Determinants of foraging profitability in two nectarivorous butterflies

ABSTRACT.

• 1 I studied flower selection and foraging energetics of Agraulis vanillae L. (Nymphalidae) and Phoebis sennae (Pieridae), two butterfly species common to north central Florida. I identified the major nectar resources exploited by several populations of these butterflies and, for each plant species, measured available nectar volumes and concentrations, corolla lengths, and density. I quantified foraging behaviour of each butterfly species at each nectar source (flower visitation rate and percentage of foraging time in flight), and used these data to estimate the net rate of energy intake of each butterfly species at each nectar source.
• 2 Estimated mean energy contents of individual flowers of the eleven exploited plant species spanned three orders of magnitude, ranging between 0.015 and 9.27 joules. Mean energy content of individual flowers was strongly correlated with mean foraging profit of both butterfly species.
• 3 Mean nectar volume strongly influenced energy content and varied widely within and among species, ranging from 0.0076 to 1.853 μ1. Nectar concentration varied between 17.1% and 40.4% sucrose-equivalents. Nectar volume was the best single predictor of foraging profitability (correlation coefficients of 0.994 and 0.984 for Phoebis and Agraulis respectively). Corolla length also strongly affected foraging profitability for both butterfly species; flower species with longer corollas were generally more profitable.
• 4 Flower density and nectar concentration showed weak or nonsignificant associations with foraging profitability.
• 5 The usefulness and limitations of these floral characteristics as bases for foraging selectivity, and the selective pressures foraging butterflies might place on the visited plants are discussed.

     

Foraging rate of ants collecting honeydew or extrafloral nectar, and some possible constraints

ABSTRACT.

• 1 In a given ant species, the number of ants collecting honeydew in an aphid colony or extrafloral nectar on a plant is proportional to the productivity of the colony or plant. Thus, the number of ants per resource unit and the ingestion rate per ant are constant for a species.
• 2 Mean number of ants per resource unit and ingestion rate per ant differed considerably between the investigated species. The ingestion rate increases with the body size of the species and decreases with an increase of the mean number of ants per resource unit.
• 3 Ingestion rates were higher in ants foraging singly at the resource than in ants foraging in the normal way in a group.
• 4 It is suggested that the ingestion rate per ant is reduced below a maximum level by the number of ants present per resource unit because a certain number of ants is needed to defend the resource against alien ants. Small species need more individuals for this purpose than large species, and consequently suffer a larger reduction of their ingestion rate.

     

Optimal foraging: food patch depletion by ruddy ducks

Summary I studied the foraging behavior of ruddy ducks (Oxyura jamaicensis) feeding on patchily distributed prey in a large (5-m long, 2-m wide, and up to 2-m deep) aquarium. The substrate consisted of a 4x4 array of wooden trays (1.0-m long, 0.5-m wide, and 0.1-m deep) which contained 6 cm of sand. Any tray could be removed from the aquarium and loaded with a known number of prey. One bird foraged in the aquarium at a time; thus, by removing a food tray after a trial ended and counting the remaining prey, I calculated the number of prey consumed by the bird. I designed several experiments to determine if ruddy ducks abandoned a food patch in a manner consistent with the predictions of a simple, deterministic, patch depletion model. This model is based on the premise that a predator should maximize its rate of net energy intake while foraging. To accomplish this, a predator should only remain in a food patch as long as its rate of energy intake from that patch exceeds the average rate of intake from the environment. In the majority of comparisons, the number of food items consumed by the ruddy ducks in these experiments was consistent with the predictions of the foraging model. When the birds did not forage as predicted by the model, they stayed in the patch longer and consumed more prey than predicted by the model. An examination of the relation between rate of net energy intake and time spent foraging in the food patch indicated that by staying in a patch longer than predicted, the ruddy ducks experienced only a small deviation from maximum rate of net energy intake. These results provided quantitative support for the prediction that ruddy ducks maximize their rate of net energy intake while foraging.     

Scales and costs of habitat selection in heterogeneous landscapes

Summary Two scales of habitat selection are likely to influence patterns of animal density in heterogeneous landscapes. At one scale, habitat selection is determined by the differential use of foraging locations within a home range. At a larger scale, habitat selection is determined by dispersal and the ability to relocate the home range. The limits of both scales must be known for accurate assessments of habitat selection and its role in effecting spatial patterns in abundance. Isodars, which specify the relationships between population density in two habitats such that the expected reproductive success of an individual is the same in both, allow us to distinguish the two scales of habitat selection because each scale has different costs. In a two-habitat environment, the cost of rejecting one of the habitats within a home range can be expressed as a devaluation of the other, because, for example, fine-grained foragers must travel through both. At the dispersal scale, the cost of accepting a new home range in a different habitat has the opposite effect of inflating the value of the original habitat to compensate for lost evolutionary potential associated with relocating the home range. These costs produce isodars at the foraging scale with a lower intercept and slope than those at the dispersal scale.Empirical data on deer mice occupying prairie and badland habitats in southern Alberta confirm the ability of isodar analysis to differentiate between foraging and dispersal scales. The data suggest a foraging range of approximately 60 m, and an effective dispersal distance near 140 m. The relatively short dispersal distance implies that recent theories may have over-emphasized the role of habitat selection on local population dynamics. But the exchange of individuals between habitats sharing irregular borders may be substantial. Dispersal distance may thus give a false impression of the inability of habitat selection to help regulate population density.     

Adaptive foraging by predators as a cause of predator-prey cycles

Summary When foraging has costs, it is generally adaptive for foragers to adjust their foraging effort in response to changes in the population density of their food. If effort decreases in response to increased food density, this can result in a type-2 functional response; intake rate increases in a negatively accelerated manner as prey density increases. Unlike other mechanisms for type-2 responses, adaptive foraging usually involves a timelag, because foraging behaviours do not often change instantaneously with changes in food density or risks. This paper investigates predator-prey models in which there are explicit dynamics for the rate of adaptive change. Models appropriate to both behavioural and evolutionary change are considered. Both types of change can produce cycles under similar circumstances, but under some evolutionary models there is not sufficient genetic variability for evolutionary change to produce cycles. If there is sufficient variability, the remaining conditions required for cycles are surprisingly insensitive to the nature of the adaptive process. A predator population that approaches the optimum foraging strategy very slowly usually produces cycles under similar conditions as does a very rapidly adapting population.     

The use of optimal foraging theory in the understanding of fishing strategies: A case from Sepetiba Bay (Rio de Janeiro State,Brazil)

The community of Gamboa is located on Itacuruçá Island, Sepetiba Bay (State of Rio de Janeiro, Brazil) and includes 26 families, mostly of artisanal fishermen who use paddled or motor canoes, and encircling nets for fishing. In this study, predictions from optimal foraging theory (patch model), in particular of patch residence time, are compared to the observed behavior of fishermen on fishing trips. Fishermen's strategies differ depending on their intended prey. They spend more time in patches and use fewer patches for shrimp than for fish. Gamboa's fishermen tend to leave a patch later than predicted by the model. The difficulty in evaluating stock availability, the comparatively few patches available for shrimp, and the competitive aspects of fishing contribute to the explanation of this behavior.     

Patterns of nest provisioning and parental investment in the solitary digger wasp Ammophila sabulosa

Abstract.

• 1 In nest-building wasps there is a continuum from species that provision each offspring cell with one large prey item (single provisioning) to species that provision each cell with many small items (multiple provisioning). The significance of number of prey per cell was examined during two seasons in Ammophila sabulosa (L.), a species that provisions between one and five caterpillars per cell.
• 2 There was no difference between the total weight of prey in singly-provisioned and multiply-provisioned cells: individual caterpillars placed in multiply-provisioned cells were smaller.
• 3 Small caterpillars were captured and transported to the nest faster than large ones.
• 4 If the first prey provisioned was large, the cell was usually then permanently closed. If the first prey was small, additional prey were captured, and females appeared to become more selective: additional prey were rarely large.
• 5 It is suggested that the flexible provisioning strategy used by A.sabulosa allows a wide range of prey sizes to be utilized without affecting maternal control of offspring size and sex. This could be important when available prey sizes vary temporally and spatially.
• 6 There was a male-biased first generation investment ratio. Female-producing cells were provisioned with a higher total prey weight than male-producing cells, and conversion of prey weight to adult weight was more efficient for females.
• 7 Total prey weight provisioned is probably a good indirect measure of parental time investment in A.sabulosa. Other measures, particularly number of prey provisioned, will be less accurate.

     

Importance of communal foraging grounds outside the reed marsh for breeding great reed warblers

Foraging habitat selection of breeding great reed warblers was studied at a shore of Lake Biwa. The foraging grounds of parent warblers during the nesting period were not restricted to the breeding territory of the reed marsh, their nestling habitat. The paddy field outside the reed marsh was used communally by them throughout the breeding season. Females with early stage nestlings did not visit the paddy field whereas when nestlings were older than 3 days, more than half of their total food was collected there. Females with nests adjacent to the paddy field tended to exploit the paddy field more often than those with nests distant from it. Monogamously mated females tended to exploit the paddy field more often than polygynously mated females. Food collected in the paddy field was larger than that in the reed marsh and parent birds were prepared to travel longer distances to exploit the rich source of food in the paddy field. The importance of the communal foraging ground outside the reed marsh as a background of the polygynous mating system of this species is discussed.     

The parasitoidOoencyrtus nezarae (Hymenoptera: Encyrtidae) prefers hosts parasitized by conspecifics over unparasitized hosts

Summary Two laboratory experiments were conducted to examine the ovipositional preferences of the egg parasitoidOoencyrtus nezarae Ishii (Hymenoptera: Encyrtidae) for parasitized and unparasitizedMegacopta punctatissimum Montandon (Hemiptera: Plataspidae). Females that had never oviposited or that had not oviposite for 3 days preferred recently parasitized hosts more than unparasitized hosts. The preference for recently parasitized hosts appeared to be mediated by the punctures in already parasitized hosts made by the ovipositor of the first female. Survival of the parasitoid progeny was lower in recently parasitized hosts than in unparasitized hosts. However, handling time of parasitized hosts was extremely short relative to that of unparasitized hosts, because the superparasitizing female could use the punctures made by the previous females. It is concluded that the females preferred the parasitized hosts over unparasitized hosts because the benefit of saving time and energy for drilling was more than the cost of progeny survival.