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Visual constraints in foraging bumblebees: flower size and color affect search time and flight behavior.

In optimal foraging theory, search time is a key variable defining the value of a prey type. But the sensory-perceptual processes that constrain the search for food have rarely been considered. Here we evaluate the flight behavior of bumblebees (Bombus terrestris) searching for artificial flowers of various sizes and colors. When flowers were large, search times correlated well with the color contrast of the targets with their green foliage-type background, as predicted by a model of color opponent coding using inputs from the bees' UV, blue, and green receptors. Targets that made poor color contrast with their backdrop, such as white, UV-reflecting ones, or red flowers, took longest to detect, even though brightness contrast with the background was pronounced. When searching for small targets, bees changed their strategy in several ways. They flew significantly slower and closer to the ground, so increasing the minimum detectable area subtended by an object on the ground. In addition, they used a different neuronal channel for flower detection. Instead of color contrast, they used only the green receptor signal for detection. We relate these findings to temporal and spatial limitations of different neuronal channels involved in stimulus detection and recognition. Thus, foraging speed may not be limited only by factors such as prey density, flight energetics, and scramble competition. Our results show that understanding the behavioral ecology of foraging can substantially gain from knowledge about mechanisms of visual information processing.

Animals↗

Phase transition between disordered and ordered foraging in Pharaoh's ants.

The complex collective behavior seen in many insect societies strongly suggests that a minimum number of workers are required for these societies to function effectively. Here we investigated the transition between disordered and ordered foraging in the Pharaoh's ant. We show that small colonies forage in a disorganized manner, with a transition to organized pheromone-based foraging in larger colonies. We also show that when food sources are difficult to locate through independent searching, this transition is first-order and exhibits hysteresis, comparable to a first-order phase transition found in many physical systems. To our knowledge, this is the first experimental evidence of a behavioral phase transition between a maladaptive (disorganized) and an adaptive (organized) state.

Animals↗

Enemy deterrence in the recruitment strategy of a termite: Soldier-organized foraging in Nasutitermes costalis.

The nasute soldiers of the neotropical termite Nasutitermes costalis function as scouts by exploring new terrain for food in advance of the worker caste and regulate foraging activity by laying trails composed of sternal gland pheromone. Additional soldiers are at first recruited in large numbers, and subsequently workers appear as the pheromone concentration increases. The role of the nasutes in the organization of foraging is extremely unusual for the soldier caste in social insects and appears to be a component of a foraging/defense system that controls the recruitment of foragers and effectively deters attacks by ants, the most fierce and important predators of termites.

Journal Article↗

Foraging ability of rufous hummingbirds on hummingbird flowers and hawkmoth flowers.

We examine the suitability of ornithophilous flowers and sphingophilous flowers in Ipompsis and Aquilegia for nectar foraging by the hummingbird Selasphorus rufus. In S. rufus, bill length averages 18.9 mm in females and 17.3 mm in males. Maximal tongue extension approximates bill length, suggesting that birds can feed from floral tubes up to 33.5 mm in length. However, their ability to do so is limited by two factors. First, the maximal depth at which S. rufus can extract nectar decreases with the width of the floral tube. Second, feeding time is shortest in short floral tubes and progressively increases as the tubes lengthen because of increased time required for tongue extension and retraction. Hence, nectar foraging occurs with optimal efficiency in moderately broad floral tubes with lengths that do not exceed or only slightly exceed the bill length plus </=5-mm tongue extension. In most ornithophilous taxa of Ipomopsis and Aquilegia, the floral tubes have lengths and widths in the optimal range for nectar foraging by S. rufus, whereas in the sphingophilous taxa, the basal nectar either may be reached but at much higher cost or, more frequently, is beyond reach of bills and tongues. The flower-visiting habits of S. rufus and other hummingbirds in nature are generally congruent with these conclusions and support the case for coadaptation between these plants and pollinators.

Journal Article↗

Mutations in the larval foraging gene affect adult locomotory behavior after feeding in Drosophila melanogaster.

Previous studies have shown a correlation between the locomotory component of larval and adult foraging behavior in the fruit fly. Here we show that this relationship is far more than mere correlation. It can be attributable to different alleles at the same genetic locus of the behavioral gene foraging (for). The for gene offers us the unique opportunity to study the genetic basis and evolutionary significance of a naturally occurring behavioral polymorphism. Until now, only the effect of for on Drosophila melanogaster larval behavior was studied. Larvae with the rover allele (forR) move significantly more while eating during a set time period than those homozygous for the sitter alleles (fors). Here, we show that rover and sitter larval strains derived from nature differ in the distance adults walk after feeding per unit time and that this variation results from different alleles at the foraging locus, the very gene originally defined on the basis of larval behavior. We hypothesize that for may be involved in the way flies evaluate a food resource.

Alleles↗

Evolution of foraging behavior in Drosophila by density-dependent selection.

One of the rare examples of a single major gene underlying a naturally occurring behavioral polymorphism is the foraging locus of Drosophila melanogaster. Larvae with the rover allele, forR, have significantly longer foraging path lengths on a yeast paste than do those homozygous for the sitter allele, fors. These variants do not differ in general activity in the absence of food. The evolutionary significance of this polymorphism is not as yet understood. Here we examine the effect of high and low animal rearing densities on the larval foraging path-length phenotype and show that density-dependent natural selection produces changes in this trait. In three unrelated base populations the long path (rover) phenotype was selected for under high-density rearing conditions, whereas the short path (sitter) phenotype was selected for under low-density conditions. Genetic crosses suggested that these changes resulted from alterations in the frequency of the fors allele in the low-density-selected lines. Further experiments showed that density-dependent selection during the larval stage rather than the adult stage of development was sufficient to explain these results. Density-dependent mechanisms may be sufficient to maintain variation in rover and sitter behavior in laboratory populations.

Animals↗

Finding food in the open ocean: foraging strategies in Humboldt penguins.

Penguins are excellent "model" organisms allowing us to study the behaviour of marine homeotherms at sea. Penguins regularly return to their breeding colonies, enabling biologists to equip them with remote sensing devices such as physiological or behavioural data-loggers, radio- or satellite transmitters. Foraging trips at sea can last from days to weeks and after return of the birds to their breeding sites, the devices can easily be removed for analysis of on-board stored data, yielding a wealth of information. Investigation of penguin behaviour at sea becomes particularly revealing when other sources of information can be matched to the data set, such as satellite data on wind, temperature, ice cover, and chlorophyll-a concentrations. Penguins and other marine homeotherms are true inhabitants of the high seas. Depending on the season, the marine behaviour varies: during reproduction, penguins are central-place foragers, and must return regularly to their nest to feed their chicks. During the remainder of the year, there are no constraints and the birds travel large distances at sea. Breeding Humboldt penguins react to climatic change by varying their daily foraging range and dive duration. Similar to other representatives of the family Spheniscidae, Humboldt penguins avoid food shortages by migrating into more productive marine areas. Navigational clues such as daylength, sea surface temperature, local wind direction and olfaction might provide important aids in finding patchily distributed prey in the open ocean. DMS, a chemical compound produced by decaying algae, seems to be a further clue that indirectly points the way to feeding areas.

Journal Article↗

Tunnel specificity and forager movement in subterranean termites (Isoptera: Rhinotermitidae and Termitidae).

The movement of foragers of two species of Australian, subterranean, mound-building termites, Coptotermes lacteus (Froggatt) (Rhinotermitidae) and Nasutitermes exitiosus (Hill) (Termitidae), was investigated in their natural habitat using artificial feeding sites along trenches dug to mimic natural forager tunnels that radiate out from the central mound-nests. Termites were dyed by self-feeding on cardboard soaked with histological fat-stains on one or two trenches and then termites were collected from other feeding sites at two and four weeks after the fat-stains were placed. At two and four weeks after marking commenced, 60-75% of marked termites were found in trenches containing the marked paper, and 2-16% were found in trenches on the opposite side of the nest. The proportion of marked termites in a sample was three to eight times greater in the trenches containing the marked paper relative to other trenches. Although difficulties with fat-stains used as markers might explain some of the observed patterns, it is evident that C. lacteus and N. exitiosus foragers do not move randomly between feeding sites in their natural habitat.

Animals↗

Comparing mark-recapture and constant removal protocols for estimating forager population size of the subterranean termite Coptotermes lacteus (Isoptera: Rhinotermitidae).

Forager population sizes of colonies of Coptotermes lacteus(Froggatt) in New South Wales were estimated using two methods: mark-recapture and constant removal, in two disturbed habitats: a pine plantation and cleared farmland. Mark-recapture population estimates were unrealistic and unreliable: they could be improbably large, over 200 million foragers, and they varied enormously between samples for each colony without any pattern. The constant removal population estimates could also be unrealistic: they could be negative or quite different when calculated using regression and maximum likelihood methods. However, the unrealistic results could be predicted reliably, and explained by the lack of re-contact with the sampling devices (bait stations) - a violation of an assumption of the method. This happened more frequently in the plantation than in the farmland, probably because of the greater abundance of alternative food sources in the plantation. Of the two methods, constant removal provided reasonable forager population estimates, relative to direct counts, at least some of the time, plus a mechanism by which reliability could be tested, whereas mark-recapture provided neither. Further refinement and testing of constant removal methods are urged to provide a more reliable population estimation technique for termites.

Animals↗

Foraging and denning patterns of brushtail possums, and their possible relationship to contact with cattle and the transmission of bovine tuberculosis.

Radio-tracking and direct observation were used over 18 months in 1990-92 to investigate both the use of sleeping dens and foraging activity by possums (Trichosurus vulpecula) on a 21 ha site in the Wairarapa used for a longitudinal study of bovine tuberculosis. Males had larger home ranges than females, and both sexes had larger activity areas during the autumn mating season than at other times of the year. Possums typically foraged in only a small area of their home ranges (termed an activity area) on any one night, and the areas used by individuals were commonly very similar over a series of nights. Activity areas overlapped extensively among possums. Possums used a limited number of dens, commonly in a small and in most cases a circumscribed part of their home range. No simultaneous den-sharing was found, with the exception of mother-joey pairs. The mortality of juveniles after independence was 36%. Only two of 25 juveniles under surveillance to detect dispersal dispersed more than 500 m off the study site, and both subsequently died. Grazing patterns of cattle meant that almost all accessible areas of the paddock were covered by at least some grazing cattle, and so all activity areas of possums within the paddock were covered by areas where cattle foraged. However, possums avoided contact with cattle, and when some cattle were excluded from access to the part of the paddock principally used by both tuberculous and healthy possums for denning, transmission of Mycobacterium bovis from possums to these cattle ceased, although there was subsequent transmission to deer. Cattle which grazed the area used principally for possum denning continued to become infected, and these denning areas appeared to be of importance in the transmission of tuberculosis.

Journal Article↗

[Investigations on the fermentation quality of silage from low-nitrate green forage].

In 3-year-experiments green forage from Dactylis glomerata and Festuca species, produced by differentiated cutting frequencies, was ensiled under laboratory conditions. The forage was almost without nitrate. Because of the higher content of water-soluble carbohydrates the fermentability coefficients of Festuca species were above of Dactylis glomerata. The fermentability coefficients of both plant species were inferior in the second and third cut, even if the first cut was delayed until July. Despite of similar fermentability coefficients of the two plant species silages contained different amounts of lactic and butyric acid. The portion of lactic acid amounted to 52 to 57% and 60 to 73% of total acids, the proportion of butyric acid 30 to 35% and 18 to 25% for Dactylis glomerata and Festuca species, respectively. According to the high fermentability coefficients silages free of butyric acid would be expected, however, by both species butyric acid always was formed, in some cases at high concentrations. In the contrary the amounts of ammonia, acetic acid and alcohol as well as the pH-value were almost extremely low. It can be concluded that the quality of silages produced of green forages with low nitrate cannot be assessed adequately by the current evaluation key.

Acetic Acid↗

Digestibility, nitrogen excretion, and mean retention time by North American porcupines (Erethizon dorsatum) consuming natural forages.

North American porcupines (Erethizon dorsatum) subsist predominantly on low-protein, high-fiber, high-tannin diets. Therefore, we measured the porcupine's ability to digest dry matter, fiber, and protein by conducting digestion trials on eight natural forages and one pelleted ration varying in concentration of fiber, nitrogen, and tannins. On these diets, dry matter intake ranged from 5 to 234 g/kg(0.75)/d and dry matter digestibility ranged from 62% to 96%. Porcupines digested highly lignified fiber better than many large hindgut fermenters and ruminants. The porcupine's ability to digest fiber may be explained, in part, by their lengthy mean retention time of particles (38.43+/-0.56 h). True nitrogen digestibility was 92% for nontannin forages and pellets. Endogenous urinary nitrogen was 205 mg N/kg(0.75)/d, and metabolic fecal nitrogen was 2.8 g N/kg dry matter intake. Porcupines achieved nitrogen balance at relatively low levels of nitrogen intake (346 mg N/kg(0.75)/d). Tannins reduced the porcupines' ability to digest protein. However, the reduction in protein digestion was not predictable from the amount of bovine serum albumin precipitated. Like many herbivores, porcupines may ameliorate the effects of certain tannins in natural forages on protein digestibility through physiological and behavioral adaptations.

Animals↗

Of hummingbirds and helicopters: hovering costs, competitive ability, and foraging strategies.

Wing morphology and flight kinematics profoundly influence foraging costs and the overall behavioral ecology of hummingbirds. By analogy with helicopters, previous energetic studies have applied the momentum theory of aircraft propellers to estimate hovering costs from wing disc loading (WDL), a parameter incorporating wingspan (or length) and body mass. Variation in WDL has been used to elucidate differences either among hummingbird species in nectar-foraging strategies (e.g., territoriality, traplining) and dominance relations or among gender-age categories within species. We first demonstrate that WDL, as typically calculated, is an unreliable predictor of hovering (induced power) costs; predictive power is increased when calculations use wing length instead of wingspan and when actual wing stroke amplitudes are incorporated. We next evaluate the hypotheses that foraging strategy and competitive ability are functions of WDL, using our data in combination with those of published sources. Variation in hummingbird behavior cannot be easily classified using WDL and instead is correlated with a diversity of morphological and physiological traits. Evaluating selection pressures on hummingbird wings will require moving beyond wing and body mass measurements to include the assessment of the aerodynamic forces, power requirements, and power reserves of hovering, forward flight, and maneuvering. However, the WDL-helicopter dynamics model has been instrumental in calling attention to the importance of comparative wing morphology and related aerodynamics for understanding the behavioral ecology of hummingbirds.

Animals↗

Genetic localization of foraging (for): a major gene for larval behavior in Drosophila melanogaster.

Localizing genes for quantitative traits by conventional recombination mapping is a formidable challenge because environmental variation, minor genes, and genetic markers have modifying effects on continuously varying phenotypes. We describe "lethal tagging," a method used in conjunction with deficiency mapping for localizing major genes associated with quantitative traits. Rover/sitter is a naturally occurring larval foraging polymorphism in Drosophila melanogaster which has a polygenic pattern of inheritance comprised of a single major gene (foraging) and minor modifier genes. We have successfully localized the lethal tagged foraging (for, 2-10) gene by deficiency mapping to 24A3-C5 on the polytene chromosome map.

Animals↗

Effects of clothianidin on Bombus impatiens (Hymenoptera: Apidae) colony health and foraging ability.

We conducted laboratory experiments to investigate the lethal and sublethal effects of clothianidin on bumble bee, Bombus impatiens Cresson, colony health and foraging ability. Bumble bee colonies were exposed to 6 ppb clothianidin, representing the highest residue levels found in field studies on pollen, and a higher dose of 36 ppb clothianidin in pollen. Clothianidin did not effect pollen consumption, newly emerged worker weights, amount of brood or the number of workers, males, and queens at either dose. The foraging ability of worker bees tested on an artificial array of complex flowers also did not differ among treatments. These results suggest that clothianidin residues found in seed-treated canola and possibly other crops will not adversely affect the health of bumble bee colonies or the foraging ability of workers.

Animals↗

Response of Argentine ants and red imported fire ants to permethrin-impregnated plastic strips: foraging rates, colonization of potted soil, and differential mortality.

This study investigated the effects of the permethrin-impregnated plastic on ant mortality and foraging rates, and tested its potential for preventing ants from colonizing potted soil. Direct exposure to the plastic for as short as 1 min caused significant mortality of both red imported fire ants, Solenopsis invicta Buren, and Argentine ants, Linepithema humile (Mayr); however, red imported fire ants were more susceptible than Argentine ants. Knockdown of virtually all ants initially occurred within 15 min after exposure. However, some moribund ants recovered from the effects within 24 h. For example, after 1 min of direct exposure to the permethrin-impregnated plastic, 70% of Argentine ants and 5% of red imported fire ants recovered from the treatment. In established colonies of Argentine ants, significantly fewer ants foraged for food up posts treated with the plastic compared with untreated posts. In addition, colonies responded to introduction of the treatment by significantly reducing their overall foraging rates, even on untreated posts. When pots filled with moistened soil were introduced into established ant colonies, 82% of Argentine ants and 99% of red imported fire ants moved into the soil. In contrast, when a 1-cm-wide coil of the plastic was placed under the pot, no ants moved into the soil. The potential for use of these materials in nursery production is discussed.

Animals↗

Low genetic variation among killer whales (Orcinus orca) in the eastern north Pacific and genetic differentiation between foraging specialists.

Killer whales from the coastal waters off California through Alaska were compared for genetic variation at three nuclear DNA markers and sequenced for a total of 520 bp from the mitochondrial control region. Two putative sympatric populations that range throughout this region were compared. They can be distinguished by social and foraging behavior and are known as "residents" and "transients". We found low levels of variation within populations compared to other cetacean species. Comparisons between fish (resident) versus marine mammal (transient) foraging specialists indicated highly significant genetic differentiation at both nuclear and mitochondrial loci. This differentiation is at a level consistent with intraspecific variation. A comparison between two parapatric resident populations showed a small but fixed mtDNA haplotype difference. Together these data suggest low levels of genetic dispersal between foraging specialists and a pattern of genetic differentiation consistent with matrifocal population structure and small effective population size.

Alleles↗

Feeding green-cut forage contaminated by radioactive fallout to dairy cows.

Grass contaminated by radioactive fallout from the Chernobyl accident was cut with 150 mm (Treatment 1) and 50 mm (Treatment 2) stubble height and fed to 20 dairy cows. The transfer of 137Cs and 131I from the greencut forage to milk was studied during a four-week and a two-week period, respectively. The four-week period was followed by a three-week period with nearly uncontaminated feeds. Cutting with high stubble height significantly reduced the activity concentration in forage compared with low stubble cutting, the average contents kg-1 dry matter being 385 Bq 137Cs and 24 Bq 131I in Treatment 1 versus 6656 Bq 137Cs and 249 Bq 131I in Treatment 2. Apparent treatment effects were also demonstrated by the 137Cs concentration in milk, with maximum weekly means of 23 Bq kg-1 (Treatment 1) and 92 Bq kg-1 (Treatment 2). Single day maximum concentrations of 131I in milk were 4 Bq kg-1 (Treatment 1) and 9 Bq kg-1 (Treatment 2). The mean transfer coefficients (d kg-1) were calculated to be 0.67 X 10(-2) and 0.19 X 10(-2) for 137Cs (weeks 2-4) and 1.8 X 10(-2) and 0.2 X 10(-2) for 131I (week 2) in Treatments 1 and 2, respectively. The lower value for 137Cs might possibly be associated with an enhanced ash content of the forage. Feeding nearly uncontaminated rations rapidly decreased the 137Cs levels in milk in both treatments with calculated effective half-times over the three-week period of 10 and 7 d, respectively.

Accidents↗