Variation in body mass dynamics among sites in Black Brant Branta bernicla nigricans supports adaptivity of mass loss during moult

Birds employ varying strategies to accommodate the energetic demands of moult, one important example being changes in body mass. To understand better their physiological and ecological significance, we tested three hypotheses concerning body mass dynamics during moult. We studied Black Brant in 2006 and 2007 moulting at three sites in Alaska which varied in food availability, breeding status and whether geese undertook a moult migration. First we predicted that if mass loss during moult were simply the result of inadequate food resources then mass loss would be highest where food was least available. Secondly, we predicted that if mass loss during moult were adaptive, allowing birds to reduce activity during moult, then birds would gain mass prior to moult where feeding conditions allowed and mass loss would be positively related to mass at moult initiation. Thirdly, we predicted that if mass loss during moult were adaptive, allowing birds to regain flight sooner, then across sites and groups, mass at the end of the flightless period would converge on a theoretical optimum, i.e. the mass that permits the earliest possible return to flight. Mass loss was greatest where food was most available and thus our results did not support the prediction that mass loss resulted from inadequate food availability. Mass at moult initiation was positively related to both food availability and mass loss. In addition, among sites and years, variation in mass was high at moult initiation but greatly reduced at the end of the flightless period, appearing to converge. Thus, our results supported multiple predictions that mass loss during moult was adaptive and that the optimal moulting strategy was to gain mass prior to the flightless period, then through behavioural modifications use these body reserves to reduce activity and in so doing also reduce wing loading. Geese that undertook a moult migration initiated moult at the highest mass, indicating that they were more than able to compensate for the energetic cost of the migration. Because Brant frequently change moult sites between years in relation to breeding success, the site‐specific variation in body mass dynamics we observed suggests individual plasticity in moult body mass dynamics.     

Nitrogenous excretion in the Antarctic plunderfish

The nitrogenous excretion rates (total ammonia nitrogen, urea, and primary amines) of plunderfish Harpagifer antarcticus were related significantly to length and to wet mass (mass exponents of 0·94, 1·01, 1·07 and 0·93 for total ammonia nitrogen, urea, primary amines, and total nitrogen, respectively). The routine total ammonia excretion rates [22·23 & 2·0 mg N kg⁻¹ day⁻¹ (mean±S.E.)] of plunderfish measured in Antarctica are 10–69% lower than those of comparable non-polar species. Plunderfish are ammonotelic, but the proportion of the total nitrogenous waste attributable to each category was variable between individuals. On average (ranges in parentheses), total ammonia nitrogen, urea, and primary amines accounted for c .82 (57–97), 13 (2–28), and 5 (0·6–22)%, respectively, of the total nitrogen excreted. Polar fish differ from their non-polar relatives only in the rate, and not the nature, of their nitrogenous waste excretion processes.     

Wing moult and mass change in free‐living mallard Anas platyrhynchos

Captured free‐living male mallard Anas platyrhynchos at Abberton in southern Britain showed peak mass gain immediately prior to simultaneous remex moult. Individuals of both sexes were heavier before shedding wing feathers than when flightless confirming literature accounts that show mallard accumulate fat stores in anticipation of moult to contribute to meeting energy needs during remex re‐growth. Over the course of four seasons, males lost 13 17% of initial body mass on average during re‐growth of flight feathers, females 13 23%. Based on energy expenditure of 1.3 times BMR, male mallard were estimated to be able to fulfil 42 60% and females 41 82% of their energy needs throughout moult from stores. Free‐flying male mallard fed ad libitum in a predator‐free environment did not differ in starting body mass or rate of mass loss during wing moult compared to free‐living Abberton birds, suggesting depletion of fat stores, irrespective of available sources of exogenous energy. Based on this evidence, we reject that the hypotheses that mass loss in moulting mallard is due to 1) simple energy stress and 2) restrictions on feeding and consider that 3) attaining the ability to fly at an earlier stage on incompletely grown flight feathers is not the primary factor shaping this trait. Rather, we consider the accumulation and subsequent depletion of fat stores, together with reductions in energy expenditure, enable mallard to re‐grow feathers as rapidly as possible by exploiting habitats that offer safety from predators, but do not necessarily enable them to balance energy budgets during the flightless period of remex feather re‐growth.     

Non‐breeding Cackling,Ross's and Snow Geese on Baffin Island show no loss of body mass during wing moult

Non‐breeding Cackling Branta hutchinsii, Ross's Anser rossii and Lesser Snow Geese Anser caerulescens caerulescens captured during remigial moult on Baffin Island in 2015 showed no loss of body mass with moult stage, and individual variation in mass was largely explained by sex and measures of body size (tarsus length). Exceptional conditions in 2015 resulted in almost no reproductive effort or success in that year, so captured geese of all three species were likely to have been non‐breeding individuals that initiated moult early, whereas there were almost no failed or successful breeders, which would normally moult later. This suggests that in a non‐breeding year (i.e. in the absence of competition from large numbers of goslings), locally moulting geese can obtain sufficient exogenous energy to meet their needs during the flightless wing moult period without losing body mass. This also is consistent with the hypothesis that in other species of geese, accumulation of fat stores prior to, and depletion of such stores during, wing moult is adaptive and likely to be a feature of individual plasticity to meet particular needs, such as undertaking moult migration to remote sites where precise foraging and predation conditions cannot be anticipated, or where competition from more dominant individuals may restrict their access to a reliable food supply.     

Changes in body mass and organ size during wing moult in non-breeding greylag geese Anser anser

The "cost-benefit" hypothesis states that specific body organs show mass changes consistent with a trade-off between the importance of their function and cost of their maintenance. We tested four predictions from this hypothesis using data on non-breeding greylag geese Anser anser during the course of remigial moult: namely that (i) pectoral muscles and heart would atrophy followed by hypertrophy, (ii) leg muscles would hypertrophy followed by atrophy, (iii) that digestive organs and liver would atrophy followed by hypertrophy and (iv) fat depots be depleted. Dissection of geese captured on three different dates during wing moult on the Danish island of Saltholm provided data on locomotory muscles and digestive organ size that confirmed these predictions. Locomotory organs associated with flight showed initial atrophy (a maximum loss of 23% of the initial pectoral muscle mass and 37% heart tissue) followed by hypertrophy as birds regained the powers of flight. Locomotory organs associated with running (leg muscles, since geese habitually run to the safety of water from predator-type stimuli) showed initial hypertrophy (a maximum gain of 37% over initial mass) followed by atrophy. The intestines and liver showed initial atrophy (41% and 37% respectively), consistent with observed reductions in daily time spent feeding during moult, followed by hypertrophy. The majority of the 22% loss in overall body mass (mean 760 g) during the flightless period involved fat utilisation, apparently consumed to meet shortfalls between daily energetic needs and observed rates of exogenous intake. The results support the hypothesis that such phenotypic plasticity in size of fat stores, locomotor and digestive organs can be interpreted as an evolutionary adaptation to meet the conflicting needs of the wing moult.     

Changes in body mass and organ size during remigial moult in common scoter Melanitta nigra

The “cost‐benefit” hypothesis states that avian body organs show mass changes consistent with the trade‐off between their functional importance and maintenance cost, which may vary throughout the annual cycle. Flightless moulting common scoter Melanitta nigra in Danish marine waters select rich undisturbed offshore feeding areas lacking predators, suggesting active feeding during moult. We tested four predictions relating to organ size during flightlessness in moulting male common scoter under this hypothesis. Namely that (i) pectoral muscles would show atrophy followed by hypertrophy, but that there would be no change in (ii) leg muscles and heart (the locomotory architecture required to sustain diving for food), (iii) digestive organs and liver (required to process food), or (iv) fat deposits (because birds could fulfil daily energy requirements from locally abundant food resources). Dissection of scoters collected at different stages during wing moult south of the Danish island of Læsø provided data on organ size that were consistent with these predictions. Pectoral muscle mass showed a c.23% atrophy during the middle of the flightless period relative to that at the end of moult. There was no significant loss in leg muscle, heart, digestive organs (except gizzard mass), liver, fat reserves or body mass with remigial growth. These findings are consistent with the hypothesis that common scoter moult in a rich feeding area, and rely on their diet to meet the nutritional requirements of remigial moult. These results differ in detail from those of a similar study of terrestrial feeding moulting greylag geese Anser anser, but because of the widely differing ecology of the species concerned, both sets of findings provide strong support for the hypothesis that variations in phenotypic plasticity in size of fat stores, locomotor and digestive organs can be interpreted as evolutionary adaptations to meet the conflicting needs (feather growth, nutritional challenges and predator avoidance) of the flightless moult period in different Anatidae species.     

Nutrient absorption and utilization by wing and flight muscle morphs of the cricket Gryllus firmus: implications for the trade-off between flight capability and early reproduction

Absorption efficiency (AD, approximate digestibility, assimilation efficiency) of various macronutrients and conversion of absorbed nutrients to biomass (ECD) were compared among the two types of flightless morph and the flight-capable morph of the cricket, Gryllus firmus. No biologically significant phenotypic or genetic difference in AD for carbohydrate, protein or lipid was observed among morphs fed either a high-nutrient (100%) or a low-nutrient (25%) diet. Thus, previously-documented differences among adult morphs in carbohydrate and lipid content must be caused by processes other than variation in nutrient absorption by morphs during adulthood. Relative absorption efficiency of total dry mass of food by morphs of G. firmus appears to be a valid indicator of relative AD of total calories. Morphs did not differ phenotypically or genetically in the excretion of end products of nitrogen metabolism (uric acid, hypoxanthine plus xanthine) on either the high nutrient or the low nutrient diet. Nutritional indices corrected for excreted nitrogenous metabolites were very similar to uncorrected indices, and the pattern of variation among the morphs was the same for corrected or uncorrected values. Each of the two types of flightless morph converted a greater proportion of absorbed nutrients into body mass, mainly ovaries, and allocated a smaller proportion of assimilated nutrients to respiration than did the flight-capable morph. Moreover, the trade-off between respiration and early reproduction was substantially magnified on the low nutrient diet. These results extend previous findings of a trade-off between flight capability and early reproduction in wing-polymorphic Gryllus species (1) to diets of very different nutrient quantity, and (2) to flightlessness arising from different causes: blockage of flight muscle development in juveniles vs histolysis of fully-developed flight muscles in adults.     

Late‐moulting Black‐necked Grebes Podiceps nigricollis show greater body mass in the face of failing food supply

From August to December, thousands of Black‐necked Grebes Podiceps nigricollis concentrate during the flightless moult period in salt ponds in the Odiel Marshes, southern Spain, where they feed on the brine shrimp Artemia parthenogenetica. We predicted that because Black‐necked Grebes moulted in a food‐rich, predator‐free environment, there would be no net loss of body mass caused by the use of fat stored to meet energy needs during remigial feather replacement (as is the case for some other diving waterbirds). However, because the food resource disappears in winter, we predicted that grebes moulting later in the season would put on more body mass prior to moult because of the increasing risk of an Artemia population crash before the moult period is completed. Body mass determinations of thousands of birds captured during 2000–2010 showed that grebes in active wing‐moult showed greater mass with date of capture. Early‐moulting grebes were significantly lighter at all stages than late‐moulting birds. Grebes captured with new feathers post‐moult were significantly lighter than those in moult. This is the first study to support the hypothesis that individual waterbirds adopt different strategies in body mass accumulation according to timing of moult: early‐season grebes were able to acquire an excess of energy over expenditure and accumulate fat stores while moulting. Delayed moulters acquired greater fat stores in advance of moult to contribute to energy expenditure for feather replacement and retained extra stores later, most likely as a bet hedge against the increasing probability of failing food supply and higher thermoregulatory demands late in the season. An alternative hypothesis, that mass change is affected by a trophically transmitted cestode using brine shrimps as an intermediate host and Black‐necked Grebes as final host, was not supported by the data.     

Do captive waterfowl alter their behaviour patterns during their flightless period of moult?

Many different behavioural changes have been observed in wild waterfowl during the flightless stage of wing moult with birds frequently becoming inactive and reducing time spent foraging. Increased predation risk, elevated energetic demands of feather re-growth and restriction of foraging opportunities are thought to underlie these changes. By studying captive populations of both a dabbling and a diving duck species at the same site, we determined whether captive birds would reflect the behavioural responses of wild waterfowl to moult. The time-budgets of 42 Common Eiders, Somateria mollissima, (a diving duck) and 18 Garganeys, Anas querquedula, (a dabbling duck) were recorded during wing moult (July–August) and non-moult (January) with behaviour recorded under six categories. Despite captivity providing a low predation risk and constant access to food, birds altered their behaviour during the flightless period of wing moult. Time allocated to foraging and locomotion decreased significantly during moult compared to non-moult periods, while resting time increased significantly. Moulting Eiders underwent a greater reduction in time spent foraging and in locomotion compared with Garganeys, which is likely to be in response to a higher energetic cost of foraging in Eiders. It is possible that increased resting in both diving and dabbling ducks reduces their likelihood of detection by predators, while allowing them to remain vigilant. We demonstrate that there is much potential for using captive animals in studies that can augment our knowledge of behaviours of free-living conspecifics, the former being a hitherto under-exploited resource.     

Do Seaducks Minimise the Flightless Period?: Inter- and Intra-Specific Comparisons of Remigial Moult

Remigial moult is one of the crucial events in the annual life cycle of waterfowl as it is energetically costly, lasts several weeks, and is a period of high vulnerability due to flightlessness. In waterfowl, remigial moult can be considered as an energy-predation trade-off, meaning that heavier individuals would minimise the flightless period by increasing feather growth rate and energy expenditure. Alternatively, they could reduce body mass at the end of this period, thereby reducing wing-loading to increase flight capability. We studied timing of remigial moult, primary growth rates, flightlessness duration, and the pattern of body mass variation in 5 species of captive seaducks (Melanitta fusca, M. perspicillata, Clangula hyemalis, Histrionicus histrionicus, and Somateria mollissima) ranging in size from 0.5 to 2.0 kg. Their feather growth rates weakly increased with body mass (M^0.059) and no correlation was found at the intra-specific level. Consequently, heavier seaduck species and especially heavier individuals had a longer flightless period. Although birds had access to food ad libidum, body mass first increased then decreased, the latter coinciding with maximum feather growth rate. Level of body mass when birds regained flight ability was similar to level observed at the beginning of remigial moult, suggesting they were not using a strategic reduction of body mass to reduce the flightlessness duration. We suggest that the moulting strategy of seaducks may be the result of a compromise between using an intense moult strategy (simultaneous moult) and a low feather growth rate without prejudice to feather quality. Despite the controlled captive status of the studied seaducks, all five species as well as both sexes within each species showed timing of moult reflecting that of wild birds, suggesting there is a genetic component acting to shape moult timing within wild birds.     

Greater energy stores enable flightless moulting geese to increase resting behaviour

Many species of waterfowl undergo a post‐breeding simultaneous flight feather moult (wing moult) which renders them flightless and vulnerable to predation for up to 4 weeks. Here we present an analysis of the correlations between individual time‐budgets and body mass states in 13 captive Barnacle Geese Branta leucopsis throughout an entire wing moult. The daily percentage of time spent resting was positively correlated with initial body mass at the start of wing moult. Behaviour of individual birds during wing moult is dependent on initial physiological state, which may in turn be dependent on foraging ability; the storage of energy before the start of wing moult will help birds to reduce exposure to the dangers of predation.     

Seasonal hyperphagia does not reduce digestive efficiency in an Arctic grazer

Muskoxen (Ovibos moschatus) consume fibrous plants that grow rapidly over the short Arctic summer. We studied responses of eight castrated male muskoxen to a diet of grass hay and mineral supplements during spring, autumn, and winter. Animals gained body mass in spring (239+/-39 kg) as body fat content increased from 26% to 38% of ingesta-free mass in winter without changes in lean mass and protein. Intakes of dry matter (DM) increased by 74% between spring and autumn as digestible energy increased from 554 to 923 kJ kg(-0.75) d(-1) during mass gain. Digestibility of cellulose (72%-76%) was not affected by increasing food intake between spring and autumn but was reduced to 65% in winter. Digestibility of nitrogen compounds was 61%-66%, even though intake increased by 134% between spring and autumn. Excess dietary nitrogen from hay and supplements increased urea concentrations in plasma and urine. High loads of solutes such as potassium did not affect plasma or urinary osmolality but were associated with increased rates of glomerular filtration and urinary excretion. Low intakes of sodium from grasses may limit intake and digestion during summer, but high food intakes can support deposition of nitrogen, calcium, magnesium, copper, and zinc in body tissue even when dietary concentrations are low. Seasonal increases in digestive and metabolic functions allow muskoxen to rapidly accumulate energy and nutrients in body tissue during the short season of plant growth.     

Evidence of differences in growth and food intake regulation in different genetic strains of channel catfish

Norris and USDA-103 strains of channel catfish Ictalurus punctatus were compared for growth rate and food conversion ratio under satiation feeding and restricted feeding (1% body weight day⁻¹) regimes. At the start of the experiment Norris fish weighed 2·8 g, USDA-103 fish weighed 14·0 g. Therefore, a regression of the loge of specific growth rate against the log_e of mean body size with an empirically derived fixed slope of -0·37 was used to compare growth rates. Under both feeding regimes the USDA-103 strain had faster specific growth rates and more efficient food conversion. In subsequent studies, voluntary food intake of size matched fish (60 g average) from these two strains was compared using a radiographic method. Fish were acclimatized to tank conditions for 3 weeks prior to voluntary food intake measurement. Half of the groups were deprived of food for 2 days prior to food intake measurement, while the remaining groups were fed 1% body weight day⁻¹. The USDA-103 strain fish ate significantly more food and grew faster than the Norris strain fish. Previously fasted Norris fish subsequently ate more than their fed counterparts, whereas the fed USDA-103 fish consumed more food than the fasted USDA-103 group. When the USDA-103 strain fish were deprived of food for 4 , 2 or 0 days, all groups subsequently consumed between 4·5 and 5·0% of body weight in one meal. The USDA-103 fish, unlike the Norris fish were not stimulated to consume more after short-duration fasting. Taken together, these results suggest that there are genetic differences in growth, food conversion ratio and regulation of food intake between Norris and USDA-103 strains.     

Species‐specific habitat use of wing‐moulting waterbirds in response to temporary flightlessness

The choice of the moulting habitat is of paramount importance for wing‐moulting waterbirds that have to cope with a flightless period of several weeks. However, some species might have more restricted habitat requirements during moult than others, for example due to a highly specialized feeding ecology. The moult‐related habitat use of five species (Gadwall Anas strepera, Red‐crested Pochard Netta rufina, Common Pochard Aythya ferina, Tufted Duck Aythya fuligula, Coot Fulica atra) was compared at a European inland moulting site that offered a variety of water bodies characterized by different levels of nutrient concentration, water depth, shoreline vegetation density and disturbance. To determine location‐ and species‐specific densities, birds were regularly counted throughout the moulting seasons of 2010 and 2011. In 2011, additional data on Gadwalls were used to assess differences in requirements between the flightless phase of moult and the periods before and after. Furthermore, habitat choice of 38 tagged Gadwalls was compared among two to four successive years. During the moulting season, all species showed clear preferences for specific levels of nutrient content, suggesting an active choice of suitable food sources in both food specialists and generalists. Species showing the strongest attachment to shallow water (Gadwall and Coot) were most sensitive to human disturbance and increasing water depths, and species averse to diving (Gadwall) used ponds with dense shore vegetation while flightless. For Gadwalls, habitat conditions rather than nutrient supply became increasingly important during the flightless phase. Average return rates of 59 and 54% were recorded for male and female Gadwalls, respectively, and the repeated use of familiar locations could be demonstrated in the majority of returning birds (65%). Familiarity with the habitat apparently plays an important role and may enable individuals to compensate for suboptimal conditions at the moulting site.     

Phenotypic flexibility of a southern African duck Alopochen aegyptiaca during moult: do northern hemisphere paradigms apply?

Phenotypic flexibility during moult has never been explored in austral nomadic ducks. We investigated whether the body condition, organ (pectoral muscle, gizzard, liver and heart) mass and flight‐feather growth Egyptian geese Alopochen aegyptiaca in southern Africa show phenotypic flexibility over their 53‐day period of flightless moult. Changes in body mass and condition were examined in Egyptian geese caught at Barberspan and Strandfontein in South Africa. Mean daily change in primary feather length was calculated for moulting geese and birds were dissected for pectoral muscle and internal organ assessment. Mean body mass and condition varied significantly during moult. Body mass and condition started to decrease soon after flight feathers were dropped and continued to do so until the new feathers were at least two‐thirds grown, after which birds started to regain body mass and condition. Non‐moulting geese had large pectoral muscles, accounting for at least 26% of total body mass. Once moult started, pectoral muscle mass decreased and continued to do so until the flight feathers were at least one‐third grown, after which pectoral muscle mass started to increase. The regeneration of pectoral muscles during moult started before birds started to gain overall body mass. Gizzard mass started to increase soon after the onset of moult, reaching a maximum when the flight feathers were two‐thirds grown, after which gizzard mass again decreased. Liver mass increased significantly as moult progressed, but heart mass remained constant throughout moult. Flight feather growth was initially rapid, but slowed towards the completion of moult. Our results show that Egyptian geese exhibit a significant level of phenotypic flexibility when they moult. We interpret the phenotypic changes that we observed as an adaptive strategy to minimize the duration of the flightless period. Moulting Egyptian geese in South Africa undergo more substantial phenotypic changes than those reported for ducks in the northern hemisphere.     

The bioenergetics of grass carp, Ctenopharyngodon idella (Val.): the influence of body weight, ration and dietary composition on nitrogenous excretion

Nitrogenous excretion by grass carp, Ctenopharyngodon idella (Val.), was measured in the form of ammonia and urea. Endogenous nitrogen excretion (ENE) was estimated as the daily rate of excretion by grass carp which had been starved for 2 days. ENE was scaled allometrically with body weight with weight exponents of 0.75 for ammonia, 0.63 for total nitrogen and 0.63 for the energy lost. The proportion of nitrogen attributable to urea was smaller than that attributable to ammonia and decreased from 25 to 12% as fish weight increased from 2 to over 10 g.
Linear relationships were found between daily rates of ammonia, total nitrogen and energy loss and daily rates of food intake. High carbohydrate and high lipid diets were not shown to have a protein-sparing action compared to a high protein diet. Differences in the amount of nitrogen excreted were explained by the differing nitrogen contents of the diets. Nitrogen budgets were erected and their implications discussed.     

SOME NOTES ON THE BIRDS OF THE ISOKA DISTRICT OF THE NORTHERN PROVINCE OF NORTHERN RHODESIA

Dean, W. R. J. 1978. Moult seasons of some Anatidae in the western Transvaal. Ostrich 49:76-84.

Spurwinged Geese Plectropterus gambensis, Egyptian Geese Alopochen aegyptiacus, Yellow-billed Ducks Anas undulata, Redbilled Teal A. erythrorhyncha and Southern Pochard Netta erythrophthalma have a flightless moult mainly during the dry season, from April to August, in the western Transvaal. South African Shelduck Tadorna cana moult during October to February after breeding during July and August. The Cape Shoveller Anas smithii has two main flightless periods, April-May and October-January. Cape Teal A. capensis have been recorded in flightless moult in October, December and January.

The duration of the flightless period correlates with wing length; larger and longer winged Anatidae require proportionally more time for wing moult than do smaller and shorter winged Anatidae.

Geese and shelducks moult on large open lakes with an open shore. Ducks have been recorded flightless on lakes and dams, with or without emergent vegetation.     

Can grazing sheep compensate for a daily foraging time constraint?

1. Theoretical studies of large herbivore foraging assume that total daily grazing time is a key constraint on daily intake and diet choice. We experimentally tested this assumption and investigated the effects of food availability on the ability of grazing sheep to compensate for restriction of available daily grazing time.
2. Foraging behaviour, intake and diet digestibility by sheep, were measured on grass pastures in a replicated 2 × 2 factorial experiment, in which overnight access to pasture was varied (restricted overnight and continuous access) on two sward heights (5·5 and 3·0 cm), representing high and low food availability.
3. Regardless of food availability, the overnight-restricted sheep fed for almost all of the available grazing time by grazing for fewer, longer foraging bouts, but still had much shorter total daily grazing time than the continuous access sheep.
4. In response to overnight penning, the sheep had a significantly higher instantaneous rate of intake achieved mainly via larger bites. The continuous access sheep were hence not maximizing their short-term rate of intake, whilst grazing according to the daily schedule considered normal for sheep.
5. The behavioural responses to overnight food restriction were able to counteract the reduction in daily grazing time only where food availability was high. In contrast on short swards overnight grazing restriction led to a reduction in total daily intake. We suggest that the interactions between the factors considered as constraints on foraging behaviour of herbivores are, as yet, only poorly quantified.     

Fecundity, lifespan and egg mass in butterflies: effects of male-derived nutrients and female size

1. Effects of larval reserves and nutrients received as adults on fecundity and lifespan in female Danaus plexippus (the Monarch Butterfly) were measured to determine the relative importance of different sources of nutrients for reproduction and somatic maintenance.
2. Egg-laying lifespan was correlated with female size but not with the amount of male-derived nutrients or adult food concentration.
3. Lifetime fecundity was higher when females received a large first spermatophore, but was not affected by female size when lifespan was controlled or by adult food concentration.
4. At the end of their lives, females contained unlaid eggs and retained, on average, 88% of their initial mass. This proportion was unchanged in two years, although mean egg-laying lifespan varied from 22·5 to 28·7 days.
5. Egg mass decreased over the female lifespan, and was correlated with female size.
6. These results suggest that larval reserves are more important for somatic maintenance than adult income, but that the protein-rich nutrients received from males contribute to egg production. This supports theoretical predictions and empirical studies of other Lepidoptera showing that larval reserves are less likely to affect fecundity when the adult income can contribute substantially to egg production.     

Repeated grazing of a salt marsh grass by moulting greylag geese Anser anser– does sequential harvesting optimise biomass or protein gain?

The effects of simulated goose grazing on common saltmarsh-grass Puccinellia maritima plants were tested on a Danish salt marsh during the flightless moulting period of greylag geese Anser anser (3–21 June 1998). Plants in an area exclosed from the influence of grazing and the nutrient effects of goose faeces were subject to removal of youngest lamina at 3-, 6-, 9- and 18-day intervals during this period. Average biomass and protein accumulation between harvests was highest at defoliation intervals of 9 days or more. Field observations from two separate study areas demonstrated geese returned to regraze the Puccinellia sward after 6–8 days and oesophageal contents from feeding geese showed selection for lamina lengths consistent with the results of clipping every 6 days. Geese therefore regrazed Puccinellia patches at shorter intervals than expected were they to maximise their intake of biomass or protein at each visit. However, total cumulative lamina elongation, equivalent to the long term gain during the entire moult period, showed no significant difference between the three most intensive defoliation treatments, which were significantly greater than those of plants defoliated at 18 day intervals. Highest overall lamina protein levels were maintained at 6- and 9-day defoliation intervals. This suggests geese regrazed Puccinellia patches at a rate that maximised their number of harvests during the flightless period, but maintained highest protein levels and overall biomass in the sward. This suggests, in line with earlier studies, that moulting greylag geese combine dietary selection, reduced nitrogen excretion and regrazing patterns to meet protein demands during regrowth of flight feathers.