Foraging rhythms in Adélie penguins (Pygoscelis adeliae) at hope bay,Antarctica; determination and control

Summary We studied diel periodicity in activity of Adélie penguins (Pygoscelis adeliae) during the austral summer solstice and one month later by counting, hourly, numbers of birds leaving colonies to forage, numbers arriving back after foraging, numbers in a colony and numbers resting outside the colonies. During the solstice large numbers of birds were arriving at and departing from the colony at all times of the day although there was a tendency for more birds to be at sea when light intensity was highest at mid-day. Generally, birds not brooding chicks did not rest on land. A month later, when visibility was poor at mid-night, the percentage of birds at sea was highly positively correlated with light intensity. Birds returning from foraging in the evening fed chicks immediately and then either rested in the colonies or on snow patches between the colonies and the sea until the following morning.     

Temperature dependence of feeding behaviour in the mopane worm (Lepidoptera)

We related foraging activity by the caterpillars of the mopane emperor moth (Imbrasia belina) to temperature in the laboratory and then used these data to predict foraging activity from temperature in the field. Predictions were poor when based on shade temperature, better when based on operative temperature, but always over-estimated foraging activity. We suggest that temperature is not the only factor affecting foraging activity by I. belina the field and that other factors, especially predation pressure, also have a strong influence.     

The influence of daily variation in foraging cost on the activity of small carnivores

The daily activity of some predators is correlated with the activity pattern of their prey. If capture efficiency varies as a function of prey activity, a predator that synchronizes its foraging activity with the time of day that prey are most vulnerable should capture more prey, and at lower cost, than a predator that initiates foraging at random. Mink, Mustela vison, and weasels, M. erminea and M. nivalis, were presented with an opportunity to maximize their intake in similar circumstances in the laboratory. An animal's ability to synchronize its foraging activity with the time of day when food was most available was tested. Energy costs (wheel revolutions) were varied to encourage an animal to redistribute its activity from a preferred 12-h phase of the light-dark cycle to the other phase, and then back again. The degree to which activity changed to coincide with the most economical phase was analysed for each individual, and weasels were compared to mink. The activity of six (of seven) animals shifted in response to modified foraging costs, but only two animals (both mink) approached the 180° redistribution of activity expected of an energy-maximizing forager. In general, all animals were active during a favoured phase (usually dark) of the light-dark cycle and relatively large variations in foraging cost during this cycle had only modest effects on activity.     

Effect of constant and fluctuating temperature on the circadian foraging rhythm of the red imported fire ant,Solenopsis invicta Buren (Hymenoptera: Formicidae)

Understanding circadian foraging rhythms activity of the red imported fire ant, Solenopsis invicta Buren (Hymenoptera: Formicidae) foragers at different temperatures is an important step towards developing control measures in Integrated Pest Management (IPM) programs. In this study, the circadian foraging rhythm activities of S. invicta foragersat different temperature were investigated under laboratory and field conditions. Results indicated that the foraging activity increased after sunrise, and maximum foraging occurred at 14:00 (foraging rate was 69.22 ± 0.57 and 72.58 ± 1.15 foragers/min in the first and second year, respectively) in the tea fields of Guangzhou during autumn. Furthermore, foragers demonstrated circadian rhythms and exhibited a unimodal after 24 h. A significant correlation was found between foraging activity and temperature. S. invicta colonies were active at moderate soil temperatures (approximately 26.65 °C to 29.24 °C). The preferred temperature of the colonies was 26 °C, followed by 22 °C and 18 °C in the laboratory. The individual S. invicta activity was maximum at 17:00 (18.67 ± 1.66 times /10 min) and minimum at 5:00 (8.33 ± 2.51 times/10 min) at 26 °C. The fluctuating temperature had a significant impact on individual locomotor activity (r = 0.8979, P < 0.01) but did not alter the rhythm activity. Our results demonstrated that temperature might play an important role in circadian foraging rhythms activity of S. invicta. These results may have implications for the development of more effective fire ant management strategies.     

Activity Pattern of the Trawling Phyllostomid Bat,Macrophyllum macrophyllum,in Panamá1

We studied activity patterns of long‐legged bats, Macrophyllum macrophyllum (Phyllostomidae), in the Barro Colorado Nature Monument, Panamá, using radio‐telemetry. Activity of four males and five females equipped with radio‐transmitters were monitored for 4–7 entire nights each between April and July 2002. Bats exhibited maximum activity around dusk and high activity during the night. Males and females foraged for equal amounts of time in continuous flight (mean: 7 min, maximum 1 h) with interspersed resting phases (mean: 15 min, maximum 3 h). Activity of M. macrophyllum was sensitive to several factors. Time of emergence and return to day roost were correlated with time of sunset and sunrise, respectively. Maximum bat activity coincided with high abundance of aerial insects. Finally, heavy rain caused bats to reduce or cease flight activity. Direct observations and field video recordings support the assumption that M. macrophyllum employs two distinct foraging modes: trawling of insects from and capture of aerial insects at low heights above water. Combination of foraging modes gives M. macrophyllum high flexibility and efficiency in prey search. Activity, foraging mode, and morphology, which are similar to trawling bats from other families, distinguish M. macrophyllum from all other phyllostomid species and grant it access to open habitat above water, a habitat no other phyllostomid bat has conquered.     

Seasonality in foraging behaviour of Constrictotermes cyphergaster (Termitidae, Nasutitermitinae) in the Caatinga of Northeastern Brazil

The foraging activity of Constrictotermes cyphergaster was investigated in the Caatinga of Northeast Brazil. Eight colonies were monitored for seven days, during both dry and wet seasons. Foraging activity occurred in exposed columns at night, generally between 22:00 and 05:00 h. During the wet season, foraging activity was significantly higher, with one bout every 1.6 ± 0.2 days, than the dry season, when foraging bouts were performed every 1.9 ± 0.3 days. Foraging activity throughout the study colonies presented high temporal synchronization. In both seasons, foraging was negatively correlated with air temperature and positively correlated with humidity. The foraging trails were often re-utilized and ranged from 1 to 18.5 meters in length. No difference between seasons in the area potentially utilized by the study colonies was observed. Approximately 51000 individuals participated in the foraging bout during the dry season, whereas some 87000 individuals participated in the foraging bout during the wet season. This corresponds to 43 and 74% of the estimated total nest population for the dry and wet seasons respectively. The average ratio soldiers:workers during foraging was 1:1.2 in the dry season and 1:2 in the wet season. The higher frequency and number of individuals foraging during the wet season in the present study are likely to be a strategy from C. cyphergaster to store energy reserves to be utilized during the dry season. Received 28 November 2005; revised 29 May 2006 and 16 August 2006; accepted 1 September 2006.     

The regulation of activity in populations of the terrestrial slugLimax maximus (Gastropoda; Limacidae)

The activity of the slug Limax maximus was studied in relation to weather. Three hundred-and-fifty-eight hourly observations of activity and weather were made on 21 nights from May until October, 1976. Factors causally important to molluscan activity were included in a step-down correlation-regression analysis of daily and seasonal behavior. The analysis was also performed using weather data from the previous hourly observation. Models using lag-weather did not explain as much variability as did concurrent weather. The regression models explained about 73% to 87% of the observed variation in activity. The most important factors included in the regression models were time of day (circadian rhythm), light intensity, changes in light intensity and surface temperature. Shelter temperature, temperature gradients, length of the night, and time of sunset were also included in some models. Age and hydration were shown to be key factors in other experiments. A model incorporating weather thresholds estimated from field data explained 83.06% of the variability in the activity of L. maximus over the season. The values predicted from the model did not differ significantly from those actually observed in the field (Kolmogorov-Smirnov test, p>0.50).     

Temperature influences the handling efficiency of an aphid parasitoid through body size-mediated effects

It is well known that increasing the ambient temperature increases the metabolic rate and consequently, the foraging rate of most insects. However, temperature experienced during the immature stages of insects affects their adult size (an inverse relationship). Because body size is generally correlated to foraging success, we hypothesized that temperature indirectly influences the foraging efficiency of adult insects through developmental effects. We first investigated the role of parasitoid: host body size ratio on the handling time of Aphidius colemani (Viereck) (Hymenoptera: Braconidae), then tested the prediction that increasing temperature during immature development increases the handling time of adults. As expected, parasitoids took longer to handle large aphids than small aphids. However, large parasitoids did not have shorter handling times than small parasitoids except when attacking large (adult) aphids. Developmental temperature had the predicted effect on parasitoids: Individuals reared at 25°C were smaller than those insects reared at 15°C. Parasitoids reared at 15°C had similar short handling times for both first instar and adult aphids, whereas parasitoids reared at 25°C took longer to handle adult aphids than first instar aphids. The size-mediated effect of temperature through development on parasitoid efficiency was opposite to the more familiar direct effect of temperature through metabolic rate. We conclude that the net effect of temperature on foraging insects will depend on its relative influence on immature and adult stages.     

Summer diel activity and movement of adult brown trout in high-elevation streams in Wyoming, U.S.A.

The variation in diel activity patterns was determined by implanting transmitters in adult brown trout of various sizes in two montane streams in two summers in the central Rocky Mountains, U.S.A. More brown trout tended to be active, and to move greater distances, each hour during twilight and night than during the day. Mean hourly light intensity was significantly negatively correlated with the proportion of fish active and with the distance moved. Maximum water temperature was not correlated with any measure of diel activity. Brown trout displayed fidelity to diurnal positions within a given diel cycle. Overall, fish were active an average of 11 h, had an average home range of 41 m, and moved an average of 121 m over the diel cycle. Fish length and distance moved were significantly positively correlated. Water temperature, predator avoidance, and foraging tactics may influence these patterns of diel behaviour in adult brown trout in U.S. streams.     

Insights into short‐ and long‐term crop‐foraging strategies in a chacma baboon (Papio ursinus) from GPS and accelerometer data

Crop‐foraging by animals is a leading cause of human–wildlife “conflict” globally, affecting farmers and resulting in the death of many animals in retaliation, including primates. Despite significant research into crop‐foraging by primates, relatively little is understood about the behavior and movements of primates in and around crop fields, largely due to the limitations of traditional observational methods. Crop‐foraging by primates in large‐scale agriculture has also received little attention. We used GPS and accelerometer bio‐loggers, along with environmental data, to gain an understanding of the spatial and temporal patterns of activity for a female in a crop‐foraging baboon group in and around commercial farms in South Africa over one year. Crop fields were avoided for most of the year, suggesting that fields are perceived as a high‐risk habitat. When field visits did occur, this was generally when plant primary productivity was low, suggesting that crops were a “fallback food”. All recorded field visits were at or before 15:00. Activity was significantly higher in crop fields than in the landscape in general, evidence that crop‐foraging is an energetically costly strategy and that fields are perceived as a risky habitat. In contrast, activity was significantly lower within 100 m of the field edge than in the rest of the landscape, suggesting that baboons wait near the field edge to assess risks before crop‐foraging. Together, this understanding of the spatiotemporal dynamics of crop‐foraging can help to inform crop protection strategies and reduce conflict between humans and baboons in South Africa.     

Patterns in foraging and nesting ecology in the neotropical ant, <Emphasis Type="Italic">Gnamptogenys moelleri</Emphasis> (Formicidae,Ponerinae)

Summary This study provides quantitative field data on the natural history and foraging behaviour of the Neotropical bromeliad-nesting ant Gnamptogenys moelleri (Ponerinae) in a sandy plain forest in Southeast Brazil. The ant nested on different bromeliad species and the nests were more frequently found in bigger bromeliads. The species used a wide array of invertebrates in its diet, hunting for live prey and scavenging the majority of the items from dead animals. The food items varied greatly in size (1 to 26 mm). Hunting was always performed by solitary workers. Retrieving was performed by solitary workers (small items), or by a group of 3 to 12 workers recruited to the food source (large items). Almost all G. moelleri foraging activity was restricted to the nest bromeliad. In the warm period more ants left the nest to forage, and foraging trips achieved greater distances compared to the cool season. Trap data revealed that overall availability of arthropod prey is higher in the summer than in the winter. The opportunism in nest site use and in foraging behaviour, the small foraging area, as well as the seasonal differences in foraging activity are discussed and compared with other tropical ants.Received 30 May 2003; revised 22 September 2003; accepted 3 October 2003.     

Critical thermal limits in Mediterranean ant species: trade-off between mortality risk and foraging performance

1. In Mediterranean ant communities, a close relationship has been found between activity rhythm in the period of maximum activity and position in the dominance hierarchy: subordinate species are active during the day, when conditions are more severe, while dominants are active during the afternoon and the night.
2. Results obtained in this study confirmed that the species foraging at higher temperatures were closer to their critical thermal limits than the species foraging at lower temperatures.
3. This enabled two extreme strategies of foraging in relation to temperature to be distinguished: (1) heat-intolerant ant species behaved as risk-averse species, foraging at temperatures very far from their critical thermal limits; and (2) heat-tolerant ant species behaved as risk-prone species, foraging very near their critical thermal limits and running a high heat mortality risk.
4. Heat-tolerant species benefited from this strategy by having better foraging performance at high temperatures.
5. This wide range of thermal niches may be one reason why Mediterranean ant faunas are so diverse in the face of limited diversity in vegetation and habitat structure: the daily range of temperature may be sufficiently great to meet the requirement both of heat-adapted and cold-adapted species as well as a spectrum of intermediate forms.     

To minimize foraging time,use high‐efficiency,energy‐expensive search and capture methods when food is abundant but low‐efficiency,low‐cost methods during food shortages

Based on a mathematical model, I show that the amount of food in the habitat determines which among alternative methods for search of prey, respectively, for pursuit‐and‐capture give the shortest daily foraging time. The higher the locomotor activity, the higher the rate of energy expenditure and the larger the habitat space a predator can search for prey per time unit. Therefore, I assume that the more efficient a foraging method is, the higher its rate of energy expenditure. Survival selection favors individuals that use foraging methods that cover their energy needs in the shortest possible time. Therefore, I take the optimization criterion to be minimization of the daily foraging time or, equivalently, maximization of the rate of net energy gain. When time is limiting and food is in short supply, as during food bottleneck periods, low‐efficiency, low‐cost foraging methods give shorter daily foraging times than high‐efficiency, energy‐expensive foraging methods. When time is limiting, food is abundant and energy needs are large, as during reproduction, high‐efficiency high‐cost foraging methods give shorter daily foraging times than low‐efficiency low‐cost foraging methods. When time is not limiting, food is abundant, and energy needs are small, the choice of foraging method is not critical. Small animals have lower rates of energy expenditure for locomotion than large animals. At a given food density and with similar diet, small animals are therefore more likely than large ones to minimize foraging time by using high‐efficiency energy‐expansive foraging methods and to exploit patches and sites that require energy‐demanding locomotion modes. Survival selection takes place at food shortages, while low‐efficiency low‐cost foraging methods are used, whereas reproduction selection occurs when food is abundant and high‐efficiency energy‐expensive foraging methods do better. In seasonal environments, selection therefore acts on different foraging methods at different times. Morphological adaptation to one method may oppose adaptation to another. Such conflicts select against foraging and morphological specialization and tend to give species‐poor communities of year‐round resident generalists. But a stable year‐round food supply favors specialization, niche narrowing, and dense species packing.     

Production of greenhouse tomatoes (Lycopersicon esculentum) using Nannotrigona perilampoides, Bombus impatiens and mechanical vibration (Hym.: Apoidea)

Importation of exotic bumblebees for greenhouse pollination may be restricted in México, thus making it necessary to evaluate the potential of native species as pollinators in enclosures. We studied the foraging activity and fruit production of tomato using one colony of Nannotrigona perilampoides (NP) and one colony of Bombus impatiens (BI) in greenhouses with ≈1000 plants. Mechanical vibration (MV) was included as a test treatment. The foraging activity was measured as the number of flowers visited within 5 min, the time spent on a flower collecting pollen, the number of visits that a flower received and the duration of a foraging trip. BI collected pollen more rapidly, visited more flowers within 5 min and did more visits per flower when compared with NP that also lasted longer in their trips. Significant correlations were found between environmental variables and the number of bees entering the hive and the number of bees on the flowers. For NP, the highest correlation was found for light intensity whilst in BI a negative effect of environmental temperature was detected. Regarding the quantity of fruit, BI resulted in higher fruit set when compared with NP, but the latter performed similarly to MV. However, the weight of the fruit and seed number was significantly higher for BI when compared with NP, and this was higher than for MV. Our results demonstrate that at the densities of tomato plants tested, one colony of BI was more efficient pollinator when compared with NP. We suggest that pollination efficiency of NP could have been limited by a reduced number of foragers on the plants at a given time and their limited flight range when compared with BI. Therefore, it will be necessary to evaluate if increasing densities of colonies of NP could improve tomato yield in tropical greenhouses.     

Antipredator Behavior in Desmognathus ochrophaeus: Threat‐Specific Responses to Chemical Stimuli in a Foraging Context

Prey species may reduce the likelihood of injury or death by engaging in defensive behavior but often incur costs related to decreased foraging success or efficiency. To lessen these costs, prey may adjust the intensity or type of antipredator behavior according to the nature of the perceived threat. We evaluated the potential for threat‐sensitive responses by Allegheny Mountain dusky salamanders (Desmognathus ochrophaeus) exposed to chemical stimuli associated with predation by asking three questions: (1) Do individual D. ochrophaeus respond to chemical cues in a threat‐sensitive manner? (2) Do salamanders exhibit the same pattern of behavioral response while foraging? and (3) Is foraging efficiency reduced when focal individuals are exposed to stimuli from predators or predation events? In our first experiment, we evaluated salamander chemosensory movements (nose‐taps), locomotor activity (steps), and edge behavior in response to chemical stimuli from disturbed and injured conspecifics as well as predatory Gyrinophilus porphyriticus and found that individual D. ochrophaeus show a significant graded increase in nose‐taps when exposed to cues from conspecifics and a reduction in activity when exposed to the predator. In our second experiment, we again observed salamander responses to the same chemical stimuli but in this instance added five Drosophila prey to the test dishes. We found that salamanders exhibited a similar pattern of response to the chemical stimuli in the presence of prey, showing a graded increase in nose‐taps to cues from conspecifics and a reduction in activity when exposed to the predator. However, foraging efficiency (i.e. the proportion of successful strikes) did not vary significantly among treatments. Our data show that individual D. ochrophaeus detect and differentially respond to chemical stimuli associated with predation, but do not significantly reduce foraging efficiency. Overall, the type and relative intensity of these responses is largely unaffected by the presence of potential prey.     

Environmental factors influencing daily foraging activity of <Emphasis Type="Italic">Vespula germanica</Emphasis> (Hymenoptera,Vespidae) in Mediterranean Australia

In social insects, workers forego reproduction in favour of foraging and other tasks to promote growth of the whole colony. Maximising individual work effort is limited by the physical constraints on foraging outside the nest. Previous studies of factors influencing activity in social insects suggest that light intensity, ground and air temperature, relative humidity, solar radiation, wind speed and/or rainfall may be important. This study aimed to determine which environmental factors influence foraging in an invasive social insect, Vespula germanica, which has been introduced to Australia, where it experiences a hotter and drier climate than in its native range. Activity was measured in terms of foraging traffic, with nests from a range of locations being monitored. Results indicate that the onset of rain reduces activity by approximately 30%, but foraging returns to previous levels immediately after rain stops. Foraging time is correlated with duration of daylight, with wasp daily activity being on average 22 min longer than the time between sunrise and sunset. Low light was found to restrict wasp activity, as were low and high temperatures. A linear mixed-effects model developed to explain the influence of these variables on numbers of foragers was highly significant. Under hot conditions, V. germanica individuals thermoregulate their body temperature by regurgitating water. They also use water in evaporative cooling to keep nests at optimum temperatures. Thus, in this species, hot temperatures increase the need for water, and so populations may be severely impacted in seasonally hot regions where water is limited. Received 30 March 2007; revised 10 March 2008; accepted 9 April 2008.     

Time‐series models to quantify short‐term effects of meteorological conditions on bumblebee forager activity in agricultural landscapes

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Crowding increases foraging efficiency in the leaf-cutting ant <Emphasis Type="Italic">Atta colombica</Emphasis>

Many animals, including humans, organize their foraging activity along well-defined trails. Because trails are cleared of obstacles, they minimize energy expenditure and allow fast travel. In social insects such as ants, trails might also promote social contacts and allow the exchange of information between workers about the characteristics of the food. When the trail traffic is heavy, however, traffic congestion occurs and the benefits of increased social contacts for the colony can be offset by a decrease of the locomotory rate of individuals. Using a small laboratory colony of the leaf-cutting ant Atta colombica cutting a mix of leaves and Parafilm, we compared how foraging changed when the width of the bridge between the nest and their foraging area changed. We found that the rate of ants crossing a 5 cm wide bridge was more than twice as great as the rate crossing a 0.5 cm bridge, but the rate of foragers returning with loads was less than half as great. Thus, with the wide bridge, the ants had about six times lower efficiency (loads returned per forager crossing the bridge). We conclude that crowding actually increased foraging efficiency, possibly because of increased communication between laden foragers returning to the nest and out-going ants. Received 15 December 2006; revised 16 February 2007; accepted 19 February 2007.