The effect of time and digestion constraints in Common Eiders while feeding and diving over Blue Mussel beds

1. To maintain energy intake Common Eiders ( Somateria mollissima ) in winter should compensate for reduced day-length by increasing both the proportion of time spent feeding and diving efficiency, defined as the proportional duration of a dive bout within a dive cycle. Common Eiders swallowed Blue Mussel ( Mytilus edulis ) whole with their shells where any behavioural compensation in relation to short days may be limited by digestive processes.
2. Based on time budget studies conducted from mid-December to the end of April in the Gulf of St. Lawrence, Québec, Canada, diving and feeding efficiency were compared for three seasonal periods varying in day-length (from 557 to 890 min).
3. Results showed that eiders were compensating for short days by feeding 56% of the time in mid-winter compared with 33% in spring. However, diving efficiency remained constant through the season and apparently no compensation occurred at this level of their foraging behaviour. Despite this, the daily rate of prey ingestion was much higher in mid-winter than in spring.
4. Ingestion rate values for mid-winter individuals approached or even exceeded the rate at which prey items are defecated and it was concluded that shell crushing performed by the muscular gizzard is physiologically more demanding during that period. On this basis, gizzard mass should be larger in winter when ingestion rates are higher. Data presented support that hypothesis and suggest that compensation in relation to short days can be both behavioural and physiological.     

State-dependent incubation behaviour in the high arctic barnacle geese

Energetic trade-offs between time spent on incubation and times spent on foraging for nesting birds give individuals in good body condition the possibility to incubate more continuously. In the present paper, the incubation behaviour of female barnacle geese Branta leucopsis was quantified in a colony in Svalbard. Females were weighted in early incubation and feeding recess lengths and frequencies were recorded. The feeding behaviour during the course of incubation was significantly correlated to by body mass, and heavy females had both fewer and shorter feeding recesses than lighter females. Moreover, there was an increase in the number of feeding recesses and the total feeding time as the incubation period progressed. Neither clutch size nor egg laying date had an effect on incubation behaviour. However, clutch size was positively related to female body mass suggesting that high-quality females produce large clutches but also allocate more body reserves to incubation. Females left the nest to feed at all times of the day, but more frequently during day time. This was not related to their body mass. Females presumably leave their nest at the time of day when the costs to reheat eggs are at a minimum. The diurnal rhythm may also be adjusted to the activity by egg predators. Overall the results support the state-dependent hypothesis for incubation behaviour, suggesting that body condition at the start of incubation is an important factor for incubation behaviour in barnacle geese.     

Feeding ecology of barnacle geese on their spring staging grounds in northern Iceland

The distribution and abundance of barnacle geese, Branta leucopsis. staging in northern Iceland was determined in May 1987 and 1994 A similar proportion of the total population (70%) was located in the two years The distribution differed between the two years, with a higher proportion of the geese found in more westerly sites in 1994 Over 80% of records of birds feeding were on improved agricultural pasture though this made up only 47% of the available feeding habitat Some geese fed on riverine and coastal grasslands but this was in a much smaller proportion than the availability of these habitats The number of geese using a site was positively correlated with the area of improved pasture Birds fed intensively through the day, suggesting that the area is important for the accumulation of nutrient stores No differences in habitat use or feeding behaviour were found between 1987 and 1994     

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.     

Travel schedules to the high arctic: barnacle geese trade-off the timing of migration with accumulation of fat deposits

On their way from the wintering area to the breeding grounds in Spitsbergen, barnacle geese Branta leucopsis stage on islands off the coast of Norway. The aim of this study was to describe when the geese migrate in relation to the body stores deposited and explore questions related to the concept of optimal migration schedules and on the possible mechanisms involved. We estimated fat stores by repeated assessments of the abdominal profile index of individually marked females throughout staging. Reproductive success was derived from observations of the same individuals later in the annual cycle. Females arriving late, or with low fat stores at arrival, achieved higher fat deposition rates, probably by spending more time foraging. But they were unable to match final fat scores of birds that arrived earlier or with larger fat stores. Reproductive success was correlated with the timing of migration and individuals departing at intermediate dates achieved highest success. The exact date of peak reproductive success depended on the size of fat stores accumulated, such that low-quality birds (depositing less fat) benefited most from an early departure to the breeding grounds. Observations in the breeding colonies showed that these birds did not initiate a nest earlier but they spent a longer time in Spitsbergen before settling. The length of stay in Norway was close to the prediction derived from an optimisation model relating spring events to eventual breeding success. Poorest performing birds stayed longer than expected, perhaps depositing more fat to avoid the risk of starvation. Two possible mechanisms of the timing of migration were contrasted and it seemed that the geese departed for migration as soon as they were unable to accumulate any more fat stores.     

Sward height and bite size affect the functional response of barnacle geese Branta leucopsis

Intake rate, the rate in which herbivores can process their food, is presumed to be an important factor in habitat selection down to the scale of the foraging patch. Much attention has been given to the selection of swards of high nutritional quality, but much less has been given to the influences of sward structure on patch selection in small herbivores. In this study we tested the effects of sward density and height on the functional foraging response of barnacle geese, Branta leucopsis. The functional response curve for herbivores describes how intake rate is affected by food availability. We conducted feeding trials to determine intake rate and bite size of barnacle geese on experimentally manipulated swards. Results indicate that intake rate is mainly dependent on sward height and that there is a strong correlation between bite size and intake rate. Sward density does not influence the rate of food consumption; it is, however, a crucial parameter affecting potential total yield. We conclude that bite size is the crucial parameter influencing intake rate. Bite size is explained both by sward height and individual differences in bill morphology. Furthermore, intake rate seems to be dependent on the physical structure of the grass species consumed.     

Lack of competition between barnacle geese Branta leucopsis and pink-footed geese Anser brachyrhynchus during the pre-breeding period in Svalbard

The feeding ecology of barnacle geese and pink-footed geese was studied in Sassendalen, Svalbard during the pre-nesting period (late May) to assess the potential for inter-specific competition. Barnacle geese fed almost exclusively (97%) by grazing above-ground plant material, mostly (79%) along snow edges in moss-mat habitats. Pink-footed geese fed mostly (93%) by excavating below-ground parts of plants, mostly (56%) away from snow and were more evenly distributed between habitat types. Barnacle goose faeces contained mostly (62%) moss, that of pink-footed geese mostly (48%) below-ground plant storage organs (especially Bistorta viviparum L.). Principal components analysis of dropping contents showed no overlap in species diet in allopatry or sympatry. There was little overlap in diet and feeding ecology of the two species at this pre-nesting feeding site. Hence, unless increased goose feeding densities affect future vegetation density and composition, under present circumstances, increasing numbers of either species is unlikely to affect foraging conditions for the other at this important stage in the annual cycle. However, such changes could have local density-dependent intra-specific effects.     

Patch Use in Free‐Ranging Goats: Does a Large Mammalian Herbivore Forage like Other Central Place Foragers?

Classic central place foraging theory does not focus on the foraging of central place herbivores. This is especially true with regard to large mammalian herbivores. To understand the foraging dynamics of these neglected foragers, we measured giving‐up densities (GUDs) in artificial food patches. We did this at different distances away from the central point (i.e. corral) for a herd of free‐ranging domestic goats. To determine temporal changes, we conducted the study over a 3‐mo period during an extended dry season. Throughout our study, goats foraged across a gradient of food availability where forage was more available farther away from the central point. In contrast to the prediction that predation risk and/or increased travel costs were the main drivers of foraging decisions, we found that the goats increased their feeding effort (i.e. achieved lower GUDs) the farther away they moved from the central point. This suggests that either metabolic or missed opportunity costs were the main factors that influenced foraging decisions. In addition, we suggest that social foraging may have also played a role. With increases in foraging opportunities away from the central point, a herd will likely move slowly while foraging. As a result, individuals can feed intensively from patches but remain part of the group. Ironically, owing to the sustained close proximity of other group members, individuals may perceive patches farther from the central point as being safer. Temporally, the goats increased their feeding effort throughout the dry season. This suggests there was a decline in food quality and/or availability across the environment as the study progressed. Despite this increase in feeding effort, the negative relationship with distance did not change. Ultimately, our results provide key insight into how metabolic, missed opportunity and perceived predation costs influence the feeding decisions of large central place herbivores.     

Patch choice of avian herbivores along a migration trajectory–From Temperate to Arctic

Migratory waterfowl species seem to track temporal and spatial pulses of optimal forage availability on their way from temperate wintering to arctic breeding sites. In order to unravel the relative contribution of forage quality and forage biomass to foraging choices in avian herbivores, we experimentally manipulated biomass and quality of main forage plants through fertilisation and grazing exclusion at three sites along the flyway of barnacle geese, Branta leucopsis. Fertilisation increased the nitrogen content of the forage and grazing exclusion increased biomass levels. Manipulated plots were offered to wild geese in a random block experimental design and goose visitation was measured through dropping counts. At all sites there was a trend towards a higher preference of plots with increased quality and average biomass above plots with an average quality and increased biomass. Generally, geese preferred plots with highest standing crop of nitrogen. The numerical response of the geese to forage changes was supported by behavioural observations at the Baltic site. We conclude that for migrating barnacle geese the bottlenecks in the standing crop of nitrogen appear to lie in the limited biomass availability at the Baltic stopover site and the limited nutrient content of food in the Arctic breeding site, restricting the potential nutrient intake on these sites.     

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.     

Resting metabolic rate in migratory and non‐migratory geese following range expansion: go south,go low

While many species suffer from human activities, some like geese benefit and may show range expansions. In some cases geese (partially) gave up migration and started breeding at wintering and stopover grounds. Range expansion may be facilitated and accompanied by physiological changes, especially when associated with changes in migratory behaviour. Interspecific comparisons found that migratory tendency is associated with a higher basal or resting metabolic rate (RMR). We compared RMR of individuals belonging to a migratory and a sedentary colony of barnacle geese Branta leucopsis. The migratory colony is situated in the traditional arctic breeding grounds (Russia), whereas the sedentary colony has recently been established in the now shared wintering area (the Netherlands). We measured RMR by oxygen consumption () during two ontogenetic phases (juvenile growth and adult wing moult). We also investigated juvenile growth rates and adult body mass dynamics. Mass‐independent was 13.6% lower in goslings from the sedentary colony than in goslings from the migratory colony. Similarly, in adult geese, mass‐independent was 15.5% lower in sedentary than in migratory conspecifics. Goslings in the Netherlands grew 36.2% slower than goslings in Russia, while we found no differences in body dimensions in adults. Adult geese from both colonies commenced wing moult with similar body stores, but whereas Russian barnacle geese maintained this level throughout moult, body stores in geese from the Netherlands fell, being 8.5% lower half‐way through the moult. We propose that the colony differences in resting metabolic rate, growth rate and body mass dynamics during moult can be explained by environmental and behavioural differences. The less stringent time constraints combined with poorer foraging opportunities allow for a smaller ‘metabolic machinery’ in non‐migratory geese. Our analysis suggests that range expansion may be associated with changes in physiology, especially when paired with changes in migratory tendency.     

Sharing habitat: Effects of migratory barnacle geese density on meadow breeding waders

• 1.Following targeted conservation actions most goose populations have increased. The growing goose populations caused an increase in human-wildlife conflicts and have the potential to affect nature values. As meadow birds, including meadow-breeding waders, were declining throughout Western Europe, the possible negative effect of rising numbers of foraging barnacle geese on their breeding success has been questioned.
• 2.We used GPS-transmitter data to measure the density of foraging barnacle geese during daylight hours. Using dynamic Brownian Bridge Movement Models (dBBMM), we investigated the effect of barnacle goose density on the territory distribution of five wader species, and on nest success of the locally common Northern lapwing. We used model selection methods to identify the importance of barnacle goose density related to other environmental factors.
• 3.Our results showed an insignificant positive association between barnacle goose density and nest territory density of the Northern lapwing and common redshank. Barnacle goose density had no influence on territory selection of godwit, oystercatcher and ringed plover. We did, however, find a negative correlation between barnacle geese density and the nest success of the Northern Lapwing.
• 4.We infer that either barnacle goose foraging leads to improved territory conditions for some wader species, or that both barnacle geese and waders prefer the same type of habitat for foraging and nesting. Higher barnacle goose density was correlated with fewer Northern lapwing nests being successful.
• 5.Synthesis and application: Experimental research is needed to disentangle the causal chain, but based on our observational findings, we suggest to increase water logging that may attract both barnacle geese and wader species. Further investigation on the effects of barnacle geese on wader species is necessary to identify the cause of the negative correlation between barnacle geese density and nest success of lapwings; nest protection experiments could give further insight.

     

Goose grazing selectivity along a depletion gradient

The micro-habilat food selectivity along a gradient of increasing grazing intensity of wintering pink-footed geese Anser brachyrhynchus and barnacle geese Branta leucopsis foraging on winter wheat Triticum aestivum and smooth meadow grass Poa pratensis was investigated. Protein content of ungrazed primary and secondary leaves of winter wheat did not differ, whereas the protein content of tertiary leaves was significantly lower. Secondary leaves were significantly longer than both primary and tertiary leaves, whereas tertiary leaves were significantly longer than primary leaves. Protein content of primary and secondary leaves of Poa pratensis did not differ, whereas the protein content of tertiary leaves was significantly lower. Primary leaves were significantly shorter than both secondary and tertiary leaves, which did not differ significantly.
For all leaf categories, there was an increase in the removal with increasing goose grazing intensity. The geese showed a sequential selection of winter wheat leaves according to the protein content, initially preferring primary and secondary leaves, but increasingly selecting tertiary leaves of poorer quality as high-quality leaves became depleted. Foraging on Poa pratensis. geese selected secondary leaves according to the protein content, whereas tertiary leaves were preferred to primary leaves, although these had a protein content equal to secondary leaves. This, however, may be explained by the primary leaves being too short to maintain intake rates.
It is concluded that feeding preferences cannot be established on the basis of protein contents of food plants alone and that green biomass estimates not necessarily reflect resource availability.     

Temporal Allocation of Foraging Effort in Female Australian Fur Seals (Arctocephalus pusillus doriferus)

Across an individual''s life, foraging decisions will be affected by multiple intrinsic and extrinsic drivers that act at differing timescales. This study aimed to assess how female Australian fur seals allocated foraging effort and the behavioural changes used to achieve this at three temporal scales: within a day, across a foraging trip and across the final six months of the lactation period. Foraging effort peaked during daylight hours (57% of time diving) with lulls in activity just prior to and after daylight. Dive duration reduced across the day (196 s to 168 s) but this was compensated for by an increase in the vertical travel rate (1500–1600 m·h⁻¹) and a reduction in postdive duration (111–90 s). This suggests physiological constraints (digestive costs) or prey availability may be limiting mean dive durations as a day progresses. During short trips (<2.9 d), effort remained steady at 55% of time diving, whereas, on long trips (>2.9 d) effort increased up to 2–3 d and then decreased. Dive duration decreased at the same rate in short and long trips, respectively, before stabilising (long trips) between 4–5 d. Suggesting that the same processes (digestive costs or prey availability) working at the daily scale may also be present across a trip. Across the lactation period, foraging effort, dive duration and vertical travel rate increased until August, before beginning to decrease. This suggests that as the nutritional demands of the suckling pup and developing foetus increase, female effort increases to accommodate this, providing insight into the potential constraints of maternal investment in this species.     

Behavioral responses of Barbary macaques (Macaca sylvanus) to variations in environmental conditions in Algeria

In this study, the behavioral responses of Barbary macaques to seasonal and interhabitat variations in resource availability were analyzed over an entire annual cycle. Two groups, one in an evergreen cedar–oak forest (Djurdjura) and the other in a deciduous oak forest (Akfadou), were observed. In this paper, references to data on resource availability published elsewhere are made. Time budget has been studied. Variations in foraging and moving time, in day–range lengths, and in time moving in trees have been considered to estimate the variations in foraging effort and thus energy expenditure. Great monthly variations in foraging effort and other activities were observed in both habitats. In early spring, when resource availabilities were maximal, foraging effort was low while monkeys maximized their feeding time (about 5 h/day). In June, during the peak of the birth season and the rearing period, monkeys minimized their feeding time to the benefit of social interactions (to 1.6–2.7 h/day), whatever the food availability, which was low in Akfadou and high in Djurdjura. In addition, foraging effort remained low in Djurdjura, while it increased in Akfadou. Thus, at the beginning of the dry summer period, monkeys in Akfadou were in a less favorable position than those in Djurdjura. At both sites, in periods of food shortage in summer or in winter, monkeys displayed two different strategies. In the former case, their foraging effort increased, while in the second one it remained relatively low. Whatever the foraging effort, monkeys did not reach the same amount of feeding time as in early spring. In the poorest site of Akfadou, foraging effort was globally greater than in the richest site of Djurdjura, especially for adults. At both sites, adult males spent more time feeding than juveniles and less time in social interactions. Results are discussed according to rearing period, temperatures, and day length constraints. The limits of adaptability to different habitats are considered in light of the demographic parameters. Am. J. Primatol. 43:285–304, 1997. © 1997 Wiley-Liss, Inc.     

Lateralization in monogamous pairs:wild geese prefer to keep their partner in the left hemifield except when disturbed

Behavioural lateralization, which reflects the functional specializations of the two brain hemispheres, is assumed to play an important role in cooperative intraspecific interactions. However, there are few studies focused on the lateralization in cooperative behaviours of individuals, especially in a natural setting. In the present study, we investigated lateralized spatial interactions between the partners in life-long monogamous pairs. The male-female pairs of two geese species (barnacle, Branta leucopsis, and white-fronted, Anser albifrons geese), were observed during different stages of the annual cycle in a variety of conditions. In geese flocks, we recorded which visual hemifield (left/right) the following partner used to monitor the leading partner relevant to the type of behaviour and the disturbance factors. In a significant majority of pairs, the following bird viewed the leading partner with the left eye during routine behaviours such as resting and feeding in undisturbed conditions. This behavioural lateralization, implicating the right hemisphere processing, was consistent across the different aggregation sites and years of the study. In contrast, no significant bias was found in a variety of geese behaviours associated with enhanced disturbance (when alert on water, flying or fleeing away when disturbed, feeding during the hunting period, in urban area feeding and during moulting). We hypothesize that the increased demands for right hemisphere processing to deal with stressful and emergency situations may interfere with the manifestation of lateralization in social interactions.     

Foraging time and dietary intake by breeding Ross's and Lesser Snow Geese

We compared foraging times of female Ross's (Chen rossii) and Lesser Snow Geese (Chen caerulescens caerulescens) breeding at Karrak Lake, NT, Canada and examined variation due to time of day and reproductive stage. We subsequently collected female geese that had foraged for known duration and we estimated mass of foods consumed during foraging bouts. Female Ross's Geese spent more time foraging (mean % - SE =28.4ǃ.3%; P=0.0002), on average, than did female Lesser Snow Geese (21.5 - 1.4%). Foraging time by female geese differed among reproductive stages, but differences were not consistent among time periods (stage-by-time block interaction, P=0.0003). Females spent considerably more time foraging during prelaying and laying than during incubation. Ross's Geese also spent a greater percent of time feeding (83.0DŽ.8%) during incubation recesses than did Lesser Snow Geese (60.9Dž.6%). Consumption of organic matter during foraging bouts was minimal; estimated consumption averaged 9.6dž.0 and 12.4dž.6 g (mean - SE) dry mass/day before incubation and 5.9DŽ.0 and 5.7DŽ.1 g dry mass/day during incubation for Lesser Snow and Ross's Geese, respectively. Diets consisted primarily of mosses (bryophytes), Chickweed (Stellaria spp.) and Sedges (Carex spp.). Before incubation, eggshell consumption was estimated as 4.3Dž.2 and 0.4ǂ.3 g dry mass/day for Lesser Snow and Ross's Geese, respectively; neither species consumed eggshell during incubation. We conclude that eggshell from nests of previous years is likely an important source of dietary calcium used to meet mineral demands of eggshell formation at Karrak Lake. Our findings of wide disparities between foraging time and food intake indicate that results from studies that do not directly measure intake rates remain equivocal. Finally, we propose four hypotheses accounting for foraging effort that evidently yields little nutritional or energetic benefit to geese nesting at Karrak Lake.     

Expensive Traditions: Energy Expenditure of Aleutian Geese in Traditional and Recently Colonized Habitats

ABSTRACT In an effort to reduce goose depredation at a traditional spring migratory stopover site, private landowners implemented a coordinated hazing plan to scare Aleutian cackling geese (Branta hutchinsii leucopareia) from private lands to adjacent public pastures that were cultivated and set aside specifically for geese. Coincidentally, some Aleutian geese began using a new stopover site 150 km farther south in their spring migratory range; numbers at the new site continue to increase. We tested the idea that when their ability to acquire resources deteriorates geese are likely to seek improved foraging conditions, especially during spring migration when individuals strive to maximize nutrient stores and minimize energy expenditure. We quantified measures of goose foraging performance in traditional and new spring staging sites by calculating foraging opportunity, foraging effort, body condition, and daily energy expenditure. Geese staging at the site with higher levels of human disturbance had less foraging opportunity and, despite increased foraging effort and more nutritious food-plants at the site, birds there experienced an elevated energy expenditure and poorer body condition than birds at the new stopover site. Reduced foraging time and increased energy expenditure at the traditional spring staging site may have triggered the colonization process. Suitability assessment of habitat for migratory geese should include measures of foraging opportunity, disturbance risks, and daily energy expenditure in addition to quantity and quality of foods.     

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.     

Providing water for goats in arid landscapes: effects on feeding effort with regard to time period, herd size and secondary compounds

In arid regions, herbivores contend with a wide range of variables that influence their foraging ability. These may include plant secondary compounds (e.g. tannins and oxalates), water availability, time of day, and herd size. To determine the relative importance of these variables for goats living in a semi-desert, we measured the remaining food resources after foraging events (giving up densities-GUDs) as an index of foraging effort in artificial food patches. Time of day and the availability of drinking water had the greatest effect on GUDs. Goats achieved lower GUDs (i.e. ate more) in the afternoon than they did during the morning. We suggest this was due to missed opportunity costs of future foraging opportunities being lower in the afternoon. Ultimately, this implies that goats made short-term foraging decisions based on estimates of future feeding opportunities. When water was available, feeding effort increased and GUDs dropped almost 50%. For goats living in an arid environment, food and water are strong complementary resources. The presence of low to moderate levels of tannic or oxalic acid in food was a slight deterrent, raising the GUDs similarly. Larger herds ate more and thus obtained lower GUDs. In addition, there was an interaction of herd size and water, suggesting that in arid environments, the effect of water intensifies with population density. Our approach allowed us to quantify and rank the effects that disparate environmental factors had on the goats' foraging costs: water>time of day>herd size>plant secondary compounds. These findings ultimately demonstrate how the provision of water may increase grazing impacts by herbivores in arid landscapes.