Sex‐specific growth in chicks of the sexually dimorphic Black‐tailed Godwit

Sexual size dimorphism (SSD) is common in birds and has been linked to various selective forces. Nevertheless, the question of how and when the sexes start to differentiate from each other is poorly studied. This is a critical knowledge gap, as sex differences in growth may cause different responses to similar ecological conditions. In this study, we describe the sex‐specific growth – based on body mass and five morphometric measurements – of 56 captive Black‐tailed Godwit Limosa limosa limosa chicks raised under ad libitum food conditions, and conclude that all six growth curves are sex‐specific. Females are the larger sex in terms of body mass and skeletal body size. To test whether sex‐specific growth leads to sex‐specific susceptibility to environmental conditions, we compared the age‐specific sizes of male and female chicks in the wild with those of Black‐tailed Godwits reared in captivity. We then tested for a relationship between residual growth and relative hatching date, age, sex and habitat type in which the wild chicks were born. Early‐hatched chicks were relatively bigger and in better condition than late‐hatched chicks, but body condition and size were not affected by natal habitat type. Female chicks deviated more negatively from the sex‐specific growth curves than male chicks for body mass and total‐head length. This suggests that the growth of the larger females is more susceptible to limiting environmental conditions. On average, the deviations of wild chicks from the predicted growth curves were negative for all measurements, which suggests that conditions are limiting in the current agricultural landscape. We argue that in estimating growth curves for sexually dimorphic species, it is critical first to make accurate sex and age determinations.     

Laboratory observations of moulting,growth and maturation in Antarctic krill (Euphausia superba Dana)

Summary Observations of intermoult period, growth and maturation were made on krill which were transported from Antarctic waters and maintained in the laboratory in Australia over a three year period. The mean intermoult period (IP) for each of 10 specimens, with initial body lengths of 24.7=46.8 mm, kept at -0.5° C varied from 22.0 to 29.8 days (overall mean = 26.6 days). These measurements of IP are significantly longer than those obtained in some previous studies. Differences in experimental temperatures, light, body sizes and growth patterns of the specimens between studies are unlikely to be causes of these dissimilar results. The pattern of changes in body length (BL) varies from one individual to the next. The greatest increase in BL over a series of 4–5 moults ranged from 0.024 to 0.070 mm/day, which is equivalent to 0.0020 to 0.0086/day in body weight, assuming exponential growth. This maximum growth rate is about half the rate predicted from the growth scheme of Mauchline (1980) for wild krill. Comparison of growth data for other euphausiids suggests that Mauchline's scheme produces anomalous growth rate. The slower growth rate observed in the present study would extend the estimated life span of krill from 3–4 years, as calculated by Mauchline (1980), to 4–7 years. If krill undergo body shrinkage during the Antarctic winter the estimated life span might be even longer. Examination of the external sexual characters of moults showed both progression and regression of maturity stage in association with changes in BL.     

Moulting interval and growth of juvenile Antarctic krill (Euphausia superba) fed different concentrations of the diatom Phaeodactylum tricornutum in the laboratory

Summary Juvenile Antarctic krill (Euphausia superba) were fed the diatom Phaeodactylum tricornutum at concentrations ranging from 0.0–5.0 mgCl^-1. Intermoult period (IP) decreased, but an increment of body length per moult (BL) of juvenile krill increased, up to a concentration of 1.0 mgCl^-1. No further effect of food concentrations on IP or BL was seen at concentration beyond 1.0 mgCl^-1. IP plateaued at 23.8 days and BL, 1.14 mm. The maximum daily growth rate (BL/IP) of juvenile krill was calculated to be 0.047 mm day^-1. BL and IP were correlated and the relationship is expressed as BL=-0.0066IP+2.47 (r=0.58, n=141, P<0.01). Growth conditions of krill in the Southern Ocean are discussed in terms of available food concentration in there.     

Daily activity and minimum food requirements during winter for gentoo penguins (Pygoscelis papua) in the South Shetland Islands, Antarctica

Estimates of daily activity and consequent demand for food during winter are scarce for many polar seabirds, yet essential for assessing constraints on foraging effort, demand for food, and potential competition with local fisheries. We affixed archival temperature tags to gentoo penguins (Pygoscelis papua) from two colonies in the South Shetland Islands to measure the frequency, timing, and duration of foraging trips and to estimate minimum food requirements during winter. Foraging trip frequencies ranged from 0.85 to 1.0 trips day⁻¹ and were positively correlated with day length. Early winter foraging trips more closely matched day length than late winter foraging trips. The data suggest that individuals maximize foraging time during the early winter period, likely to recover body mass following the breeding season and molt. The more attenuated response of foraging trip durations to increasing day length in late winter may be related to differences in local resource availability or individual behaviors prior to the upcoming breeding season. Minimum food requirements also exhibited a seasonal cycle with a mid-winter minimum. On average, minimum food requirements were estimated at 0.70 ± 0.12 kg day⁻¹. Extrapolated to the regional population of gentoo penguins, winter food requirements by gentoo penguins were equivalent to roughly 33% of annual krill catches by commercial fisheries in the South Shetland Island region over the past decade. Current expansion of the gentoo population and the krill fishery in the southern Scotia Sea warrants continued monitoring of gentoo penguins during winter.     

External parasite infestation depends on moult-frequency and age in Antarctic krill (<Emphasis Type="Italic">Euphausia superba</Emphasis>)

Levels of infestation by the suctorian Ephelota sp. were determined in Antarctic krill taken in the north-eastern region of the Scotia sea, close to the island of South Georgia. Individuals caught in summer between 2002 and 2004 were measured for body length, eye diameter, moult stage, sex, sexual maturity and number of suctorian parasites. All krill samples were found to contain at least two modal size-classes and the complete range of maturity- and moult stages. Sample analysis for the different characters identified moult stage as an important determinant of infestation by Ephelota sp., with 66% of those about to moult (pre-moult individuals) being infested compared with 0% of post-moult individuals. Multiple Correspondence Analysis also identified a strong link between age and degree of suctorian infestation with large-eyed (and hence old) males having very high numbers of suctorian bodies. Moulting is costly in terms of energy and vulnerability, but it is believed that krill moult at relatively high rates throughout their life to facilitate growth and “shrinkage”. Here, we demonstrate that the control of external parasitism is also a major advantage of moulting at high frequency. The capacity of krill to maintain a high moult rate decreases with age, leading to older krill enduring higher parasitic loads.     

The influence of feeding on the metabolic activity of Antarctic krill (Euphausia superba Dana)

Summary The influence of feeding on the metabolic activity of juvenile krill was assessed from 24h experiments in which krill were incubated with various concentrations of diatoms (Chaetoceros calcitrans, Phaeodactylum tricornutum, Thalassiosira eccentrica, Fragilariopsis vanheurkii), newly hatched Artemia nauplii and latex beads. Krill fed on the larger food more efficiently, with reluctant feeding on latex beads. Feeding of krill expressed as clearance rates was poorly correlated with their oxygen uptake rates. Instead, a positive correlation was found between the oxygen uptake rates and ingestion rate (except for latex beads). The result implies that the specific dynamic action is the major cause of the increased oxygen uptake of krill. Krill fed diatoms increased both ammonia and phosphate excretion with increasing ingestion rate, but only phosphate excretion was increased in parallel with ingestion rate for those fed Artemia nauplii. Assuming the daily ration of krill in the field is 5% of the body weight, and the major food source is phytoplankton, oxygen uptake, ammonia excretion and phosphate excretion rates of wild krill are estimated to be 1.6, 4.5 and 7.8, respectively, times the rates of non-feeding krill in 24h laboratory experiments. Krill offered various kinds of food showed different metabolic quotients (O/N, N/P and O/P ratios). While no functional relationship was seen between the metabolic quotient and the ingestion rate of krill fed Artemia nauplii, those fed Fragilariopsis showed a progressive decrease in O/N, N/P, and O/P ratios as their ingestion rates increased.     

The scheduling of spawning with the moult cycle in Northern krill (Crustacea: Euphausiacea): a strategy for allocating lipids to reproduction

Summary

Euphausiids moult and grow throughout their life, which implies sharing of resources between growth and reproduction for adult krill. In the Northern krill, Meganyctiphanes norvegica (M. Sars), female krill produce eggs cyclically. Spawning moult cycles alternate with vitellogenic moult cycles for lipid yolk accumulation. Histology shows that lipids are associated with the R cells of the digestive gland in both sexes, with the yolk platelets of mature oocytes and with the fat body cell membranes and blood lacunae in reproducing females. Mature female krill can have a total lipid content twice as high as males, mostly due to accumulation in the ovary, the fat body and the haemolymph. In contrast, in males, as well as in non-reproducing females, the highest percentage of lipids is found in the digestive gland and the haemolymph. In Meganyctiphanes norvegica, the most abundant lipid fractions are polar lipids and triglycerides, the latter being relatively low in reproducing female gonad and fat body. Triglycerides are believed to be a pure energy source and polar lipids are essential for membrane development in embryos. The fatty acid content and composition of the triglyceride and polar lipid fractions in females are different from males, related to both reproductive and dietary processes. Higher levels of polyunsaturated fatty acids (PUFA) in the polar lipid fraction were found in reproductive females. During the non-reproductive season, the converse was found, indicating the specific role PUFA and other fatty acids play in growth and egg production. Adaptive processes linked to reproduction were studied comparatively in three populations of the Northern krill—Clyde Sea (W, Scotland), Kattegat (E, Denmark), Ligurian Sea (Mediterranean)—all differing considerably in climatic and trophic conditions. Such adjustments in lipid synthesis and storage are viewed as reproductive strategies developed by the Northern krill in response to different environmental conditions.     

Eugregarine infection within the digestive tract of larval Antarctic krill, Euphausia superba

The infection rate of eugregarine parasites, Cephaloidophora pacifica, within the digestive tract of larval Antarctic krill, Euphausia superba, was examined using samples collected from the Indian sector of the Southern Ocean. Immature and mature eugregarine gamont stages were found at all larval stages older than Calyptopis I stage. Eugregarine infection in 14.0% (N = 108) of the first feeding stage (Calyptopis I) suggested that krill larvae are at risk from being infected during physiological transition from non-feeding to feeding stages. Eugregarine prevalence and intensity of infection at the three calyptopis stages increased with stage/krill length. Statistical analysis showed that the intensity of C. pacifica infection also increased with host density. Thus, krill density is probably a key determinant of the intensity of infection. We found gamont stage eugregarines in the host hind-gut, blocking the passage of food. Eugregarine infections in larval krill may have a negative impact on digestion and absorption in the host digestive tract.     

The krill maturity cycle: a conceptual model of the seasonal cycle in Antarctic krill

A long-term study on the maturity cycle of Antarctic krill was conducted in a research aquarium. Antarctic krill were either kept individually or in groups for 8 months under different temperature and food conditions, and the succession of female maturity stages and intermoult periods were observed. In all cases regression and re-maturation of external sexual characteristics were observed, but there were differences in length of the cycle and intermoult periods between the experimental conditions. Based on these results, and information available from previous studies, we suggest a conceptual model describing seasonal cycle of krill physiology which provides a framework for future studies and highlight the importance of its link to the timings of the environmental conditions.     

Factors affecting fledging weight of Adélie penguin (Pygoscelis adeliae) chicks: a modeling study

An individual-based model is developed to examine mechanisms that potentially underlie the observed constancy in fledging weight (2.8-3.2 kg) of Adélie (Pygoscelis adeliae) penguin chicks, in spite of large variability in the abundance of Antarctic krill (Euphausia superba), the primary food source. The model describes the energetic requirements of the chick, with growth resulting from the difference between assimilated energy and respiration. Parameterizations of these metabolic processes are based upon experimental and field observations. Ingestion of Antarctic krill by the chick is dependent on the frequency of food delivery to the chick by the adults, which is based on measured foraging times. The mass, size, and size frequency distribution of Antarctic krill fed to the chick are specified using empirical data. The energy content of the Antarctic krill provided to the chick is taken to be constant or allowed to vary with size. The simulations show that food availability is most critical in the latter portion of chick development, when growth rates and food demands are high. Low food availability during this time must be compensated by either feeding chicks with larger krill of higher caloric value or by increased assimilation efficiency. Periods when small krill with lower caloric value dominate require more frequent feeding of the chicks in order to attain their observed fledging weight. Thus, although the total food energy given to the chick is the main factor determining chick growth, the distribution of food availability relative to chick size (i.e., different net growth rates) and food quality are also factors influencing the fledging weight of penguin chicks. The simulations provide insight into the compensating effects of food delivery, food quality, and metabolic processes that allow Adélie penguin chicks to reach their observed fledging weight in spite of considerable environmental variability in food supply.     

Food and daily food consumption of southern minke whales in the Antarctic

Summary The stomach contents of 273 southern minke whales (Balaenoptera acutorostrata) taken in the region 55°S to the ice-edge and between 105°E and 115°E by a Japanese survey during 1987/88 were examined. The minke whales' dominant feeding ground coincided with a distinctive hydrographic front in the vicinity of the ice-edge at the mouth of Vincennes Bay. Krill (Euphausia superba) were the dominant food species comprising 100% and 94% by weight of stomach contents in the ice-edge and offshore zones, respectively. In the offshore zone, minke whales tend to feed on E. superba rather than Thysanoessa macrura, which is found more frequently than the former in net samples. Total food consumption by minke whales per day was estimated to be 1170 t (22.1 kg/km²) and 596 t (2.0 kg/km²) in the ice-edge and offshore zones, respectively. Feeding activity peaked in the early morning in the ice-edge zone, whereas it occurred irregularly throughout the day in the offshore zone. Two size modes of krill (25–28 mm in body length (1 year-old) and 41–48 mm (3–4 year-old)) dominated the diet. Although the former was numerically more abundant, the latter dominated the diet by weight, suggesting that larger krill are more important food for minke whales. Observed spatial pattern of krill populations in the study region suggests that movement of the subsurface cold water mass tended to carry krill offshore from the ice-edge.The earlier version of this paper was submitted to the 42nd Meeting of the Scientific Committee of International Whaling Commission as SC/42/SHM; 34.     

Environmental influences on the gametic investment of yellow dung fly males

The energetic investment per spermatozoon and in spermatogenesis is central to a male's reproductive strategy. Relatively little, however, is known about environmental influences on variation in male allocation decisions and associated trade-offs. Plasticity in sperm length and testis size in response to variable food and temperature conditions either before or after adult eclosion was investigated in Scathophaga stercoraria, a classic model organism for sperm competition. Both measures showed interesting and clear environmental effects and also a heritable component. Testis length, and thus presumably sperm production, showed a hypoallometric (b < 1), but non-linear increase with body size, indicating that the allometric relationship changed with size. Like body size, testis length decreased with increasing developmental temperatures, but also showed a complex cubic relationship with adult temperatures. In contrast, sperm length increased or showed a negative quadratic relationship with increasing temperatures. The increase of within-male variation in sperm length with increasing developmental temperature and decreasing adult food indicates that some of our treatments were stressful. Nevertheless, there was no evidence of a trade-off between testis size and sperm length. The missing effect of adult or larval food availability on testis and sperm length, despite strong effects of larval food on body size, suggests that investment into reproduction is less sensitive to food restriction than investment into growth.     

Experiments on the physiology of southern and northern krill,Euphausia superba and Meganyctiphanes norvegica,with emphasis on moult and growth – a review

Physiological data are needed for life history studies on krill, and as parameters for input into energy budgets and models. In conjunction with moult and growth data, these may also prove useful for assessing the fishable biomass of krill. Here, the development of physiological concepts in experimental krill research is briefly evaluated, with emphasis on the gaps to be filled. Krill growth is very flexible, as well as strongly temperature and nutrition dependent. The polar Antarctic krill Euphausia superba grows as fast as the boreal species Meganyctiphanes norvegica, at least during the first 2.5 years, and the species are comparable in terms of physiological plasticity. Accordingly, as krill appear to adjust quickly to specific laboratory conditions, short-term experiments are essential if field conditions are to be reflected as closely as possible. Furthermore, direct comparisons between laboratory experiments and swarming studies in the field are advantageous. For these, M. norvegica is particularly well-suited, as swarms can be followed over longer times and more easily than in E. superba. For example, processes of moult and reproduction were found to be highly coordinated in swarms and populations of Northern krill. For this species a conceptual model of reproduction was developed based on a combination of short-term laboratory observations coupled with field data on moult and ovary stages. In further physiological experiments krill should be studied as groups when swarming. Using proxies, that is applying physiological and/or biochemical methods side by side, is a promising way to enhance the reliability of life history data.     

A comparison of Antarctic petrel (Thalassoica antarctica) diets with net samples of Antarctic krill (Euphausia superba) taken from the Prydz Bay region

Samples of the stomach contents of Antarctic petrels (Thalassoica antarctica) were obtained on board ship in the Prydz Bay region of Antarctica from birds which had spontaneously regurgitated. The weight of food and the species composition of the stomach contents were measured. Antarctic krill,Euphausia superba, the sole prey item taken, were compared to krill obtained by nets in the same region as part of a large-scale krill survey. Krill from petrel stomach samples were larger in mean size than those sampled by nets. This may be attributed to selection of the larger sized krill by the petrels, it may be caused by the nets sampling different populations of krill or it may be due to net avoidance by the larger krill.     

The components of life-history traits variation in Daphnia magna population

The variation of six growth and reproduction traits in the parthenogenetic clones of Daphnia magna and its relationships with the genotypes at 5 isozyme loci were studied under controlled conditions at three food concentrations. A significant genetic component of age at maturity, growth rate and fecundity variation was detected. Nested ANOVA revealed the influences of the genotypes of the isozyme loci, that are more pronounced at the high food concentration. In some cases the genotype-environment interactions were found.The variation of body length at 6 days and total number of offspring includes a significant heterozygosity — environment interaction. The most heterozygous clones show the least differences in life history traits between food conditions.Life-history traits heritability, genotype-environment interactions and different reproductive strategies of the clones are discussed.     

Influence of sampling protocol on diet determination of gentoo penguins Pygoscelis papua and Antarctic fur seals Arctocephalus gazella

The influence of two sampling protocols on diet determination of two marine predators, the gentoo penguin (Pygoscelis papua) and Antarctic fur seal (Arctocephalus gazella), was investigated. The collection of diet samples on three occasions over a 2-week period was compared with collecting all samples during a single session, as current CCAMLR Ecosystem Monitoring Programme protocols recommend. Some differences in the mass of food recovered from penguins were found but this was attributed to the mass of penguin sampled. There were no differences in diet composition between protocols and although body mass was a significant determinant of the mean length of krill Euphausia superba recovered from penguins, there were no differences between sampling protocols. This study has shown that differences between sampling frequencies are small and a variety of sampling protocols can produce results acceptable for inter-annual monitoring. The mass of sampled individuals can account for significant variation and should be recorded, especially if sampling frequencies and sizes are low. Accepted: 24 March 1999     

Biological characteristics of euphausiids preyed upon by Adélie penguins in relation to sea-ice conditions in Lützow-Holm Bay,Antarctica

We examined the biological characteristics of euphausiids found in the stomachs of Adélie penguins in relation to sea-ice conditions in Lützow-Holm Bay over three seasons. Euphausiids, especially Euphausia superba, proved to be a staple food for Adélie penguins irrespective of the ice condition. Body length and maturity-stage compositions of euphausiids were different among seasons, probably reflecting sea-ice condition in summer. The mean body length decreased and maturity regressed during each season in E. superba, which was partly attributable to the selective feeding on large, mature female krill by Adélie penguins. The 1995/1996 year class of E. superba, which was spawned when the sea ice was most developed, was strong and conspicuous in the 1996/1997 and 1997/1998 seasons. This vigor indicates that sea ice provided females with good spawning conditions and larvae with good growth and survival rates.     

Resource dependent growth and body condition dynamics in juvenile snakes: an experiment

We examined the manner in which animals adjust the proportion of energy allocated to growth and storage in response to food availability. We compared univariate growth and length-mass relationships between juvenile adders (Vipera berus) reared under two different feeding regimes. Animals in the low- and high-food experimental groups were fed suckling mice once and twice weekly, respectively. Snout-vent length, body mass, and body condition (residual scores from log-log regression of body mass on snout-vent length) were measured shortly after birth, and at 4, 9, and 14 weeks. We found that growth in length and mass, as well as changes in length-mass relationships, differed between treatments; snakes with access to more food not only increased faster in length but were also heavier at the completion of the experiment than were similar sized less frequently fed snakes. There was no association between body condition of individuals measured at birth and at the end of the experiment, whereas size at birth was a good predictor of final size. Our results provide evidence for resource-dependent allocation strategies in V. berus, and suggest that somatic growth is less sensitive to environmental fluctuations than body condition, presumably because body size is of greater importance for fitness.     

A laboratory study on digestive processes in the Antarctic krill,Euphausia superba,with special regard to chitinolytic enzymes

Feeding experiments of 9, 14 and 20 days duration were carried out on the Antarctic krill, Euphausia superba. Two groups were fed with the chitinous diatom Cyclotella cryptica and the non-chitinous green algae Dunaliella bioculata, respectively. A control group remained unfed. The time courses of the activities of endo- and exochitinase in the stomach and the midgut gland were compared with those of the digestive enzymes protease, cellulase (1,4-β-d-glucanase) and laminarinase (1,3-β-d-glucanase). Specific activities of all enzymes were higher in the stomach than in the midgut gland. Characteristic time courses of activity were evident after 4 days. In starved animals, enzyme activities decreased to a minimum after 4 days and recovered within 14 days to initial values. In the stomach, the activities of endo- and exochitinase increased when krill were fed on Cyclotella. For animals fed with Dunaliella, activities stayed constant or decreased slightly. The results confirm chitinases as digestive enzymes and, therefore, the capability of krill to utilize various food sources. Accepted: 4 October 1998     

Effects of temperature and food level on growth and development of a planktonic water mite

We analysed the relative effects of food availability and temperature on rates of growth and development of a predatory planktonic water mite, Piona exigua. Growth in length of mites fed Daphnia, Ceriodaphnia and Chydorus was analysed by Gompertz or von Bertalanffy curves; these curves were compared by parallel curve analysis. Growth rates of nymphs and adult female mites increased with temperature; the duration of the imagochrysalis stage decreased. Females grown at 10 °C were smaller at final size than females grown at 15 °C, 18 °C or 22 °C. Females reared at food levels of 15 or 30 prey l⁻¹ grew more slowly and were smaller than those provided with 60 or 120 prey l⁻¹. Nymphs grew more slowly when Daphnia were the only prey, than when smaller prey were available. Food level did not affect nymph growth at 10 °C or 15 °C, but growth at 18 °C or 22 °C may have been slowed at the lowest food levels. Synergistic effects of temperature and food level on nymph growth were apparent only from analysis of growth curves and not from stage duration data.