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Thermal constraints for stingless bee foragers: the importance of body size and coloration.

In the dry tropics, foraging bees face significant thermal constraints as a result of high ambient temperatures and direct insolation. In order to determine the potential importance of body size and body coloration in heat gain and heat loss, passive warm-up and cooling rates were measured for freshly killed workers of 24 stingless bee species. Results accorded with biophysical principles. Small bees reached lower temperature excesses ( T(exc)) and warmed up and lost heat much more rapidly than larger bees. In addition to body size, body coloration had a clear effect on thermal parameters. Light-coloured bees warmed up less rapidly and had lower T(exc) than dark bees. An intraspecific comparison of Melipona costaricensis and Cephalotrigona capitata colour morphs confirmed that body coloration influences thermal characteristics. This study is the first to indicate that abdominal coloration in stingless bees might be involved in the regulation of body temperature in extreme thermal conditions. However, body temperatures of foraging bees of colour morphs were not very different. This is probably due to behavioural adaptations (e.g. foraging strategies) or differences in convective and evaporative heat loss or the production of metabolic heat during flight, that all mask the effect of body colour. Notwithstanding such effects and potential thermoregulatory capabilities, stingless bees show niche differentiation and biogeographic distributions that correlate with body coloration and body size. This also suggests that, in general, light bees have an advantage over black bees in hot open lowland habitats, whereas black bees might have an advantage in wet habitats and mountains. The origin, occurrence and function of flavinism (yellow integument colouring) are discussed.

Animals↗

How do frugivores track resources? Insights from spatial analyses of bird foraging in a tropical forest.

Frugivores often track ripe fruit abundance closely across local areas despite the ephemeral and typically patchy distributions of this resource. We use spatial auto- and cross-correlation analyses to quantify spatial patterns of fruit abundance and avian frugivory across a 4-month period within a forested 4.05-ha study grid in Puerto Rico. Analyses focused on two tanager species, Spindalis portoricensis and Nesospingus speculiferus, and their principal food plants. Three broad questions are addressed: (1). at what spatial scales is fruit abundance and frugivory patchy; (2). at what spatial scales do frugivores respond to fruit abundance; and (3). to what extent do spatial patterns of frugivory overlap between bird species? Fruit patch size, species composition, and heterogeneity was variable among months, despite fruit patch locations remaining relatively consistent between months. Positive correlations between frugivory and fruit abundance suggested tanagers successfully tracked fruit abundance. Frugivory was, however, more localized than fruit abundance. Scales of spatial overlap in frugivory and monthly variation in the foraging locations of the two tanager species suggested that interspecific facilitation may have been important in determining bird foraging locations. In particular, S. portoricensis, a specialist frugivore, may have relied on the loud calls of the gregarious generalist, N. speculiferus, to find new foraging areas. Such a mechanism could help explain the formation of mixed species feeding flocks and highlights the potential importance of facilitation between species that share resources.

Animals↗

Using stable isotope analysis with telemetry or mark-recapture data to identify fish movement and foraging.

Information about animal movements has often been inferred from stable isotope analysis (SIA), but is dependent on animals assimilating site-specific isotopic signatures via diet. This potential weakness in ecological interpretation can be overcome by using other investigative tools that provide precise information about individual movement patterns. In this paper, we demonstrate the value of combining SIA with telemetry or mark-recapture data from trapping, electrofishing and remote detection of individuals to study the movement and feeding ecology of fishes in different habitats. In a fjord lake system in Newfoundland, Canada, juvenile Atlantic salmon delayed downstream migration (smolts) or actively moved into a large lake (parr) where they foraged for periods reflecting different life history strategies. In the Miramichi River (New Brunswick, Canada), SIA provided evidence of distinct foraging habitats (tributary versus large river). By tracking fish implanted with passive integrated transponder (PIT) tags, we distinguished between movements related to foraging versus seeking cool water refugia during high temperature events. Finally, site fidelity and limited mobility of slimy sculpin, a small benthic fish, was established where delta13C in muscle tissue showed a progressive enrichment downstream and where a median displacement of <10 m was estimated for sculpin tagged with PIT tags. Technological improvements have permitted non-destructive tissue sampling of wild fishes for SIA, and the tagging and remote detection of animals smaller than was previously possible. These advancements and the combination of investigative tools promise new insights into animal ecology.

Animals↗

Experimental manipulation of predation risk and food quality: effect on grazing behaviour in a central-place foraging herbivore.

The relative importance of predation risk and food quality on spatial grazing pressure and activity patterns was tested in a central-place foraging herbivore: the European rabbit. Rabbits grazed less with increasing distance from their burrows into adjacent grassland, thereby creating a gradient of increasing vegetation height and plant biomass and decreasing plant nutrient concentration. When nitrogen concentration was experimentally increased by 150% through fertilizing and mowing, rabbits visited these plots four times more frequently than the untreated control plots. Addition of predator scent (mink pellets) did not result in different patch use by rabbits. The combined addition of fertilizer and mink pellets had the same effect as addition of fertilizer alone. However, the mink pellets changed the temporal activity pattern of rabbits as measured with infrared detectors. Rabbits were predominantly nocturnal but shifted their activities to the day when mink pellets were added, resulting in equal activities during night and day. We conclude that rabbits are sensitive to perceived predation risk, but that this does not influence their spatial grazing pressure. A selection for the highest food quality on the other hand can explain the observed natural rabbit grazing gradient. Food quality was highest close to the burrows, therefore rabbits selecting for high quality food should forage most intensely close to the burrows and only move further away for higher quality items or when the vegetation close to their burrows is depleted. Through intensive grazing close to the burrows rabbits facilitated for themselves either through stimulating fresh protein rich re-growth or the return of nutrients through faeces or both. This is in contrast with central-place foraging theory where intense feeding close to the burrow is assumed to lead to reduced food resources.

Animals↗

Spatial context influences patch residence time in foraging hierarchies.

Understanding responses of organisms to spatial heterogeneity in resources has emerged as a fundamentally important challenge in contemporary ecology. We examined responses of foraging herbivores to multi-scale heterogeneity in plants. We asked the question, "Is the behavior observed at coarse scales in a patch hierarchy the collective outcome of fine scale behaviors or, alternatively, does the spatial context at coarse scales entrain fine scale behavior?" To address this question we created a nested, two-level patch hierarchy. We examined the effects of the spatial context surrounding a patch on the amount of time herbivores resided in the patch. We developed a set of competing models predicting residence time as a function of the mass of plants contained in a patch and the distance between patches and examined the strength of evidence in our observations for these models. Models that included patch mass and inter-patch distance as independent variables successfully predicted observed residence times (bears: r (2)=0.67-0.76 and mule deer: r (2)=0.33-0.55). Residence times of grizzly bears (Ursus arctos) and mule deer (Odocoileus hemionus) responded to the spatial context surrounding a patch. Evidence ratios of Akaike weights demonstrated that models containing effects of higher levels in the hierarchy on residence time at lower levels received up to 34 times more support in the data than models that failed to consider the higher level context for grizzly bears and up to 48 times more support for mule deer. We conclude that foraging by large herbivores is influenced by more than one level of heterogeneity in patch hierarchies and that simple empirical models offer a viable alternative to optimal foraging models for the prediction of patch residence times.

Animals↗

Methyl parathion modifies foraging behaviour in honeybees (Apis mellifera).

We examined the effects of sublethal doses of an organophosphorus insecticide, Methyl Parathion (MeP), on the foraging behaviour of honeybees (Apis mellifera ligustica) in a flight cage. The results revealed that MeP modified the frequency of visits to a feeding station to which the bees had previously been trained. A dose of 50 ng per animal elicited an increase in the frequency of visits to the feeder, compared to control animals. A dose of 10 ng, on the other hand, led initially to a decrease in the visit frequency, followed by an increase to a level above that of the controls. A hypothesis is presented to account for the way in which MeP affects foraging behaviour. We propose that the behavioural assay presented here could be useful as a preliminary screening test to study sublethal effects of pesticides on foraging performance in honeybees.

Animals↗

Metal concentrations, foraging distances, and fledging success of great blue herons nesting along the Hanford Reach of the Columbia River.

An ecological risk assessment of the spatial distribution of metal concentrations along the Hanford Reach of the free-flowing Columbia River in southcentral Washington, identified great blue herons, Ardea herodias, at potential risk through the ingestion of contaminated riverine biota, especially fish. We measured metal concentrations in livers of pre-flight herons from the Hanford Reach and excrement samples taken from the same nests. Nests were distributed among three colonies situated upstream and downstream from nine retired plutonium production reactors along the river where metals in reactor coolant waters had been released directly into the river or disposed to shoreline retention basins and ditches. Distances traveled by parent herons to foraging areas along the river shore were determined by visually tracking parent birds as they flew from nests to upriver and downriver foraging sites. Foraging flight distances varied between colonies with mean distances ranging between 0.7 and 3.1 km. Cadmium, Cr, and Pb concentrations were higher in excrement than in the livers of pre-flight herons but the opposite was noted for Cu, Hg, and Zn. Highest metal concentrations of Cr, Cu, Zn, Cd, and Pb, were measured in excrement taken from heron nests at the colony located upstream from all reactors. These results were consistent with metal concentrations reported in river sediment from the same regions, indicating excrement from the heron nests may be a useful indicator of dietary uptake of metals by herons. Fledging success and eggshell thickness measurements were used as an index of health of the local heron population. The results indicate that the reproductive health of great blue herons nesting along the Hanford Reach is among the highest reported in the continental United States.

Animals↗

Solitary foraging in the ancestral South American ant, Pogonomyrmex vermiculatus. Is it due to constraints in the production or perception of trail pheromones?

Several North American species of Pogonomyrmex harvester ants exhibit group foraging, whereas South American species are exclusively solitary foragers. The composition of the secretions of the poison and Dufour glands in the South American species, Pogonomyrmex vermiculatus, were analyzed, and the secretions and their components were tested as trail pheromones in laboratory bioassays. The major compounds in the poison gland were the alkylpyrazines, 2,5-dimethylpyrazine, 2,3,5-trimethylpyrazine, and 3-ethyl-2,5-dimethylpyrazine. The Dufour gland contained five alkanes, from tridecane to heptadecane, with pentadecane being most abundant. In behavioral bioassays, poison gland extracts and the mixture of pyrazines produced a trail pheromone effect, whereas the Dufour gland extracts and the alkanes had no effect on ant locomotion. We conclude that group foraging in P. vermiculatus does not arise from the inability to produce or detect possible pheromones, but rather, from physiological and/or ecological factors.

Animal Communication↗

Behavioral and physiological responses of female house mice to foraging variation.

Peripubertal female house mice were required to work for their food at either 23 degrees C or 9 degrees C. We used a special caging system in which animals had to emerge from a thermally-buffered burrow and run a programmable number of running wheel revolutions to obtain a pellet of food. Of concern here were the behavioral and physiological adjustments necessary to accommodate growth and reproductive development when faced with the need to forage for different lengths of time at different temperatures. When female house mice are confronted with poor foraging at cool temperatures they allot their highest priorities to maintaining energy balance; body growth is next, and reproductive development and nonforaging activity have the lowest priorities. Our results also demonstrate that the time spent foraging while exposed to low ambient temperature is critical for this species. This relationship probably determines whether or not house mice will breed continuously or seasonally in a particular habitat.

Animals↗

Development of an ethanol model using social insects: V. Honeybee foraging decisions under the influence of alcohol.

We examined the effect of ethanol on a complex decision process in honeybees. In many contexts (e.g., flower patches containing both blue and white flowers), honeybees have been shown to make decisions concerning which flowers to visit based on the reward's caloric value offered by competing flower types. However, under some contexts (e.g., flower patches containing both blue and yellow flowers) honeybees ignore the energetic differences that exist between rewards. When this occurs, some bees show extreme fidelity to one flower type, and other bees simultaneously show extreme loyalty to the competing flower type. In this set of experiments we use different combinations of flower colors to elicit these different context-specific behaviors, and examine response under the influence of ethanol in conditions with rewards differing in either quality or quantity. Alcohol affected the same decision situation differently in the alternative contexts we gave foragers. In the blue-white dimorphic flower patch context, foragers in the alcohol condition no longer followed the species' typical behavior of selectively choosing the higher energetic reward; they chose flowers randomly as if rewards did not differ between flower colors. However, in the blue-yellow dimorphic flower patch context, foragers in the alcohol condition continued their species-typical behavior of favoring their initial flower-color choice. That is, alcohol had no effect on behavior in the latter context. This pattern of context-dependent effects of alcohol was not associated with motor impairment in either context because flower visitation rate increased when bees were given ethanol. The brain regions responsible for alternative context behaviors (using the Drosophila model), or at least receptors within the same region responsible for these context-dependent behaviors, have very different sensitivities to alcohol, which suggests that differing neural processes are involved.

Animals↗

Influence of physical stress, distress cues, and predator kairomones on the foraging behavior of Ozark zigzag salamanders, Plethodon angusticlavius.

An animal's response to environmentally induced stress can have important fitness consequences. For example, an animal may reduce activity in response to stress related to increased predation risk, but this response carries the potential cost of missed foraging or mating opportunities. We used laboratory bioassays to determine whether Ozark zigzag salamanders, Plethodon angusticlavius, decreased their feeding behavior following increased levels of threat-related stress. Stress level was manipulated in three experiments where salamanders were exposed to direct or indirect indicators of stress. The direct cue was a physical attack. The indirect cues were chemical cues from distressed conspecifics and chemical cues ("kairomones") from predatory ringneck snakes, Diadophis punctatus. For all three cues, stressed salamanders showed lower foraging success than unstressed salamanders. The physical attack did not result in a more intense response than the other treatments, possibly because the attack occurred away from the foraging area. The intensity of the response to stress may depend more on the nature of the threat (identity and imminence) than the level of physical disturbance.

Adaptation, Physiological↗

Momentary maximizing and optimal foraging theories of performance on concurrent VR schedules.

Optimal foraging theory proposes that animals obtain the highest rate of reinforcers for the least effort and momentary maximizing theory proposes that animals make the response that at that instant is most likely to be reinforced. While each theory may account for matching on concurrent schedules, the data supporting each theory are weak. Two experiments assessed these theories by considering concurrent choice as consisting of two pairs of stay and switch schedules. Symmetrical arrangements, which are equivalent to standard concurrent schedules, maintained behavior described by the generalized matching law. Weighted arrangements, in which the programmed rate of earning reinforcers was always greater at one alternative, maintained behavior that was biased towards the weighted alternative, yet the bias was less than that predicted by optimal foraging theory. Asymmetrical arrangements, in which the stay and switch schedules operating at an alternative are the same, maintained behavior that favored one alternative, even though momentary maximizing predicted indifference. The generalized matching law poorly described each rat's pooled data from all conditions but these data were described by an equation based on the stay and switch reinforcers earned per-visit and included elements of optimal foraging and momentary maximizing theories of choice.

Animals↗

Evaluation of sewage sludge, septic waste and sludge compost applications to corn and forage: yields and N, P and K content of crops and soils.

This paper presents the data from two years of experiments concerned with the application of aerobically-digested sewage sludge, anaerobic lagoon septic sludge, sewage sludge compost or fertilizer to soils for grass forage and feed corn (Zea mays L.) production at two different sites 45 km from Truro, Nova Scotia. Crop yields, plant tissue and Mehlich-1 extractable soil nutrients were evaluated; 15 elements were analyzed in the plant tissue and nine elements in the soil extracts. This paper describes the results of crop yields, plant N, P and K content and Mehlich-1 extractable P and K. The research demonstrated the fertilizer produced higher yields of grass forage than the sludge and the compost but equivalent to the sludge in corn yields. Forage and corn N, P and K contents, however, varied with treatment, crop and year, while the compost-amended soils were highest in extractable nutrients. Both sludges and the compost, therefore, could be effective sources of N, P and K for crop production. Compared to the conventional fertilizer, the nutrient availability from the organic amendments (especially N and P) was considerably lower than the 50% assumed at the start of the experiment; the sludges however, provided higher nutrient availability than the compost.

Animal Feed↗

Assessment of pesticide residues in army cutworm moths (Euxoa auxiliaris) from the Greater Yellowstone Ecosystem and their potential consequences to foraging grizzly bears (Ursus arctos horribilis).

During summer, a grizzly bear (Ursus arctos horribilis) in the Greater Yellowstone Ecosystem (GYE) (USA) can excavate and consume millions of army cutworm moths (Euxoa auxiliaris) (ACMs) that aggregate in high elevation talus. Grizzly bears in the GYE were listed as threatened by the US Fish and Wildlife Service in 1975 and were proposed for delisting in 2005. However, questions remain about key bear foods. For example, ACMs are agricultural pests and concern exists about whether they contain pesticides that could be toxic to bears. Consequently, we investigated whether ACMs contain and transport pesticides to bear foraging sites and, if so, whether these levels could be toxic to bears. In 1999 we collected and analyzed ACMs from six bear foraging sites. ACMs were screened for 32 pesticides with gas chromatography with electron capture detection (GC-ECD). Because gas chromatography with tandem mass spectrometry (GC-MS/MS) can be more sensitive than GC-ECD for certain pesticides, we revisited one site in 2001 and analyzed these ACMs with GC-MS/MS. ACMs contained trace or undetectable levels of pesticides in 1999 and 2001, respectively. Based on chemical levels in ACMs and numbers of ACMs a bear can consume, we calculated the potential of chemicals to reach physiological toxicity. These calculations indicate bears do not consume physiologically toxic levels of pesticides and allay concerns they are at risk from pesticides transported by ACMs. If chemical control of ACMs changes in the future, screening new ACM samples taken from bear foraging sites may be warranted.

Animals↗

Phenotypically linked dichotomy in sea turtle foraging requires multiple conservation approaches.

Marine turtles undergo dramatic ontogenic changes in body size and behavior, with the loggerhead sea turtle, Caretta caretta, typically switching from an initial oceanic juvenile stage to one in the neritic, where maturation is reached and breeding migrations are subsequently undertaken every 2-3 years. Using satellite tracking, we investigated the migratory movements of adult females from one of the world's largest nesting aggregations at Cape Verde, West Africa. In direct contrast with the accepted life-history model for this species, results reveal two distinct adult foraging strategies that appear to be linked to body size. The larger turtles (n = 3) foraged in coastal waters, whereas smaller individuals (n = 7) foraged oceanically. The conservation implications of these findings are profound, with the population compartmentalized into habitats that may be differentially impacted by fishery threats in what is a global fishing hotspot. Although the protection of discrete areas containing coastal individuals may be attainable, the more numerous pelagic individuals are widely dispersed with individuals roaming over more than half a million square kilometers. Therefore, mitigation of fisheries by-catch for sea turtles in the east Atlantic will likely require complex and regionally tailored actions to account for this dichotomous behavior.

Africa, Western↗

Prescribed burning effects on summer elk forage availability in the subalpine zone, Banff National Park, Canada.

The effects of prescribed burning on forage abundance and suitability for elk (Cervus elaphus) during the snow-free season was evaluated in east-central Banff National Park, Canada. Six coniferous forest and mixed shrub-herb plant communities (n=144 plots), and 5223ha of burned (n=131) vegetation <12 years old were sampled using a stratified semi-random design. Sampling units represented various combinations of vegetation, terrain conditions, and stand ages that were derived from digital biophysical data, with plant communities the basic unit of analysis. Burning coniferous forest stands reduced woody biomass, and increased herbaceous forage from 146 to 790 kg/ha. Increases commonly occurred in the percent cover of hairy wild rye (Leymus innovatus (Beal) Pigler) and fireweed (Chamerion angustifolium (L.) Holub.). The herbaceous components of mixed shrub-herb communities increased from 336-747 kg/ha to 517-1104 kg/ha in response to burning (P<0.025, Mann-Whitney U-test). Browse biomass (mostly Salix spp. and Betula nana L.) increased >or=220% (P<or=0.003, Mann-Whitney U-test) from 653 kg/ha in deciduous shrub types. Elk preferences for unburned and burned vegetation-types were assessed as low and moderate, respectively. Potential summer carrying capacity, based on forage availability, increased from eight to 28 elk/100 km2 within burned areas, whereas spring grazing potential rose from 13 to 45 elk/100 km2. Most of the increase (73%) was attributable to changes within burned Engelmann Spruce stands, which composed 58% of the burned area.

Alberta↗

Prevalence and the duration of linear enamel hypoplasia: a comparative study of Neandertals and Inuit foragers.

As a dental indicator of generalized physiological stress, enamel hypoplasia has been the subject of several Neandertal studies. While previous studies generally have found high frequencies of enamel hypoplasia in Neandertals, the significance of this finding varies with frequencies of enamel hypoplasia in comparative samples. The present investigation was undertaken to ascertain if the enamel hypoplasia evidence in Neandertals suggests a high level of physiological stress relative to a modern human foraging group, represented here by an archaeological sample of Inuit from Point Hope, Alaska. Unlike previous studies, this study focused specifically on linear enamel hypoplasia (LEH), emphasizing systemic over localized causes of this defect by considering LEH to be present in an individual only if LEH defects occur on two anterior teeth with overlapping crown formation periods. Moreover, this study is the first to evaluate the average growth disruption duration represented by these defects in Neandertals and a comparative foraging group. In the prevalence analysis, 7/18 Neandertal individuals (from Krapina and southern France) and 21/56 Neandertal anterior teeth were affected by LEH, or 38.9% and 37.5% respectively. These values do not differ significantly from those of the Inuit sample in which 8/21, or 38.1% of individuals, and 32/111, or 28.8% of anterior teeth were affected. For the growth disruption duration analysis, 22 defects representing separate episodes of growth disruption in Neandertals were compared with 22 defects in the Inuit group using three indicators of duration: the number of perikymata (growth increments) in the occlusal walls of LEH defects, the total number of perikymata within them, and defect width. Only one indicator, the total number of perikymata within defects, differed significantly between the Inuit and Neandertal groups (an average of 13.4 vs. 7.3 perikymata), suggesting that if there is any difference between them, the Inuit defects may actually represent longer growth disruptions than the Neandertal defects. Thus, while stress indicators other than linear enamel hypoplasia may eventually show that Neandertal populations were more stressed than those of modern foragers, the evidence from linear enamel hypoplasia does not lend support to this idea.

Alaska↗

Nectar foraging behaviour is affected by ant body size in Camponotus mus.

The nectivorous ant Camponotus mus shows a broad size variation within the worker caste. Large ants can ingest faster and larger loads than small ones. Differences in physiological abilities in fluid ingestion due to the insect size could be related to differences in decision-making according to ant size during nectar foraging. Sucrose solutions of different levels of sugar concentration (30% or 60%w/w), viscosity (high or low) or flow rate (ad libitum or 1microl/min) were offered in combination to analyse the behavioural responses to each of these properties separately. Differences were found depending on ant body size and the property compared. A regulated flow produced smaller crop loads for medium and large ants compared to the same solution given ad libitum. All foragers remained longer times feeding at the regulated flow source but larger ants often made longer interruptions. When sugar concentration was constant but viscosity was high, only large ants increased feeding time. Constant viscosity with different sugar concentration determined longer feeding time and bigger loads for the most concentrated solution for small but not for large ants. Small ants reached similar crop loads in a variety of conditions while large ants did not. These differences could be evidence of a possible specialization for nectar foraging based on ant body size.

Animals↗