Optimal diving behaviour for foraging in relation to body size

Overall, large animals dive longer and deeper than small animals; however, after the difference in body size is taken into account, smaller divers often tend to make relatively longer dives. Neither physiological nor theoretical explanations have been provided for this paradox. This paper develops an optimal foraging diving model to demonstrate the effect of body size on diving behaviour, and discusses optimal diving behaviour in relation to body size. The general features of the results are: (1) smaller divers should rely more heavily on anaerobic respiration, (2) larger divers should not always make longer dives than smaller divers, and (3) an optimal body size exists for each diving depth. These results explain the relatively greater diving ability observed in smaller divers, and suggest that if the vertical distribution of prey in the water column is patchy, there is opportunity for a population of diving animals to occupy habitat niches related to body size.     

Trade-offs between social behaviour and foraging by sheep in heterogeneous pastures

In heterogeneous pastures, groups of sheep may have to alter their social behaviour in order to graze patches of preferred vegetation. In this event, patch size, inter-patch distance and the contrast between patch and background vegetation are likely to affect behaviour. In this experiment, groups of five female Scottish Blackface sheep grazed for 2-h periods in 0.1ha grass plots containing seven 1.5mx1.5m patches of improved vegetation, with inter-patch distances of 1.5, 6 or 11.5m. Background vegetation was of either medium or poor quality. Control plots contained no patches. On average, sheep spent 44% of the time grazing patches, although patches comprised only 1.6% of the total plot area. Inter-patch distance did not affect accumulated time spent grazing patches during the first 30min, but patch residence time and the number of sheep on a patch increased with inter-patch distance. The distribution of nearest neighbour distances was altered when patches were 6 or 11.5m apart, compared to no patches. Accumulated time spent grazing patches and the number of sheep on a patch were greater with poor than medium backgrounds. Sheep visited patches frequently and for short periods and it is suggested that sheep often moved off patches as a result of competition. The results provide evidence that sheep make trade-offs between social and foraging behaviour and demonstrate the importance of interactions between social spacing and the size and spacing of vegetation patches, when sheep forage in heterogeneous pastures.     

Diving behaviour in relation to water temperature in the southern elephant seal: foraging implications

Summary A time-depth-temperature recorder provided a continuous record of diving by a female southern elephant seal in relation to water temperature for 27 days (1939 dives) after completion of moult. Mean maximum dive depth was 391±2.6 m and the overall maximum was 775 m. Dives lasted on average 17.5±0.09 min. Most dives showed a rapid descent to the discontinuity between the cold surface water and warmer deep water. Consequently the seal spent 57% of its time while diving at a depth of 200–400 m when it may have been foraging. This strongly suggests that the seal was exploiting a food source at the discontinuity between vertically stratified water masses. The water temperature data also indicated that the seal was diving in waters south of the Antarctic Polar Front and at some distance from the northern edge of the pack ice. The seal spent 88% of its time under water. Normal surface intervals between dives lasted an average of 2.1 ± 0.1 min whereas 16 extended surface intervals (>10 min duration) lasted 32.7±4.6 min. Dives were deeper during the day than at night and all but one extended surface interval occurred at night. The pattern of dives was similar to records from northern elephant seals but this is the first study to show how diving behaviour relates to water temperature.     

The relationship between foraging behaviour and energy expenditure in Antarctic fur seals

By using time-depth recorders to measure diving activity and the doubly-labelled water method to determine energy expenditure, the relationship between foraging behaviour and energy expenditure was investigated in nine Antarctic fur seal females rearing pups. At-sea metabolic rate (MR) (mean of 6.34 ± 0.4 W. kg^-1; 4.6 times predicted BMR) was positively correlated to foraging trip duration (mean of 4.21 ± 0.54 days; r²= 0.5, P < 0.04). There were no relationships between MR and the total number of dives, the total time spent diving or the total vertical distance travelled during the foraging trip. There was, however, a close negative sigmoidal relationship (r²= 0.93) between at-sea MR and the proportion of time at sea spent diving. This measure of diving behaviour may provide a useful, inexpensive means of estimating foraging energy expenditure in this species and possibly in other otariids. The rate of diving (m.h^-1) was also negatively related to at-sea MR (r²= 0.69, P < 0.005). Body mass gain during a foraging trip had a positive relationship to the time spent at sea (r²= 0.58, P < 0.02) and the total amount of energy expended while at sea (r²= 0.72, P < 0.004) such that, while females undertaking long trips have higher metabolic rates, the energetic efficiency with which females gain mass is independent of the time spent at sea. Therefore, within the range of conditions observed, there is no apparent energetic advantage for females in undertaking foraging trips of any particular duration.     

Food and foraging behaviour of the Snares fernbird

Abstract

The Snares fernbird (Bowdleria punctata caudata Buller, 1894) is an opportunistic insectivore, its prey ranging greatly in size and type. Birds forage over a wide variety of sites, including forest, tussock, boulder beaches, cliffs, penguin colonies, and floating kelp. Foraging methods vary with the prey sought, and at times birds adopt specific “search images” when hunting certain insects. Details of foods brought to nestlings are given. Four postures used by fernbirds when foraging actively are described.     

Optimizing foraging behaviour through learning

Manifestation of life-history strategy is through the allocation of resources acquired by foraging. Foraging efficiency can be improved by learning, as fishes adjust their behaviour to changing circumstances. We briefly review the influence of learning on the foraging behaviour of fishes and make recommendations for further research. We stress the importance of quantifying learning and memory in relation to ontogeny and life history.     

Body temperature and foraging behaviour of the Eurasian otter (Lutra lutra), in relation to water temperature

The effects of water temperature (T_w) on core body temperature (T_b) and foraging behaviour were studied in four free-ranging and one captive otter Lutra lutra in NE Scotland. The free-ranging animals were observed in and around two freshwater lochs, and measurements were made of T_w (2–16°C), T_b at the beginning and end of swimming bouts, lengths of swimming bouts, and dive times and intervals. T_b ranged from 35.9 to 40.4°C (mean 38.1°C). At the beginning of a period of activity and of a swimming bout. T_b rose significantly; during swimming it fell at a rate of 2.3°C/h, independent of T_w. Dive intervals were longer in colder waters, but this was probably due to other, seasonal environmental changes (perhaps water level). T_w had no significant effect on length of swimming bouts or on T_b. Although otters entered water with increased T_b, they exited when T_b was not significantly different from the mean resting T_b. The data suggest that otters maintain T_b irrespective of T_w, by increasing it before entering water, then either exiting at a given T_b, or at an earlier time determined by other environmental conditions.     

Pandora neoaphidis transmission and aphid foraging behaviour

Pandora neoaphidis is an aphid-specific entomopathogen that produces infective conidia. As aphid movement increases, so does the likelihood of contact with conidia. Volatile distress signals released in response to aphid infestation as an indirect defence against herbivory may affect aphid foraging and, therefore, the fungus-aphid interaction. In this study, two different methods were used to investigate the effect of plant volatiles and P. neoaphidis-sporulating cadavers on (1) the colonisation of Vicia faba plants by Acyrthosiphon pisum and (2) P. neoaphidis transmission. This study indicates that A. pisum does not avoid bean plants containing P. neoaphidis and that transmission of conidia occurs during plant colonisation and, to a lesser extent, during in situ feeding. Although significantly more aphids were recovered from damaged plants compared to undamaged plants, the likelihood of infection was not affected by previous infestation by aphids.     

Morphology and foraging behaviour of Siberian Phylloscopus warblers

There are seven species of Phylloscopus warblers breeding sympatrically in Central Siberia. In general they are very similar in morphology and behaviour. Slight differences in morphology, however, are often interpreted as having arisen through adaptation to distinct foraging ecologies, thereby mediating species coexistence. We studied the morphology and foraging behaviour of these warblers, taking into account phylogenetic relationships using Felsenstein's method of independent contrasts. Striking correlations were found between morphology and foraging techniques, as well as between morphology and microhabitat selection. Species with large hind limbs and short wings foraged in dense vegetation near the ground using gleaning techniques. In contrast, species with long wings and large bills favoured open vegetation, and foraged using sallying manoeuvres. Independently of these trends, small species foraged at higher levels in the vegetation, and used hoverflight more often than did larger species. Foraging methods were highly correlated with vegetation structure. Coniferous trees were exploited using hoverflight when they had short needles and gleaning when needles were long. Sallying predominated in light deciduous vegetation whereas gleaning was used most in dense, deciduous bushes. After controlling for the effects of vegetation structure on foraging behaviour, species differences became less pronounced, but were still significant. It is therefore concluded that morphology may be regarded as having adapted to enable more efficient exploitation of certain microhabitats. A preference for these microhabitats then further increases the observed behavioural differences between species. Morphology may therefore partly determine a species' ability to colonise new habitats.     

Territorial and foraging behaviour of two corallivorous butterflyfishes

The territorial and foraging behaviour of two Red Sea butterflyfish species was studied at three sites in the Gulf of Aqaba. Chaetodon austricaus , a generalist corallivore that exploited evenly distributed food resources, maintained exclusive pair territories. Individuals fed by grazing for brief periods on a coral colony before moving to the next. In contrast, C. trifascialis , a specialist corallivore that exploited patchily distributed food resources, demonstrated considerable variation in territorial behaviour that ranged from the defence of exclusive solitary territories, to the shared use of a large home range. Individuals showed highly variable feeding behaviour, from grazing analogous to C. austriacus , to continual use of a single large coral colony. The results of this study demonstrate how the use of space by individuals depends critically upon the distribution of key food resources. In addition, the results demonstrate that foraging behaviour of territorial butterflyfish can be quite plastic, and adapt to local conditions.     

Food and foraging behaviour of the lizard,Ctenotus taeniolatus

A sample of 261 Ctenotus taeniolatus revealed that this species of skink is principally insectivorous, the most common foods being lepidopteran and coleopteran larvae, orthopterans and formicids. The occurrence of these foods in the diet followed seasonal patterns. The prey of adults and juveniles did not differ qualitatively, although adults were capable of eating a greater diversity of prey sizes than juveniles. Lizards used both sit-and-wait and active foraging strategies, with adults and juveniles exhibiting these behaviours in different ratios.     

The relation between morphology and behaviour during ontogenetic and evolutionary changes

During development, form and function (behaviour) change while the match between them must be maintained. The quality of this match determines the importance of morphological parameters in constraining behaviour. If the match is close, the morphology of organisms will be more constraining to the behaviour than when there is a large reserve capacity that creates a certain flexibility. This leads to two questions: (1) How good is the match between form and function during development? The quality of the match necessarily changes during development because changes in structural capacity often cannot proceed at the same speed as changes in functional demand. The evidence for these changes is discussed. (2) What are the mechanisms that maintain the match between form and function during developmental and evolutionary changes? Two mechanisms for maintaining the match are discussed: (a) reserve capacity and (b) flexible muscle activity patterns. Special emphasis is given to fish examples throughout this review.     

Understanding foraging behaviour in spatially heterogeneous environments

The role of stochasticity and spatial heterogeneity in foraging systems is investigated. We formulate a spatially explicit model which describes the behaviour of grazing animals in response to local information using simple stochastic rules. In particular the model reflects the biology in that decisions to move to a new location are based on visual assessment of the sward height in a surrounding neighbourhood, whilst the decision to graze the current location is based on the residual sward height and olfactory assessment of local faecal contamination. It is assumed that animals do not interact directly, but do so through modification of, and response to a common environment. Spatial heterogeneity is shown to have significant effects including reducing the equilibrium intake rate and increasing the optimal stocking density, and must therefore be taken into account by resource managers. We demonstrate the relationship between the stochastic spatial model and its non-spatial deterministic counterpart, and in the process derive a moment-closure approximation to the full process, which can be regarded as an intermediate, or pseudo-spatial model. The role of spatial heterogeneity is emphasized, and better understood by comparing the results obtained from each approach. The relative efficiency of random and directed searching behaviour in spatially heterogeneous environments is explored for both clean and contaminated pastures, and the impact of faecal avoidance behaviour assessed.     

Status-dependent foraging behaviour in coral reef wrasses

Field observations using underwater video were used to reveal differences in the duration and frequency that fish engaged in daily behaviours such as chasing, searching, feeding, and travelling, according to their social patterns [passive or aggressive terminal phases (TPs), pair-spawning, or promiscuous groups] and intraspecific status (terminal or initial colour phases). Marked differences were apparent according to status, but this was not consistent among the three species; TP Cheilinus fasciatus tended to swim in longer bouts with less frequent searching or feeding than initial phase (IP) conspecifics; TP Cirrhilabrus punctatus exhibited less frequent feeding and travelling than IP conspecifics, while the most aggressive species Stethojulis bandanensis displayed no significant intraspecific differences. We highlight the importance of social context and individual status when examining fish foraging activities and the utility of underwater video for recording the duration and frequency that fish engage in essential daily activities.     

Learning in the nectar foraging behaviour of Helicoverpa armigera

Abstract .1. Learning may enable insects to obtain nectar from flowers more efficiently. Learning in nectar foraging has been shown primarily in studies of bees and butterflies. Here, learning is demonstrated in the nectar foraging behaviour of a noctuid moth, Helicoverpa armigera .
2. The present studies show that: (1) previous experience with a flowering host species increases the probability of that species being selected for nectar foraging, and (2) previous experience of a particular flower type (food source at bottom or top of the corolla tube) increases the likelihood of the food source being found when that flower type is being searched.
3. The implications of these findings for understanding the pattern of oviposition observed in wild populations of this important pest species are discussed.