Stomach temperature telemetry reveals temporal patterns of foraging success in a free-ranging marine mammal

1. We studied feeding frequency in free-ranging grey seals using stomach temperature telemetry to test if previously reported sex differences in the diving, movement and diet were reflected in the temporal pattern of foraging success. 2. Data were retrieved from 21 of 32 grey seals from 1999 to 2001, totalling 343 days and 555 feeding events, with individual record length varying from 2 to 40 days (mean: 16.33 +/- 2.67 days/seal). 3. Seals fed on 57.8 +/- 6.46% of days sampled and had an average of 1.7 +/- 0.26 meals per day, but individual variability was apparent in the temporal distribution of feeding as evidenced by high coefficients of variation (coefficient of variation = 69.0%). 4. Bout analysis of non-feeding intervals of six grey seals suggests that feeding intervals of individuals were varied and probably reflect differences in prey availability. Grey seals tended to have many single feeding events with long periods separating each event, as would be expected for a large carnivore with a batch-reactor digestive system. 5. We found significant sex differences in the temporal distribution of feeding. The number of feeding events per day was greater in males (2.2 +/- 0.4 vs. 1.0 +/- 0.2), as was time associated with feeding per day (56.6 +/- 5.8 min vs. 43.9 +/- 9.4 min). 6. The number of feeding events varied with time of day with the least number occurring during dawn. Feeding event size differed significantly by time of day, with greater meal sizes during the dawn and the smallest meals during the night. 7. The length of time between meals increased with the size of the previous meal, and was significantly less in males (541.4 +/- 63.5 min) than in females (1092.6 +/- 169.9 min). 8. These results provide new insight into the basis of sex differences in diving and diet in this large size-dimorphic marine predator.     

A novel flexible method to split feeding behaviour into bouts

Before meal patterns can be analysed properly, a biologically relevant meal criterion must be determined in order to group short-term feeding behaviour into meals. Existing methodologies are based on modelling of the frequency distributions of intervals between feeding events but these methods cannot be used if the proper distributions cannot be clearly identified. For such cases we developed two new methods – (1) based on the analysis of the distribution of between-meal interval lengths only and (2) based on the analysis of changes in the probability of animals starting to feed with time since the last feeding event. Both methods were developed using a data set of over 700,000 records of visits to feeders obtained with broilers (Gallus gallus) aged between 2 and 5 weeks. The two methods resulted in meal criteria estimates of 20.1 and 17.5 min, respectively, which, when applied to the data set, gave statistically significant but very small differences in meal characteristics. The new methods were tested against an independent cow (Bos taurus) data set and the resultant meal criteria compared with those predicted by an existing method. The two novel methods estimated meal criteria for cows at 27.9 and 35.5 min, compared with 28.9 min for the existing method. Again, these differences in meal criteria resulted in only very small differences in meal characteristics. Even though meal criteria were relatively similar for birds and cows, characteristics such as average daily number of meals (10.9 and 5.9), meal size (12.5 g and 7.4 kg) and meal duration (7.7 and 31.4 min) were very different. The analyses show that, if the appropriate distributions of intervals cannot be identified, meal criteria can still be estimated for species as diverse as mature cows and young birds by the novel methodologies developed here.     

Patterns in feeding: a behavioural analysis using Locusta migratoria nymphs

ABSTRACT. Feeding patterns of Locusta migratoria L. nymphs, kept under LD 12:12 at 30°C with ample food, were analysed over the first 6 days of the fifth instar. A number of differences were found between the light and dark phases. (1) More was eaten during the light than the dark, with the differences becoming progressively larger over the 6 days. (2) During light phases most insects showed an increasing tendency to feed with time since the last meal, while in the dark the probability of starting to feed remained constant. (3) There was little variation in feeding patterns within the light phase, but marked changes occurred within the dark phase, feeding declining to a low level in the last 4h. The amount eaten during a meal was regulated similarly in the light and dark, with time rather than amount eaten being of paramount importance in terminating a meal. The time when feeding started was somewhat influenced by the size of the preceding meal, but once feeding began meal size was largely independent of the preceding meal. The importance of food quality in determining the pattern of feeding is discussed.     

The structure of feeding behavior in a phytophagous insect (Hylobius abietis)

Analysis of the feeding behavior of animals using such a high temporal resolution that meals can be defined may improve our understanding of the mechanisms regulating feeding. Meals can be distinguished in an ethologically meaningful manner by using the ‘meal criterion’, the shortest non‐feeding interval between feeding bouts recognized as meals. However, such a criterion has only been determined for a few insect species. Applying a recent method developed for assessing meal criteria for vertebrates, we determined the meal criterion for Hylobius abietis (L.) (Coleoptera: Curculionidae) based on data from video recordings of single individuals feeding on seedlings of Norway spruce, Picea abies (L.) Karst. (Pinaceae). The pine weevil is an economically important pest insect, because it feeds on the stem bark of planted conifer seedlings. Weevils had 4–5 meals per day. Each meal lasted about 24 min during which about 13 mm² of bark per meal were removed. Females had longer total meal durations and longer non‐feeding intervals within meals than males. Girdling seedlings did not affect the weevils' feeding properties. The size of meals was significantly correlated with the duration of non‐feeding intervals before and after them. This study is one of few describing the feeding behavior of an insect at a temporal resolution that allows individual meals to be distinguished. With more meal‐related data from insects available, differences in meal properties may be interpreted based on phylogeny, ecology, and physiology. Our results may also assist in the setup and interpretation of studies of plant‐insect interactions, and facilitate the evaluation and development of methods to protect plants against herbivores.     

The influence of meal size on prey DNA detectability in piscivorous birds

Molecular methods allow noninvasive assessment of vertebrate predator–prey systems at high taxonomic resolution by examining dietary samples such as faeces and pellets. To facilitate the interpretation of field‐derived data, feeding trials, investigating the impacts of biological, methodological and environmental factors on prey DNA detection, have been conducted. The effect of meal size, however, has not yet been explicitly considered for vertebrate consumers. Moreover, different noninvasively obtained sample types remain to be compared in such experiments. Here, we present a feeding trial on abundant piscivorous birds, Great Cormorants (Phalacrocorax carbo), to assess meal size effects on postfeeding prey DNA detection success. Faeces and pellets were sampled twice a day after the feed of large (350–540 g), medium (190–345 g) and small (15–170 g) fish meals contributing either a large (>79%) or small (<38%) share to the daily consumption. Samples were examined for prey DNA and fish hard parts. Molecular analysis of faeces revealed that both large meal size and share had a significantly positive effect on prey DNA detection rate postfeeding. Furthermore, large meals were detectable for a significantly longer time span with a detection limit at ~76 hr and a 50% detection probability at ~32 hr postfeeding. In pellets, molecular methods reliably identified the meal consumed the previous day, which was not possible via morphological analysis or when examining individual faeces. The less reliable prey DNA detection of small meals or meal shares in faeces signifies the importance of large numbers of dietary samples to obtain reliable trophic data.     

Infection of the malaria mosquito Anopheles gambiae with the entomopathogenic fungus Metarhizium anisopliae reduces blood feeding and fecundity

The entomopathogenic fungus Metarhizium anisopliae is being considered as a biocontrol agent against adult African malaria vectors. In addition to causing significant mortality, this pathogen is known to cause reductions in feeding and fecundity in a range of insects. In the present study we investigated whether infection with M. anisopliae affected blood feeding and fecundity of adult female malaria vectors Anopheles gambiae Giles sensu stricto. Mosquitoes were contaminated with either a low or a moderately high dose of oil-formulated conidia of M. anisopliae, and offered a single human blood meal 48, 72, or 96 h later to assess feeding propensity and individual blood meal size. In a second experiment, individual fungus-infected females were offered a blood meal every third day (to a total of 8 gonotrophic cycles), and allowed to oviposit after each cycle in order to quantify feeding propensity and fecundity. Infected females took smaller blood meals and displayed reduced feeding propensity. It was found that mosquitoes, inoculated with a moderately high dose of fungal conidia, exhibited reduced appetite related to increasing fungal growth. Of the fungus-infected females, the proportion of mosquitoes taking the second blood meal was reduced with 51%. This was further reduced to 35.3% by the 4th blood meal. During 8 feeding opportunities, the average number of blood meals taken by uninfected females was 4.39, against 3.40 (low dose), and 2.07 (high dose) blood meals for the fungus-infected females. Moreover, infected females produced fewer eggs per gonotrophic cycle and had a lower life-time fecundity. Epidemiological models show that both blood feeding and fecundity are among the most important factors affecting the likelihood of a mosquito transmitting malaria, which suggests that this fungus may have potential as biocontrol agent for vector-borne disease control.     

Feeding behaviour of artificially reared Romane lambs

A consequence of increasing litter size in sheep is that a portion of the lambs have to be reared artificially. Detailed information about the pattern of milk consumption of artificially reared lambs would help improve their management. The purpose of this study is to describe the individual and group feeding behaviour of 94 Romane artificially reared lambs from 5 to 28 days of age using an electronic automatic lamb feeder. Animals were located in four pens of 8 to 15 lambs of similar age with one teat per pen. They were fed ad libitum. In our experimental situation (group rearing, continuous lightning) on average a lamb made 1.4±0.7 visits to the teat per meal and 9.5±3 meals per day. Mean meal duration was 247±158 s and the mean daily time spent feeding was 38±25 min. The mean quantity of milk intake was 176±132 ml per meal and 1.68±0.8 l per day. With age, the number of daily meals and their duration decreased while the quantity of milk consumed per meal and per day increased. Females tended to make more visits to the teat per meal and perform more meals per day but their milk consumption per meal was lower. The feed conversion ratio was 1.36±0.2. Synchrony in feeding (group meal) was estimated as the percentage of lambs that wanted to access the teat within the same short period (relative group meal size). On average 65% of lambs in the pen wanted to access the teat within the same period, but for 35% of group meals the relative group meal size was >90%. There was no consistency in the order in which lambs accessed the teat during a group meal. Our evaluation suggested that electronic automatic lamb feeders are tools that can provide, on a large scale, data describing the feeding behaviour of artificially reared lambs. It is then possible to study factors influencing these traits in order to improve the outcome of artificially reared lambs.     

The inhibitory effects of the molluscicide metaldehyde on feeding, locomotion and faecal elimination of three pest species of terrestrial slug

Single meals on maize flour pellets containing 0, 0·5, 1, 2, 4, 6, and 8% metaldehyde were analysed in Arion hortensis (agg), Deroceras reticulatum and Deroceras caruanae, by attaching the pellets to a mechanical transducer from which signals were ultimately fed into a BBC microcomputer for on-line analysis. Post-feeding tests were used to investigate toxic effects of metaldehyde and the recovery of slugs was examined daily for 7 days. Bite rates were slightly lower and more irregular on metaldehyde meals especially in A. hortensis. The meal length and number of bites per meal were markedly inhibited by metaldehyde and inversely proportional to the concentration. The size of the bites was also reduced, by about 30%, in meals on 6% pellets compared with 0%. Clear symptoms of poisoning became evident during the meal or normally in the 16-min test period immediately afterwards. In these tests slugs which fed on metaldehyde pellets showed a disruption of their ability to right themselves (after being placed on their side), less activity and a reduced probability of accepting a ‘dummy’ pellet containing no metaldehyde. It is hypothesised that feeding is prematurely terminated primarily because of the toxic effects of the metaldehyde and that aversion also acts to terminate the meal early, by decreasing the probability of continuing to feed as the meal proceeds. At higher concentrations of metaldehyde aversion may be more important than the toxic effects in ending the meal. Faecal elimination is delayed after metaldehyde meals and this contributes to mortality. The estimated lethal doses were about 0·2 mg/g body weight of slug in Deroceras spp. A. hortensis was less susceptible. To increase the likelihood of a lethal dose being ingested the inhibition of feeding needs to be overcome, at least partly, and the results presented indicate the need for a bait formulation which reduces the rate of absorption of the molluscicide from the gut.     

Can one unrestricted meal buffer the effects of previous pre-meal intervals on the feeding behaviour of sheep?

Varying the time since the last meal is one means of manipulating feeding motivation. In order to use this method effectively it is necessary to know whether and the extent to which effects of one pre-meal interval are carried over to affect the behaviour during the following meals. Pre-meal interval (PMI) is defined here for practical purposes, for short meals, as the time between the start of two successive meals. The possibility that one unrestricted meal might buffer the effects of an 8h as opposed to a 4h PMI on aspects of feeding behaviour was studied with eight Scottish Blackface sheep. They were fed on a regime in which they were given access to food until they finished their meal and lay down (this always occurred within 60min) at which time the remaining food was withdrawn. Feeding behaviour was recorded during the meal after these 4 and 8h intervals, as well as during the following meal 4h later.At a meal after a PMI of 8h, compared to 4h, sheep had a higher intake per meal (mean+/-S.T.D. for 8 and 4h PMI, respectively: 604.4+/-78.8 and 430.1+/-100.9g; P<0.001), a longer meal duration (27.1+/-7.5 and 21.8+/-8.1min; P<0.001), and a tendency for a higher intake rate (23.8+/-6.2 and 21.9+/-8.2g/min; P=0.11). During the following meal 4h later these differences were smaller, but intake per meal still tended to be higher (430.8+/-81.5 and 338.5+/-45.6g; P<0.06) for sheep who had previously had the 8h PMI. Meal duration (21.9+/-7.2 and 20.6+/-7.08min; P=0.28) and intake rate (21.2+/-6.1 and 18.7+/-7.2g/min; P=0.13) were no longer different.A single meal after the different PMIs reduced differences in all three aspects of feeding behaviour observed during the subsequent meal, 4h later, but differences in intake per meal were still apparent. It is suggested that an additional meal may overcome the carry-over effect.     

A cost-benefit analysis of feeding in female tsetse

Abstract. Three models for feeding in female tsetse are considered. Model I: there is a prolonged non-feeding phase after each meal followed by feeding at a constant rate, with a constant probability of dying as a consequence of feeding. Model II: the feeding rate increases linearly after each meal. Model III: the feeding rate increases exponentially after each meal. In Models II and III the feeding hazard is a linear function of the probability of feeding. Production of viable female offspring is estimated under each model, making allowance for losses of adults due to starvation and to background and feeding mortality, losses of pupae due to predation and parasitization, and losses of young flies if their mothers take insufficient blood during pregnancy. Under Model I, if females require three meals to produce viable pupae in 9 days, then for a non-decreasing population with a background mortality of 1%/day, and 25% pupal losses due to predation and parasitism, the feeding risk must be ≤5%/feed. At this maximum level the non-feeding phase should be 2–2.5 days for optimal productivity, with a mean feeding interval of 60–72 h. If the background mortality is 2%/day, feeding losses cannot exceed 1%/feed for a non-decreasing population. If four or five meals are required for the production of fully viable pupae, the optimal values of the non-feeding phase and mean feeding interval tend towards 1 and 2 days respectively. Under Models II and HI the mean feeding interval is 50–60 h for optimal productivity (with variances 3 times as large as for Model I), in good agreement with estimates from recent models for feeding and digestion. Field evidence suggests that feeding tsetse take greater risks as their fat levels dwindle. This should result in feeding (and feeding mortality) rates which increase during the feeding phase - as assumed in Models II and III but not in Model I. These models allow greater flexibility than Model I, because flies can feed early in the hunger cycle, at low probability, as long as the feeding risk is also low.     

川楝素对斜纹夜蛾幼虫取食行为的影响

斜纹夜蛾5龄幼虫的取食活动是非连续进行的。在每两次取食活动之间存在一个取食间间隔．在每次取食活动期间,也存在一非常短暂的取食内间隔。经川楝素处理后的斜纹夜蛾5龄幼虫．其21小时内发生的取食活动和取食间间隔的次数减少．平均每次取食活动和取食间间隔持续的时间延长;可是在每次取食活动中,用于取食的时间明显减少。这是因为．取食内间隔的次数增加了．取食内间隔的时间也延长了。所有上述指标,处理虫与对照之间均存在着显著或极显著的差异。     

Measuring meals: structure of prandial food and water intake of rats

Attempts to understand ingestion have sought to understand the control of meals. The present study evaluated a meal definition that included prandial drinking (drinking-explicit meals). The spontaneous nocturnal intake of male Wistar rats was studied. The meal breakpoint was defined as the interval between feeding or drinking events providing the most stable estimate of meal structure. Alternative breakpoints derived from prevailing methodology, log-survivorship, or frequency histogram analysis of interfeeding intervals without respect to drinking were compared (drinking-naive meals). Threshold interfeeding intervals that accounted for drinking indirectly were evaluated as surrogate breakpoints (drinking-implicit meals). Definitions were compared by determining which criterion better conformed to predictions of satiety. Microstructural differences resulting from the definitions were also studied. Under the drinking-explicit definition, rats averaged nine or ten 13-min meals/night, during which they consumed food and water equally in duration (5-6 min) and quantity (2.3 g). Individual differences were observed in microstructure measures. Meals defined by drinking-informed, but not drinking-naive, methods were followed by the behavioral satiety sequence and by an initially low likelihood of resuming feeding that monotonically increased with time. The drinking-explicit definition uniquely revealed preprandial and postprandial correlations of similar magnitude. Under drinking-informed definitions, food restriction increased meal size, whereas drinking-naive definitions increased meal frequency. Drinking-implicit definitions provided workable approximations of meal frequency and size but inferior estimates of feeding duration, eating rate, and the preprandial correlation. Thus log-survivorship analysis is not appropriate for identifying meal breakpoints, and the consideration of drinking in meal definitions can provide a better estimate of meal structure.     

Individual rainbow trout can learn and anticipate multiple daily feeding times

The present study was conducted to investigate whether individual rainbow trout Oncorhynchus mykiss can simultaneously anticipate two daily meals, and to evaluate the influence of time-restricted feeding schedules and light-dark (L:D) cycles on the timing of self-feeding activity. Individual rainbow trout were capable of anticipating not only a single meal but also two daily meals scheduled at different times; the duration of anticipation was short and precise. This seems to be the first convincing evidence to show that a single fish can simultaneously anticipate two daily meals. Change in the onset and cessation of light did not affect the duration of food-anticipatory activity, although it led to shift the phases of the light-related self-feeding activity. Moreover, under constant light and ad lib feeding conditions, only one band of rhythmic activity was observed in individual rainbow trout that had previously developed food-anticipatory activity to each of the two daily meals.     

Rates of gastric evacuation in piscivorous brown trout, Salmo trutta

SUMMARY. 1. The dry weight of food remaining in the stomachs of piscivorous trout decreased exponentially with time. Gastric evacuation rates increased exponentially with increasing temperature but were unaffected by predator size, meal size or type of fish prey.
2. Mathematical models were developed to estimate both the rate and time for the gastric evacuation of different meal sizes (expressed as dry weight), and were applicable to piscivorous trout of different sizes (length range 10–32 cm) feeding on trout fry or sticklebacks at different temperatures (range 5–18°C).
3. The wet weight of food in the stomachs also decreased exponentially with time, but evacuation rates both increased with temperature and decreased with increasing meal size; the latter relationship occurred because relative rates of water loss from a meal also decreased with increasing meal size. Use of wet or dry weights can therefore lead to different conclusions about the effect of meal size on evacuation rates.
4. When piscivorous trout were fed three consecutive meals of varying size, the models predicted the total dry weight of food left in the stomach, but not the weight remaining for each individual meal. Interactions between meals led to an increase in evacuation rates for meals consumed early in the series and a decrease in evacuation rates for later meals.
5. Evacuation rates for piscivorous trout were compared with those for trout feeding on invertebrates in an earlier study, and were close to those for caddis larvae as prey, higher than those for mealworms and lower than those for a variety of invertebrate prey. Although a great deal is now known about the daily food intake and growth rates of trout feeding on invertebrates, there is little comparable information for piscivorous trout.     

The structure of feeding behavior in the colorado potato beetle,Leptinotarsa decemlineata (Coleoptera: Chrysomelidae)

Feeding behavior, in an ad libitum situation on potato plants in the laboratory, was continuously observed for approximately 7 h/day on 2 successive days for 18 adult femaleLeptinotarsa decemlineata. Additional behaviors were also recorded including resting, walking, biting, local movements, grooming, defecating, and regurgitating. These data were used to calculate a time budget for the various behaviors. The feeding data were analyzed to describe the structure of feeding for young adult females on their normal host plant. The criterion for a meal (minimum intermeal interval) was determined to be 286 s. This criterion was used to distinguish between intra- and intermeal interruptions in feeding for all subsequent analyses. Meals taken from leaves that were young, medium aged, or old did not differ, but on average beetles took 60% of their meals from young leaves. Meal size and meal duration were equally good predictors of when a meal would end. Feeding from stems was a prominent feature for most beetles. The stem meals had much longer durations than leaf meals, but stem feeding did not affect subsequent leaf feeding. The structure of feeding by these beetles is compared with that found in other insects, especiallyLocusta migratoria.     

Increasing sodium bicarbonate level in high-concentrate diets for heifers. II. Effects on chewing and feeding behaviors

Four Holstein heifers (264 ± 12 kg initial BW) were used in a 4 × 4 Latin square design with 21-day experimental periods to determine the effect of increasing levels of sodium bicarbonate (BICARB) (0%, 1.25%, 2.5% and 5%, of concentrate dry matter (DM) basis) on chewing and feed intake behavior when fed high-concentrate diets. Concentrate (13.41% CP, 13.35% NDF) and barley straw were fed once a day at 0830 h ad libitum. Feed bunks placed on scales and video recording were used to measure 24-h feed intake and chewing behavior, respectively. The patterns of feeding behavior (feed intake, meal size and length) and chewing behavior (eating, ruminating and total chewing) were studied by dividing the day into 12 intervals of 2-h each, beginning at feeding (interval 1 through 12). Number of meals per day and eating rate decreased linearly with increasing buffer level, but meal length increased linearly. No treatment effects were observed in sum of daily meal lengths or average meal size. The treatment × interval interaction was significant on meal size, length and feed intake. The size and length of those meals occurring during the 4 h post-feeding increased linearly. However, meal size tended to decrease in the evening between 8 and 12 h, whereas feed intake decreased linearly from 6 to 10 h and from 12 to 14 h post-feeding. Buffer concentration did not affect the percentage of time spent ruminating, eating or drinking per day but the buffer level × interval interaction was significant. Time spent eating expressed as min per kg of DM or organic matter (OM) intake increased linearly with buffer levels. Proportion of time spent eating increased linearly during the intervals between 0 and 4 h post-feeding. Time spent ruminating decreased linearly during the 2 h post-feeding, and also in the evening from 12 to 14 h, and at night from 18 to 22 h post-feeding, but the effect was quadratic between 8 and 10 h when intermediate buffer levels showed the greatest ruminating time. Time spent drinking decreased linearly from 6 to 8 h but increased during the 2 h following feeding and from 10 to 12 h post-feeding. Daily eating rate and meal frequency decreased linearly as the buffer level increased, but average meal size and daily chewing times were not affected. However, significant time of the day × buffer level interactions were observed for feed intake, meal size and length and chewing behavior.     

Influence of adult breeding experience on growth and provisioning of Wandering Albatross Diomedea exulans chicks at South Georgia

For most seabirds, reproductive performance improves with age; in albatrosses this is thought not to be so (experience being acquired before starting breeding) but only one study (of chick growth in a single season at one site) has specifically addressed this. We compared the provisioning performance and growth rates of chicks of Wandering Albatrosses Diomedea exulans breeding for the first (IN), second and third (LE) and fourth or more times (EE) on Bird Island, South Georgia in the austral winters of 1996 and 1997. Eggs from EE adults were significantly heavier than the other two categories and these chicks had a greater mass and longer wings up to 160 days of age and longer culmen and tarsus up to 115 days old. However chicks from all categories fledged at the same average mass, size and age. No significant differences between categories in feeding frequency or meal size were detected but experienced adults made shorter long foraging trips and spent more time at the nest than less experienced birds. Adults that remained at the nest gave chicks smaller meals than those that left immediately after feeding the chick. Although provision of smaller but more frequent meals by experienced adults promotes more rapid chick growth, the resulting differences do not persist into the late chick-rearing period. Our results were very similar to those from Iles Crozet in the Indian Ocean, supporting the hypothesis that when Wandering Albatrosses start to breed they are fully competent foragers but that it takes a while, during early chick-rearing, for birds breeding for the first time to adapt to the additional demands of provisioning a chick.     

Individual feeding specialisation in shorebirds: population consequences and conservation implications

Individual feeding specialisation in shorebirds is reviewed, and the possilble mechanisms involved in such specialisations. Any specialisation can he seen as an individual strategy, and the optimum strategy for any given individual will be conditional upon its specific priorities and constraints. Some specialisations are related to social status and some to individual skills. Some are also probably frequency-dependent. However, most shorebird specialisations are constrained to a large extent by individual morphology, particularly bill morphology. For example, larger birds are able to handle larger prey, and birds with longer bills are able to feed on more deeply buried prey. Sex differences in bill length are uncommon in the Charardriidae, which are surface peckers, but are common in the Scolopacidae, which feed by probing in soft substrates. Sex differences in bill morphology are frequently associated with sex differences in feeding specialisation. There is evidence that different feeding specialisations are associated with different payoffs, in which case the probability of failing to reproduce or of dying will not be distributed equally throughout the population. I consider the population consequences of such feeding specialisations, particularly the different risks and benefits associated with different habitats or diets. I also consider the way in which individuals may differ in their response to habitat loss or change. I suggest that population models designed to predict the effect of habitat loss or change on shorebirds should have the ability to investigate the differential response of certain sections of the population, particularly different ages or sexes, that specialise in different diets or feeding methods.     

Individual differences in feeding efficiencies and feeding preferences of captive great tits

The preferences of eight, adult, hand-reared great tits for feeding on four different types of feeding sites have been tested. There were significant differences between individuals in these preferences, and the preferences for the different objects were correlated with the birds' individual feeding efficiencies on the objects. This correlation was thought to be a result of the birds' experience and learning during training for the experiment. During the tests the birds tended to make runs of visits to the same object type. This effect was more marked for less-preferred objects, and when the birds had just made a successful visit.