be related to stress in chicks, resulting from an inability to adapt to the rearing environment. The high subsequent mortality percentages of low live mass chicks that survived to 28 days after hatching could probably be attributed to residual setbacks suffered earlier. Abetter understanding of the underlying principles involved in ostrich chick mortality in intensive rearing environments is required for progress in this field, resulting in more predictable survival rates under these conditions.

Birds living in desert environments have been the preferred models for the study of physiological adaptations to water scarcity. Passerine birds living in marine coastal habitats face similar problems, yet physiological adaptations to water conservation in such species have been poorly documented. We measured total evaporative water loss (TEWL) and rates of oxygen consumption (VO2) in three species of passerine birds dwelling in marine and fresh water habitats. Mass specific total evaporative water loss was significantly lower in the marine species, Cinclodes nigrofumosus, than in species inhabiting areas near freshwater sources. We found a positive relationship between TEWL and VO2. The ratio of TEWL to VO2 (relative evaporative water loss, RTEWL) showed significant variation among Cinclodes species, and was highest for the fresh-water living species, C. oustaleti and C. fuscus. The variation in TEWL found in Cinclodes is likely a consequence of differential exploitation of marine prey with high osmotic loads, which, in turn, may impose the need for water conservation