Migratory constraints on yolk precursors limit yolk androgen deposition and underlie a brood reduction strategy in rockhopper penguins

Hormonally mediated maternal effects link maternal phenotype and environmental conditions to offspring phenotype. The production of lipid-rich maternal yolk precursors may provide a mechanism by which lipophilic steroid hormones can be transported to developing yolks, thus predicting a positive correlation between yolk precursors in mothers and androgen levels in eggs. Using rockhopper penguins (Eudyptes chrysocome), which produce a two-egg clutch characterized by extreme egg-size dimorphism, reversed hatching asynchrony and brood-reduction, we examined correlations between circulating concentrations of the primary yolk-precursor vitellogenin (VTG) and levels of yolk androgens. Previous work in Eudyptes penguins has shown that egg-size dimorphism is the product of migratory constraints on yolk precursor production. We predicted that if yolk precursors are constrained, androgen transport to developing yolks would be similarly constrained. We reveal positive linear relationships between maternal VTG and androgens in small A-eggs but not larger B-eggs, which is consistent with a migratory constraint operating on the A-egg. Results suggest that intra-clutch variation in total yolk androgen levels depends on the production and uptake of yolk precursors. The brood reduction strategy common to Eudyptes might thus be best described as the result of a migratory constraint.     

Hatching order and size-dependent mortality in relation to brood sex ratio composition in chinstrap penguins

The differential environmental sensitivity of the sexes hasstrong implications in the evolutionary history of species asit can alter sexual size dimorphism, population sex ratios,and the faculty of parents to manipulate offspring sex in relationto environmental conditions. We studied sexual differences inhatching patterns and evaluated sex- and size-related mortalityin relation to hatching order and brood sex ratios in the chinstrappenguin Pygoscelis antarctica, a moderately size-dimorphic species,with a modal clutch size of 2 eggs. We found that male, second-hatched,and large eggs showed shorter hatching periods than female,first-hatched, and small eggs. We also found a male-biased mortalityof nestlings in the colony. However, male mortality patternsdiffered depending on the brood sex ratio composition. Mortalityof male chicks in all-male broods was higher than in mixed broodsand higher than female mortality in all-female broods. Contrary,females from mixed brood showed higher mortality than theirmale nest mates and higher too than females in all-female broods.Second-hatched chicks also suffered from higher mortality thanfirst-hatched chicks. Our results indicate that both the superiorcompetitive capacity and the higher energy demand of the largersex constitute 2 causal factors explaining patterns of sex-biasedmortality. Both factors occur in the same species and in differentsituations of sibling competition shaped by brood sex ratiocomposition. This study constitutes a good example of how patternsof sex-related mortality can vary depending on nest environmentalcircumstances. Furthermore, our study suggests that hatchingperiod can be a mechanism underlying sexual differences in theembryonic period of birds.     

Relevance of forage fish in the diet of Magellanic penguins breeding in northern Patagonia,Argentina

We quantified the trophic niche of Magellanic penguins (Spheniscus magellanicus) breeding and moulting in Golfo San Jorge, Argentina, through conventional stomach content and stable isotope analysis. A total of 112 adults were flushed during the early and late chick stages of 2011 and 2012 at Isla Vernacci Norte, and at least 15 prey taxa were found, including fishes, cephalopods, crustaceans and polychaetes. Overall, Argentine anchovy (Engraulis anchoita) showed the highest contribution in terms of importance by mass (68.1–85.3%, depending on chick stage and year), except for the old chick stage in 2011 when the shortfin squid (Illex argentinus) was the main prey consumed (56.0%). Based on carbon and nitrogen isotopic values from a total of 256 blood samples, corresponding to young and old chicks and to adults of both sexes sampled throughout the incubation, chick and moult stages at the above mentioned colony and years, Bayesian mixing model outputs showed that Argentine anchovy was always the main prey (48–86%). Bayesian mixing model outputs obtained from adults of both sexes and their chicks during the late chick stage of 2013 at Isla Vernacci Norte, Isla Tova and Isla Leones also showed that Argentine anchovy was the main prey consumed. This is the first comprehensive assessment of Magellanic penguin diet composition in northern Patagonia, quantifying the relative contribution of prey in the diet of adults and chicks at different stages of the annual cycle and years, and confirms the relevance of a forage fish such as the Argentine anchovy in its trophic ecology.     

Brood reduction and brood division in coots

The probability of starvation of chicks increases through hatching order in broods of the coot, Fulica atra. After hatching chicks accompany, and are fed by, parents as they swim around the territory. The time that chicks are able to spend outside the nest increases rapidly with age, so that the earlier hatching chicks gain a feeding advantage over the later. Starvation of chicks occurs within 4–5 days of hatching. Even after this initial mortality there persist large differences in the parental feeding rates of individual chicks within a brood. These do not correlate with age and do not seem to be the result of sibling competition. Instead, the parents regulate which chicks accompany them on foraging trips and therefore actively maintain feeding differences within the brood. Chicks cannot counter this parental regulation and the least fed of the brood grow more slowly in spite of an increased self-feeding effort. The possible functions of this parental behaviour are discussed.     

A game theoretical approach to conspecific brood parasitism

We constructed a game theoretical model to predict optimal patternsof egg laying in systems where individuals lay in the nestsof others as well as in their own nests. We show that decreasingthe effect of position within an egg-laying sequence on theworth of an egg should lead to reduced parasitism. Indeed,parasitism can only flourish if the worth of an egg to its biological parent declines with the total number of eggs laid in that nest.Further, we found that increasing the intrinsic costs of eggproduction should lead to an increased propensity for conspecificbrood parasitism. The model also predicts that variation inhosts' ability to reject parasitic eggs has little effect on parasitism until this ability is well developed.     

An experimental approach to Neolithic shifting cultivation

Archaeobotanical data from Late Neolithic lake-shore dwellings (4300-3500 cal B.C.) in the northern Pre-alpine lowlands are interpreted in different ways. The presence of permanent arable fields as well as arable fields with short fallow phases and shifting cultivation with slash-and-burn has been discussed. To test these hypotheses experimentally we have been conducting tests in a forest northeast of Stuttgart since 1994. The slightly south-exposed experimental area of approximately 4.5 ha on a loess soil is covered by mixed deciduous forest, is available for at least 20 years and has been divided into 34 plots of 30 × 30 m. Up to 2001, five plots were used for experiments. The normal procedure is clearing, burning the dry small timber (less than 10 cm in diameter) and then growing winter cereals (bread wheat) for one season. The yields were between 2000 and 4000 kg grains per ha. The harvest was more or less free from weeds. First attempts with summer crops gave much lower yields of about 1100 kg grains per ha. Continuous cereal growing on the same place in the following years resulted in minimal to zero yields, mainly due to vigorous weed growth. These weeds are not crop weeds, but forest perennials of clearings and forest fringes. Ploughing to remove the weeds is not possible, because of the presence of roots and tree-stumps, most of them still living. Weed regulation by hoeing, burning or cattle grazing remains to be tested. Protection of the crop from game and birds is by fences and nets, but protection from mice seems difficult. In the spectra from the pollen traps, clearing and burning are strongly indicated, but cereal growing only slightly. Received July 31, 2001 / Accepted April 9, 2002     

An experimental approach to the fundamental principles of hemodynamics

An experimental model has been developed to give students hands-on experience with the fundamental laws of hemodynamics. The proposed experimental setup is of simple construction but permits the precise measurements of physical variables involved in the experience. The model consists in a series of experiments where different basic phenomena are quantitatively investigated, such as the pressure drop in a long straight vessel and in an obstructed vessel, the transition from laminar to turbulent flow, the association of vessels in vascular networks, or the generation of a critical stenosis. Through these experiments, students acquire a direct appreciation of the importance of the parameters involved in the relationship between pressure and flow rate, thus facilitating the comprehension of more complex problems in hemodynamics.     

Relationships between brood size and parental provisioning performance in chinstrap penguins during the chick guard phase

 Chinstrap penguins (Pygoscelis antarctica) normally lay two eggs, but brood size is often reduced by mortality during incubation or after hatching. We hypothesized that this variation in brood size would affect the parents’ foraging behavior and their chick provisioning performance. We studied patterns of adult foraging trip duration and frequency, food load delivery, and chick growth rates in relation to brood size during the guard phase in four breeding seasons (1991–1994) on Seal Island, Antarctica. Within a given year, parents with two chicks made more frequent foraging trips to sea and may have transported larger food loads to the nest; however, the duration of foraging trips was unrelated to brood size. Overall, parents with two chicks spent ∼15% more time at sea than parents with only one chick. Both the frequency and duration of foraging trips varied between years. Foraging trip duration may partly reflect the birds’ foraging radius, which probably varies with time in response to shifts in krill distribution. Chick growth rate varied betwen years, but was related to brood size only in 1992, when chicks from two-chick broods grew significantly more slowly than chicks from one-chick broods. Food loads transported to chicks, as well as chick growth rates, were highest in 1994, when concurrent hydroacoustic studies indicated that regional krill biomass was severely depressed. This apparent anomaly suggests that the spatial scale of the krill survey may have been too coarse to detect some high-density krill aggregations within the penguins’ foraging range. Received: 26 September 1995 / Accepted: 12 May 1996     

Delayed start of full incubation is related to a delayed brood patch development in Magellanic Penguins Spheniscus magellanicus

Capsule: Most bird species start full incubation before the clutch is complete, others long after laying the last egg, gradually increasing the incubation temperature as incubation progresses, as is the case in Magellanic Penguins Spheniscus magellanicus.

Aims: In this study, we analysed the possible mechanisms underlying this delayed start of full incubation in Magellanic Penguins. We postulated two non-mutually exclusive, possible mechanisms: delayed development of the brood patch and/or variation in the adults’ incubation postures.

Methods: We conducted our work at Isla Quiroga, Argentina during 2011–13. We measured the development of the brood patch area, its temperature, the egg temperature and observed the postures of adult penguins.

Results: We found no association between the adults’ postures and the delayed start of full incubation, nor between brood patch temperature and the increase in egg temperature as incubation proceeded (egg age). The intercept of the curve between egg temperature and egg age was positively related to brood patch area, while the slope of the curve was negatively related.

Conclusion: Our results support the hypothesis that the shape of the curve of the increase in egg temperature as incubation proceeds is a consequence of the increase in the extension of the brood patch area.     

Egg temperature and initial brood patch area determine hatching asynchrony in Magellanic penguin Spheniscus magellanicus

In birds, the adaptive significance of hatching asynchrony has been under debate for many years and the parental effects on hatching asynchrony have been largely assumed but not often tested. Some authors suggest that hatching asynchrony depends on the incubation onset and many factors have been shown to influence hatching asynchrony in different species. Our objective was to analyze the exact timing of the onset of incubation and if this affects hatching asynchrony; and, in addition, which other factors (brood patch development, incubation position, adult body condition, intra‐clutch egg dimorphism, laying date and year) affect hatching asynchrony in Magellanic penguins Spheniscus magellanicus. We first estimated the eggshell temperature at which embryo development starts, with a non‐destructive and novel method. We then recorded individual egg temperatures in 61 nests during incubation, and related them, and other breeding parameters, to hatching asynchrony. We also observed incubation positions in 307 nests. We found a significant positive relationship between hatching asynchrony and the temperature that the first‐laid egg experienced during egg laying and between hatching asynchrony and the initial brood patch area. We also found a negative relationship between hatching asynchrony and the difference in temperature between second and first‐laid eggs within a clutch, measured after the egg‐laying period was finished. We ruled out position of the eggs during incubation, adult body condition, egg volume, laying date, and study year as factors influencing hatching asynchrony. The egg temperature during laying and the difference in temperature between eggs of a clutch are determinants of hatching asynchrony in Magellanic penguins.