Juvenile hormone levels in honey bee (Apis mellifera L.) foragers: foraging experience and diurnal variation

A rising blood titer of juvenile hormone (JH) in adult worker honey bees is associated with the shift from working in the hive to foraging. We determined whether the JH increase occurs in anticipation of foraging or whether it is a result of actual foraging experience and/or diurnal changes in exposure to sunlight. We recorded all foraging flights of tagged bees observed at a feeder in a large outdoor flight cage. We measured JH from bees that had taken 1, 3-5, or >100 foraging flights and foragers of indeterminate experience leaving or entering the hive. To study diurnal variation in JH, we sampled foragers every 6h over one day. Titers of JH in foragers were high relative to nurses as in previous studies, suggesting that conditions in the flight cage had no effect on the relationship between foraging behavior and JH. Titers of JH in foragers showed no significant effects of foraging experience, but did show significant diurnal variation. Our results indicate that the high titer of JH in foragers anticipates the onset of foraging and is not affected by foraging experience, but is modulated diurnally.     

Foraging strategies are related to skull morphology and life history traits of Melanerpes woodpeckers

Links between morphology and foraging strategies have been well established for many vertebrate groups. Foraging strategies of Melanerpes woodpeckers are especially variable, with at least six species being proficient flycatchers; the remainder of the better known species do not flycatch. Our objective was to examine variation in foraging tactics as it relates to skull morphology and other life history traits among these species to better understand the biology of these diverse woodpeckers. We measured eight skull characters from 241 individuals representing 19 species, but focused on eight species for which we had the most data. We used the log-geometric mean and a principal components analysis (PCA) to calculate size-scaled shape variables. Cluster analysis based on PCA scores clearly separated birds by foraging behavior. Species with similar foraging behaviors (i.e., flycatchers vs non-flycatchers) also share a number of other life history characteristics including similar plumage, diets, and migratory behavior. Diversity within Melanerpes may imply a high degree of plasticity or that species have been incorrectly placed in a polyphyletic group. Woodpeckers currently in the genus Melanerpes share few uniting characters and historically have been placed in as many as eight different genera. Additional life history, morphological, and genetic studies of the group, especially of Caribbean and Neotropical species, is warranted.     

Intercolony variation in foraging flight characteristics of black‐headed gulls Chroicocephalus ridibundus during the incubation period

Using GPS loggers, we examined the influence of colony, sex, and bird identity on foraging flight characteristics of black‐headed gulls Chroicocephalus ridibundus during the incubation period. We studied tracks of 36 individuals breeding in one urban and two rural colonies in Poland. Birds from both rural colonies performed the furthest flights (mean max distance 8–12 km, up to 27 km) foraging mainly in agricultural areas. Gulls from the urban colony performed shorter flights (mean 5 km, up to 17 km) visiting mainly urbanized areas and water bodies. We found that females performed longer flights and their flight parameters were less repeatable compared to males. Males from both rural colonies visited water bodies more frequently than females. In all colonies, males (but not females) used habitats unproportionally to their availability in the vicinity. Relatively low interindividual and relatively high intraindividual overlap in home ranges indicated considerable foraging site fidelity. Individuals specialized in the use of a particular type of habitat performed shorter foraging flights compared to individuals using diverse habitats during their foraging flights. Our results indicate diverse foraging strategies of black‐headed gulls, including generalists that explore various habitats and specialists characterized by high foraging site and habitat fidelity.     

Foraging ecomorphology within North American flycatchers and a test of concordance with southern African species

In both the Old and New Worlds, independent clades of sit-and-wait insectivorous birds have evolved. These independent radiations provide an excellent opportunity to test whether the relationships between morphology and ecology are concordant among different communities of flycatchers. First, with canonical correlation analysis, I test for significant relationships between foraging behavior and morphology in North American (NA) and southern African (SA) flycatcher communities. Second, using ordination and analysis of covariance, I test for concordant ecomorphological relationships between the localities. Also, I accounted for phylogeny to see if observed patterns were adaptive. Morphology predicted the foraging behavior in both NA and SA flycatchers. Smaller bill features and longer legs are related to perching and attacking prey items near or on the ground. Larger bill features and short legs are associated with flycatching behavior and high, open perches. The ANCOVA reveals concordance in their ecomorphological relationships, but only in a single ecomorphological axis. The lack of complete concordance may be due to differences in habitat or broad scale historical influences between NA and SA flycatchers.     

Pollinator flight directionality and the assessment of pollen returns

Summary Nectar-foraging pollinators often exhibit a directional pattern of movement between plants when the energetic costs of revisiting previously utilized areas can significantly reduce foraging efficiency. However, bumblebees (Bombus spp.) foraging for pollen on flowers of Aquilegia caerulea rarely moved in a straight line among successively visited plants. Most flights from plants visited were either to closely neighboring plants or were longer and involved bypassing near neighbor plants. Bees biased their flights toward plants with relatively large numbers of flowers yet visited only a small fraction of the flowers on each plant. Such foraging tactics might result when the energetic costs of revisiting plants are minor. Alternatively we suggest that bumblebees foraging for pollen may not perceive revisitations and their associated costs because they do not assess pollen returns on a per plant basis. In this case energetic-efficiency arguments predicting the pattern of foraging movements among plants may be inappropriate. A better level of analysis would be where the bees assess net energy returns, perhaps between bouts of pollen-combing and corbiculae-packing.     

Radar Tracking and Motion-Sensitive Cameras on Flowers Reveal the Development of Pollinator Multi-Destination Routes over Large Spatial Scales

Central place foragers, such as pollinating bees, typically develop circuits (traplines) to visit multiple foraging sites in a manner that minimizes overall travel distance. Despite being taxonomically widespread, these routing behaviours remain poorly understood due to the difficulty of tracking the foraging history of animals in the wild. Here we examine how bumblebees (Bombus terrestris) develop and optimise traplines over large spatial scales by setting up an array of five artificial flowers arranged in a regular pentagon (50 m side length) and fitted with motion-sensitive video cameras to determine the sequence of visitation. Stable traplines that linked together all the flowers in an optimal sequence were typically established after a bee made 26 foraging bouts, during which time only about 20 of the 120 possible routes were tried. Radar tracking of selected flights revealed a dramatic decrease by 80% (ca. 1500 m) of the total travel distance between the first and the last foraging bout. When a flower was removed and replaced by a more distant one, bees engaged in localised search flights, a strategy that can facilitate the discovery of a new flower and its integration into a novel optimal trapline. Based on these observations, we developed and tested an iterative improvement heuristic to capture how bees could learn and refine their routes each time a shorter route is found. Our findings suggest that complex dynamic routing problems can be solved by small-brained animals using simple learning heuristics, without the need for a cognitive map.     

The skeleton flight apparatus of North American bluebirds (Sialia): Phylogenetic thrushes or functional flycatchers?

To better understand ecological traits of organisms, one can study them from two, not necessarily mutually exclusive perspectives: how the traits evolved, and their current adaptive utility. In birds, foraging behavior and associated morphological traits generally are explained by a combination of adaptive and phylogenetic predictors. The avian skeleton and more specifically, the skeletal flight apparatus is under well‐known functional and phylogenetic constraints. This is an interesting area to partition the relative contributions of adaptive correlated evolution and phylogenetic constraint to species clustering in morphological space. A prediction of convergent evolution is that nonphylogenetic morphological clustering is a characteristic of ecological similarity. We tested this using representatives of North American birds from two clades, one with a mixture of foraging modes (Turdid thrushes, solitaires, and bluebirds) and one with more canalized foraging behaviors (Tyrannid flycatchers). Nine characters on the skeletal flight apparatus from 19 species were used to characterize the morphological space and test for ecomorphological clustering. When body size and phylogeny are considered, the three bluebird species and Townsend's solitaire cluster with the ecologically similar flycatchers rather than with their phylogenetic close relatives. Furthermore, sit‐and‐wait foragers tend to exhibit relatively long distal elements and a long keel while active ground foragers have deeper keels and a longer humerus. Distal elements, expected to be relatively shorter and more bowed in the flycatchers and bluebirds, were actually longer and narrower. A reduction of distal element mass may be more important for facilitating maneuverability than surface area for insertion of wing‐rotational musculature. J. Morphol. 274:909–917, 2013. © 2013 Wiley Periodicals, Inc.     

Can environmental fluctuation prevent competitive exclusion in sympatric flycatchers?

Ecology has been characterized by a central controversy for decades: namely, whether the distribution and abundance of organisms are determined by species interactions, such as competitive exclusion, or by environmental conditions. In part, this is because competitive exclusion has not been convincingly demonstrated in open, natural systems. In addition, traditional theoretical models cannot predict the outcome of competitive interactions in the presence of environmental variability. In this paper we document the limiting influence of strong interspecific competition on population dynamics and nestling mortality in a mixed population of pied flycatchers (Ficedula hypoleuca) and collared flycatchers (F. albicollis in a narrow zone of sympatry. Whereas the former species was limited mainly by interspecific competition, the latter species was limited by the concerted influences of intraspecific competition and climate. The analysis suggests a march towards competitive exclusion of the pied flycatcher during warm periods. However, competitive exclusion is apparently prohibited on a local scale because intraspecific competition among individual collared flycatchers intensifies when they are forced to cope with severe environmental conditions, promoting the temporary and local presence of pied flycatchers.     

Wintering birds avoid warm sunshine: predation and the costs of foraging in sunlight

Wintering birds can gain significant thermal benefits by foraging in direct sunlight. However, exposure to bright sunlight might make birds easier to detect by predators and may also cause visual glare that can reduce a bird’s ability to monitor the environment. Thus, birds likely experience a trade-off between the thermal benefits and predation-related costs of foraging in direct sunlight. To examine this possible thermoregulation-predation trade-off, we monitored the behavior of mixed-species flocks of wintering emberizid sparrows foraging in alternating strips of sunlight and shade. On average, these sparrows routinely preferred to forage in the shade, despite midday air temperatures as much as 30 °C below their thermoneutral zone. This preference for shade was strongest at relatively high temperatures when the thermal benefits of foraging in sunlight were reduced, suggesting a thermoregulation-predation trade-off. Glare could be reduced if birds faced away from the sun while feeding in direct sunlight, but we found that foraging birds tended to face southward (the direction of the sun). We speculate that other factors, such as the likely direction of predator approach, may explain this southerly orientation, particularly if predators use solar glare to their advantage during an attack. This interpretation is supported by the fact that birds had the weakest southerly orientation on cloudy days. Wintering birds may generally avoid foraging in direct sunlight to minimize their risk of predation. However, given the thermal benefits of sunshine, such birds may benefit from foraging in habitats that provide a mosaic of sunlit and shaded microhabitats.     

Central place water collection in a Japanese paper wasp, Polistes Chinensis antennalis

The water collection behaviour of colony foundresses of a Japanese paper wasp, Polistes chinensis antennalis, was recorded. A single round-trip of water collection consists of (1) an outward flight, (2) water-sucking, and (3) a homeward flight. The rate of water intake decreased as the length of water-sucking increased. There was a positive correlation between the duration of the outward flight and that of water-sucking. The variance of time for outward flights was larger than that for homeward flights. The data were consistent with the predictions of the central place foraging model of Orians & Pearson (1979).     

Search strategies of tyrant flycatchers

Aspects of searching behaviour among free-living South American flycatchers (Aves: Tyrannidae) are compared quantitatively. Flycatchers forage with stationary searching periods, followed either by an attempted prey capture (sally) or a ‘give-up’ flight to a new perch. Search times are proportional to body size within each of three categories of foraging behaviour: aerial hawking, sally-gleaning, and perch-gleaning. Over the family as a whole, search times are directly proportional to the size of the visual field scanned during the search. Intraspecific variations in search times are caused by local variations in prey density or visual complexity of the habitat. Between foraging modes, differences in searching and movement patterns are related to prey dispersion characteristics. Aerial hawkers regularly return to favoured perches, but foliage gleaners, which reduce the resources surrounding a perch by sallying only once, rarely return to a perch. In contrast to aerial hawkers, foliage gleaners appear to follow an organized scanning procedure on each perch, by searching nearby surfaces before they examine more distant prey substrates. Throughout the family, the median flight distance after a perch is abandoned is approximately twice the median search radius. Comparisons of search time distributions preceding sallies with those preceding give-up flights suggest that there is no single, optimal give-up time in a given habitat. Foliage-gleaning species appear to assess the amount of search time each perch warrants, presumably based on the degree of complexity of the search area. They either sally at prey before that time, or give-up when the allotted time has elapsed.     

On a wing and a prayer: the foraging ecology of breeding Cape cormorants

The Cape cormorant Phalacrocorax capensis is unusual among cormorants in using aerial searching to locate patchily distributed pelagic schooling fish. It feeds up to 80 km offshore, often roosts at sea during the day and retains more air in its plumage and is more buoyant than most other cormorants. Despite these adaptations to its pelagic lifestyle, little is known of its foraging ecology. We measured the activity budget and diving ecology of breeding Cape cormorants. All foraging took place during the day, with 3.6 ± 1.3 foraging trips per day, each lasting 85 ± 60 min and comprising 61 ± 53 dives. Dives lasted 21.2 ± 13.9 s (maximum 70 s), attaining an average depth of 10.2 ± 6.7 m (maximum 34 m), but variability in dive depth both within and between foraging trips was considerable. The within-bout variation in dive depth was greater when making shallow dives, suggesting that pelagic prey were targeted mainly when diving to <10 m. Diving ecology and total foraging time were similar to other cormorants, but the time spent flying (122 ± 51 min day⁻¹, 14% of daylight) was greater and more variable than other species. Searching flights lasted up to 1 h, and birds made numerous short flights during foraging bouts, presumably following fast-moving schools of pelagic prey. Compared with the other main seabird predators of pelagic fish in the Benguela region, Cape gannets Morus capensis and African penguins Spheniscus demersus , Cape cormorants made shorter, more frequent foraging trips. Their foraging range while feeding small chicks was 7 ± 6 km (maximum 40 km), similar to penguins (10–20 km), but less than gannets (50–200 km). Successful breeding by large colonies depends on the reliable occurrence of pelagic fish schools within this foraging range.     

Foraging habitat shift in the narcissus flycatcher, Ficedula narcissina, due to the response of herbivorous insects to the strengthening defenses of canopy trees

The indirect effect of tree defense on bird foraging habitat via lepidopterous larvae was studied by focusing on phenological changes in each component. The seasonal changes in foraging habitat of the narcissus flycatcher (Ficedula narcissina), the distribution pattern of arthropod populations, and the leaf characteristics related to the defense system against insect herbivores were studied in a cool-temperate deciduous forest in northern Japan. Narcissus flycatchers foraged in the canopy from late May to mid June, on the ground from late June, with lepidopterous larvae being the most preferred prey (about 80% of total prey items), and again in the canopy in July. The biomass of lepidopterous larvae was much more abundant within 3 weeks after budbreak in the canopy in late May, and thereafter decreased rapidly in mid June. By contrast, the larvae were abundant on the forest floor from early to late June. These were mainly caused by their downward migration in early to mid June. After budbreak, the leaf toughness and tannin content of two deciduous tree species (maple and oak) increased, while nitrogen and water content decreased. The strengthening of defense traits of canopy leaves caused changes in the abundance and distribution of the lepidopterous larvae. The foraging habitat of the flycatcher subsequently shifted in response to the changes in abundance and distribution of the lepidopterous larvae. In this study, the evidence of the indirect effects of trees on bird foraging habitat was presented by focusing on the phenological changes in relationships among each component in this three-trophic system.     

Prey selection and the search strategy of the spotted flycatcher (Muscicapa striata): A field study on optimal foraging

Flycatchers switched between different foraging strategies and selected prey so as to maximize energy intake. Changes in diet were influenced by the absolute abundance of the preferred prey and not by that of the alternative prey. Adults could recognize wasps (Vespula) and bees (Bombus) and removed their stings before swallowing them. The differences between adult and nestling diet and factors influencing capture success of prey are described. The flycatchers' behaviour in leaving perches and their choice of a ‘giving-up time’ was consistent with the view that they were maximizing the number of prey they caught in a given time. I conclude that flycatchers often search for and select prey on the basis of maximizing energy intake but I give reasons for not expecting them always to do so.     

Phenetic relationships among four Apodemus species (Rodentia, Muridae) inferred from skull variation

Phenetic relationships among four Apodemus species (A. agrarius, A. epimelas, A. flavicollis and A. sylvaticus) inferred from skull (mandible and cranium) variation were explored using landmark-based geometric morphometrics. Analysis of size variation revealed that mandibles and crania of A. epimelas were the largest, followed by those of A. flavicollis, while A. agrarius and A. sylvaticus had the smallest ones. Phenetic relationships inferred from mandible shape variation better reflected phylogenetic relationships among the analyzed Apodemus species than those inferred from cranial differences. Concerning cranial shape variation, the most differentiated species was A. epimelas, whose ecology clearly differs from the other three species. Thus, differentiation of the mandible provided a pattern fully concordant with the phylogeny, while the cranium differentiation was in agreement with ecology expectations. The most evident shape changes of mandible and cranium involved the angular process and facial region, respectively. We also found that allometry had a significant influence on shape variation and that size-dependent shape variation differed among the analyzed species. Moreover, mandible and cranium are differently influenced by allometric changes. Different phenetic relationships inferred from mandible and cranium shape variation imply that phylogeny, ecology, together with factors related to size differences are all involved in the observed morphological divergence among the analyzed Apodemus species.     

Foraging ecology drives social information reliance in an avian eavesdropping community

Vertebrates obtain social information about predation risk by eavesdropping on the alarm calls of sympatric species. In the Holarctic, birds in the family Paridae function as sentinel species; however, factors shaping eavesdroppers' reliance on their alarm calls are unknown. We compared three hypothesized drivers of eavesdropper reliance: (a) foraging ecology, (b) degree of sociality, and (c) call relevance (caller‐to‐eavesdropper body‐size difference). In a rigorous causal‐comparative design, we presented Tufted Titmouse (Baeolophus bicolor) alarm calls to 242 individuals of 31 ecologically diverse bird species in Florida forests and recorded presence/absence and type (diving for cover or freezing in place) of response. Playback response was near universal, as individuals responded to 87% of presentations (N = 211). As an exception to this trend, the sit‐and‐wait flycatcher Eastern Phoebe (Sayornis phoebe) represented 48% of the nonresponses. We tested 12 predictor variables representing measures relevant to the three hypothesized drivers, distance to playback speaker, and vulnerability at time of playback (eavesdropper's microhabitat when alarm call is detected). Using model‐averaged generalized linear models, we determined that foraging ecology best predicted playback response, with aerial foragers responding less often. Foraging ecology (distance from trunk) and microhabitat occupied during playback (distance to escape cover) best predicted escape behavior type. We encountered a sparsity of sit‐and‐wait flycatchers (3 spp.), yet their contrasting responses relative to other foraging behaviors clearly identified foraging ecology as a driver of species‐specific antipredator escape behavior. Our findings align well with known links between the exceptional visual acuity and other phenotypic traits of flycatchers that allow them to rely more heavily on personal rather than social information while foraging. Our results suggest that foraging ecology drives species‐specific antipredator behavior based on the availability and type of escape cover.     

The composition and foraging behaviour of mixed-species flocks of forest-living birds in Madagascar

Mixed-species flocks of forest living birds were investigated in a rainforest at Perinet/ Analamazaotra (Andasibe), Madagascar. Most insectivorous birds in the area participated in mixed-species flocks. Flocks were composed of foliage gleaners, foliage-branch gleaners, trunk gleaners and flycatchers. Species whose foraging techniques and foraging locations in vegetation were similar were different from each other in foraging heights. The nuclear species often occurred in small conspecific flocks in and out of mixed-species flocks.     

Foraging behavior of individual workers and foraging dynamics of colonies of three Sumatran stingless bees

The foraging behavior of three stingless bees, Trigona (Tetragonula) minangkabau, T. (Trigonella) moorei and T. (Heterotrigona) itama, was studied to describe patterns of resource harvest in disturbed forest areas in Sumatra, Indonesia.

1. Average daily total number of foraging flights per colony was 1200 in T. minangkabau, 2400 in T. moorei and 7000 in T. itama and it was proportional to colony population size. Foragers collecting nectar, pollen or plant resin were respectively 70–80%, 10–20% and <10% in the three species. Pollen was collected most in the morning. Nectar collection peaked in midday in T. itama but continued almost evenly until dusk in T. minangkabau and T. moorei. Resin was collected evenly throughout day.
2. In all the three species the volume of a nectar load carried by a returning forager did not decrease until 1600, followed by slight decline. In the morning the sugar concentration of nectar was almost constantly 30%, and in the afternoon its maximum value reached 60% although some remained 30%. As a result, the mean sugar weight in a nectar load gradually increased until dusk.
3. In T. minangkabau, resin was collected by specialized foragers. Nectar foragers switched to collect pollen and vice versa. There were two types of foraging: “exploitatory” flights, the repitition of short, rather constant flights bringing back full resource loads, and “exploratory” flights, prolonged flight and/or reduction in amount of resources carried. Exploitatory flights followed the exploratory flights which led to discovery of rich sources. Mean duration of exploitatory flights was 7 min during nectar collection, 12 min for pollen and 23 min for resin. Sites of exploited flowers were estimated to be 84–434 m distant from the nest site. Mean duration of flights for nectar collection was 13 min in the morning and 6 min in the late afternoon.

     

Seasonal Activity and Foraging Preferences of the Leaf-Cutting Ant Atta sexdens piriventris (Santschi) (Hymenoptera: Formicidae)

Leaf-cutting ants are known for their habit of foraging. These habits can be influenced by several factors, including variations in topography, soil, and climate among others. The objective of this research was to study the seasonal activity and foraging preferences of the leaf-cutting ant Atta sexdens piriventris (Santschi). The study was carried out from January to October of 2007 in grasslands of the state of Santa Catarina, Brazil. Nests of A. sexdens piriventris were randomly selected and the forager’s activities were recorded. Damaged plant species, soil fertility, and climatic conditions were recorded. The maximum foraging activity in the summer was recorded during periods of darkness or low light (between 7 p.m. and 5 a.m.). In autumn and spring, foraging was the highest in the morning, but during winter time the foraging activity was high in the afternoon. Fourteen plant species were frequently visited during our study with the lowest foraging activity being recorded at very low (1 to 4°C) or very high temperatures (above 26°C). The number of exploited plant species was higher in winter (13) than in summer (9). Baccharis trimera (Asteraceae) (43%) and Paspalum spp. (Poaceae) (33%) were the most exploited plants among the identified species.     

Stream meanders increase insectivorous bird abundance in riparian deciduous forests

Adult aquatic insects emerging from streams are a fundamental resource sustaining riparian bird communities in broad-leaved deciduous forests. We investigated how stream geomorphology affects the aquatic insect flux and insectivorous bird abundance in 26 riparian-forest plots during spring season in northern Japan. Lateral dispersal of emergent aquatic insects into the riparian forest exponentially decreased with distance from the stream. Similar to aquatic insect distribution, flycatchers and gleaners concentrated their foraging attacks around the stream channel, preying intensively upon emergent aquatic insects. In contrast, bark probers consumed fewer emergent aquatic insects. The abundance of flycatchers and gleaners was closely related to stream geomorphology, whereas that of bark probers was associated with snag density in the study plots. A path analysis showed that the study plots with longer stream channels had greater aquatic insect abundance. This can be interpreted as a consequence of the increased amount of both stream edge and stream surface, where emergent aquatic insects readily penetrate. The increased flux of aquatic insects by stream meanders elevated gleaner abundance in the study plots. In addition, their abundance was directly affected by stream length per se. On the other hand, flycatcher abundance was only directly affected by stream length. Flycatchers, which mainly consumed emergent aquatic insects in the air, may have increased in response to the increase in suitable foraging sites (i.e., open spaces adjacent to perches) accompanying longer stream channels. Although the causal links affecting bird abundance differed among guilds, meandering streams apparently support abundant insectivorous birds in riparian forests. Therefore, to conserve riparian bird communities, it will be necessary to maintain the functions of stream geomorphology that affect the magnitude of energy transfer across the forest-stream interface.