Seasonal changes in a honeyeater assemblage in Banksia woodland near Perth,Western Australia

Abstract

Honeyeaters were the most numerous birds in banksia woodland near Perth, Western Australia, throughout the year. Numbers were greatest in a Banksia littoralis swamp, but only during those few months when it contained large amounts of nectar. In the surrounding woodland, numbers were lower but fairly constant during the year. This reflects the smaller amounts of nectar produced throughout the year, by the overlapping flowering patterns of several Banksia and Adenanthos species.

Large and medium-sized honeyeaters (wattlebirds and New Holland Honeyeaters) and flocking silvereyes dominated the swamp when it flowered. In contrast, small honeyeaters (spinebills and Brown Honeyeaters), many of whom were highly territorial residents, comprised the majority of the woodland assemblage throughout the year. These observations support a model based upon aggressive defence of rich nectar sources by the larger honeyeater species, and more efficient exploitation of dispersed flowers by smaller honeyeaters.     

Movements of groups of tui (Prosthemadera novaeseelandiae) in winter and settlement of juvenile tui in summer

Abstract

An individually colour-banded population of tui (Meliphagidae) was studied and individuals were shown to move over sometimes large distances in search of nectar during most of the year. However, individuals were localised into small foraging areas when nectar was abundant during summer breeding.

Tui forage a regular route, taking their newly independent young with them through winter. These family groups sometimes consist of previous years' young and also other adults. These same group members nest near each other during summer and their offspring establish nesting areas nearby in following years.     

The feeding ecology of Rüppell's Parrot,Poicephalus rueppellii,in the Waterberg,Namibia

Rüppell's Parrot, Poicephalus rueppellii, inhabits a markedly seasonal environment in Namibia and feed on plant seeds that respond to that seasonality. Although Rüppell's Parrots were seen feeding at 37 plant species they showed clear preferences. The diet was dominated by seeds, but also included other fruit, flowers (especially nectar), leaves, arthropods and bark. The arthropods were the larvae of lepidoptera, coleoptera and hemiptera, and spiders. Feeding was observed on 19 plant species during a year period and the proportion of observations/species/month was determined. Terminalia prunioides, Albizia anthelmintica and Acacia er/o/oba represent preferred foods and provided most of the diet at particular times of the year. The fruiting periods of most plants were highly seasonal and the parrots changed diet frequently; however, T. prunioides holds its pods all year round and was used more than any other species. Monthly diet was also analysed by food type. Seeds formed the greatest part of the diet from December to August, but flowers and insects formed a major part of the diet prior to the onset of the rains. In particular, A. anthelmintica provided nectar and harboured insects when there was very little other food available. It may be critically important in the east of this parrot's distribution where the winter-fruiting Faidherbia albida is not found. For two species there Was some evidence that soft, slightly unripe drupes were preferred over riper, harder fruits. Rüppell's Parrot is omnivorous but mostly a seed predator. It survives in a highly seasonal environment by exploiting a wide range of food items.     

Diet of young Great Bustards Otis tarda in Spain: sexual and seasonal differences

Capsule Arthropods are the most important diet component of Great Bustards Otis tarda in the first months of life.

Aims To determine the diet composition of young Great Bustards in Spain.

Methods The diet was estimated by stomach content analysis (n?=?49).

Results Stomach contents' dry weight consisted of 33% arthropods, 30% green plant material and 23% seeds. Gastroliths were only found in summer and autumn. The diet composition changed significantly between seasons. In summer, diet consisted mainly of arthropods (50%), with green plant material being the main component in winter (56%). Volume of stomach contents and mean size of ingested arthropods were higher in males than in females. Diet composition did not differ between sexes. In summer, ground-dwelling and plant-visiting arthropods such as Mantidae, Tenebrionidae and caterpillars were the most abundant. In winter, weeds, legumes and cultivated seeds were more frequent than arthropods. Cereal plants were the least consumed in all seasons, although Barley and Wheat seeds played an important role during winter and autumn.

Conclusion The results highlight the importance of arthropods and weeds as a fundamental component of the diet of young Great Bustards. Because previous studies show that arthropods and weeds are usually more abundant in extensive farming, we recommend the implementation of agri-environmental measures in Great Bustard breeding areas.     

Why do nectar-foraging bees and wasps work upwards on inflorescences?

Summary Wasps (Dolichovespula and Vespula spp.) worked predominantly upwards when foraging for nectar on inflorescences of the protogynous Scrophularia aquatica, in which the standing crop of nectar sugar per flower showed no clear pattern of vertical distribution up an inflorescence. Bumblebees taking nectar (Bombus hortorum visiting legally, and certain individuals of B. terrestris which positioned themselves head-upwards while taking nectar through holes bitten in the corolla) worked predominantly upwards on the racemose inflorescences of Linaria vulgaris, although the standing crop of nectar sugar per open flower increased up the raceme. Individuals of B. terrestris which robbed Linaria flowers in a head-down position worked predominantly downwards on inflorescences. The upward or downward directionality of intra-inflorescence movements by foraging insects may depend in part on the position these adopt during their flower visits.     

Are nectar guide colour changes a reliable signal to pollinators that enhances reproductive success?

Background: Ageing and post-pollination changes in floral colour occur widely in flowering plants, but it remains an open question as to whether or not colour changes in nectar guides are associated with the quantity of floral rewards that ultimately influence pollinator visitations and reproductive success.

Aims: To examine whether nectar guide changes should be considered as a reliable signal to pollinators and to assess the effects of nectar guide changes on reproductive success.

Methods: We studied the process and adaptive value of colour changes in the nectar guides of Arnebia szechenyi whose flowers typically display conspicuous nectar guides at the onset of anthesis, after which they begin to fade, and disappear completely on the second day.

Results: Changes in nectar guide colour in A. szechenyi were intrinsic and age-dependent, although pollination somewhat accelerated the change. By the time that the nectar guides disappeared completely, floral rewards were reduced almost to zero. Artificial removal of nectar guides decreased both fruit set and pollen export. Flowers without nectar guides do not appear to increase the overall attractiveness of the plants.

Conclusions: Nectar guides and their changes represent reliable signals to pollinators and enhance both male and female reproductive success.     

Dietary Differences Among Sympatric Callitrichinae in Northern Bolivia: Callimico goeldii,Saguinus fuscicollis and S. labiatus

Studies of sympatric species can provide important data to define how dietary and habitat requirements differ among them. I collected dietary data during a first yearlong comparative study of wild groups of Callimico goeldii, Saguinus labiatus and S. fuscicollis. Dietary overlap was highest between Saguinus fuscicollis and Saguinus labiatus throughout the year, and lowest between Saguinus labiatus and Callimico goeldii. All three species had high dietary overlap in February and March when a few abundant fruit species dominated their diets. Although all three species rely heavily on many of the same fruits and arthropods, there are several important distinctions among their diets. Surprisingly, Callimico goeldii consume large quantities of fungus throughout the year: 29% of annual feeding records. Mycophagy is more frequent in the dry season when fruits are scarce. In contrast, Saguinus labiatus rarely eat fungus during the period of fruit scarcity, and instead rely on nectar, a resource never exploited by Callimico goeldii. Saguinus fuscicollis also rely on nectar during periods of low fruit availability and increase their intake of arthropods and exudates. During April, a period of fruit scarcity, exudates comprise >50% of the feeding records of Saguinus fuscicollis. The use of different food resources during fruit scarcity, and differences in the heights at which each species feeds and forages appear to define a distinct ecological niche for each of them and allow them to maintain long-term associations throughout the year. Furthermore, I hypothesize that the limited distribution of Callimico goeldii may result from their restriction to forests that have high disturbance rates, where microhabitats appropriate for fungal growth are abundant, but which also contain abundant fruit and insects.     

Butterfly Density and Behaviour in Uncut Hay Meadow Strips: Behavioural Ecological Consequences of an Agri-Environmental Scheme

Sparing zones from mowing has been proposed, and applied, to improve local conditions for survival and reproduction of insects in hay meadows. However, little is known about the efficiency of refuge zones and the consequences for local populations. We studied population densities of butterflies before and after mowing in the refuge zone of 15 meadows in 2009 and 2011. We also studied the behaviour of the meadow brown (Maniola jurtina) comparing nectar use, interactions and flights in the refuge zone before and after mowing. Densities of grassland butterflies in this zone doubled on average after mowing. The density of females of M. jurtina increased on average fourfold, while males showed a more modest increase. In line with the idea of increased scramble competition in the refuge zone after mowing, M. jurtina increased the time spent on nectar feeding, the preferred nectar source was visited more frequently, and females made more use of non-preferred nectar sources. Maniola jurtina did not interact more with conspecifics after mowing, but interactions lasted longer. Flight tracks did not change in linearity, but were faster and shorter after mowing. After mowing, only a part of the local grassland butterflies moved to the uncut refuge zone. The resulting concentration effect alters the time allocated to different activities, nectar use and movements. These aspects have been largely ignored for agri-environmental schemes and grassland management in nature reserves and raise questions about optimal quantities and quality of uncut refuge sites for efficient conservation of grassland arthropods in agricultural landscapes.     

Effects of Shade‐Tree Species and Crop Structure on the Winter Arthropod and Bird Communities in a Jamaican Shade Coffee Plantation1

I examined the effects of two farm management variables, shade‐tree species and crop structure, on the winter (dry season) arthropod and bird communities in a Jamaican shade coffee plantation. Birds and canopy arthropods were more abundant in areas of the plantation shaded by the tree Inga vera than by Pseudalbizia berteroana. The abundance of arthropods (potential pests) on the coffee crop, however, was unaffected by shade‐tree species. Canopy arthropods, particularly psyllids (Homoptera), were especially abundant on Inga in late winter, when it was producing new leaves and nectar‐rich flowers. Insectivorous and nectarivorous birds showed the strongest response to Inga; thus the concentration of birds in Inga may be a response to abundant food. Coffee‐tree arthropod abundance was much lower than in the shade trees and was affected little by farm management variables, although arthropods tended to be more abundant in dense (unpruned) than open (recently pruned) areas of the plantation. Perhaps in response, leaf‐gleaning insectivorous birds were more abundant in dense areas. These results underscore that although some shade coffee plantations may provide habitat for arthropod and bird communities, differences in farm management practices can significantly affect their abundances. Furthermore, this study provides evidence suggesting that bird communities in coffee respond to spatial variation in arthropod availability. I conclude that /. vera is a better shade tree than P. berteroana, but a choice in crop structures is less clear due to changing effects of prune management over time.     

Argentine ants displace floral arthropods in a biodiversity hotspot

Argentine ant (Linepithema humile (Mayr)) invasions are often associated with the displacement of ground‐dwelling arthropods. Argentine ant invasions can also exert other effects on the community through interactions with plants and their associated arthropods. For example, carbohydrate resources (e.g. floral or extrafloral nectar) may influence foraging behaviour and interactions among ants and other arthropods. In South Africa's Cape Floristic Region, Argentine ants and some native ant species are attracted to the floral nectar of Leucospermum conocarpodendron Rourke (Proteaceae), a native tree that also has extrafloral nectaries (EFNs). Despite having relatively low abundance in pitfall traps, Argentine ants visited inflorescences more frequently and in higher abundance than the most frequently observed native ants, Camponotus spp., though neither native nor Argentine ant floral foraging was influenced by the EFNs. Non‐metric multidimensional scaling revealed significant dissimilarity in arthropod communities on inflorescences with Argentine ants compared to inflorescences with native or no ants, with Coleoptera, Diptera, Hymenoptera, Arachnida, Orthoptera, and Blattaria all being underrepresented in inflorescences with Argentine ants compared to ant‐excluded inflorescences. Native honeybees (Apis mellifera capensis Eschscholtz) spent 75% less time foraging on inflorescences with Argentine ants than on inflorescences without ants. Neither Argentine ant nor native ant visits to inflorescences had a detectable effect on seed set of Le. conocarpodendron. However, a pollen supplementation experiment revealed that like many other proteas, Le. conocarpodendron is not pollen‐limited. Flower predation was negatively associated with increased ant visit frequency to the inflorescences, but did not differ among inflorescences visited by native and Argentine ants. Displacement of arthropods appears to be a consistent consequence of Argentine ant invasions. The displacement of floral arthropods by Argentine ants may have far‐reaching consequences for this biodiversity hotspot and other regions that are rich in insect‐pollinated plants.     

Arthropods in diet of Little Bustards Tetrax tetrax during the breeding season in western France

Capsule Seasonal, age- and sex-related variations in the diet were studied during March to October.

Aims To determine which arthropod orders contributed most during the laying–rearing period and those important in determining breeding success of the species in an intensive agricultural habitat.

Methods Faeces (n = 388, 345 g) and three gut samples were dissected to determine which arthropod orders contributed most to the diet, especially during the laying–rearing period.

Results Coleoptera were the most numerous prey followed by Orthoptera, Dermaptera and Hymenoptera. No difference was observed in the diet of adult males and females. Although adults ate arthropods throughout the breeding season, plant material contributed 97–99% of faecal contents by weight in each month. Chicks, however, only ate arthropods, notably Coleoptera and Orthoptera, at least until 2–3 weeks old.

Conclusion Increasing arthropod availability would be a useful management tool for maintaining endangered Little Bustard populations by potentially increasing chick survival.     

Honeybees and the nectar of Echium plantagineum L. in southeastern Australia

An account is given of the flower of Echium plantagineum in south-eastern Australia, including stages and timing of flowering, behaviour of raindrops in the flower and aspects of floral microclimate. The concentration of nectar solutes varied with time and site, with means varying from 2 to 62% (as g sucrose/100 g solution). There was a significant negative correlation between nectar solute concentration and ambient relative humidity: the drier the air, the more concentrated the nectar. Rates of nectar secretion per flower varied with the bagging method, with long-term bagging reducing net secretion rates, possibly because of re-absorption. Rates varied with time, day and site, with a temporal pattern of change suggesting a link between rates of photosynthesis and secretion. Maximum nectar secretion rates in short-term bagging experiments were ca. 300 μg sugar/flower/hr (equivalent to > 2 mglflower/24 hr). Secretion rate was correlated with flower density. As flower density increased, secretion rate per flower decreased; rate of sugar production per unit area increased relatively more slowly than flower density. E. plantagineum could produce > 500 mg sugar/m²/day. Honeybees foraged on E. plantagineum only at ambient air temperatures above ca. 17°C unless irradiance exceeded ca. 750 W m^-2. Foragers collected nectar or pollen alone, or both, with the type of visit significantly correlated with nectar solute concentration. Below 35% (as g sucrose/100 g solution) most bees took pollen only; above 40%, most took nectar. Mean standing crop of nectar was generally < 100 μg/flower when most bees were taking nectar, but could exceed 1000 μg/flower when bees were absent or foraging mainly for pollen. Honeybees did not always remove all nectar from flowers they probed. Reabsorption of residual nectar may augment the following day's secretion.     

Breeding system features and a novel method for locating floral nectar secretion in a South American nightshade (Jaltomata quipuscoae)

Abstract

The location of nectar secretion in flowers of Jaltomata has not been identified with certainty until now: removal of the corolla and androecium from one side of living flowers allowed us to see, in progress, nectar secretion by the ovarian nectary. We studied Jaltomata quipuscoae, a wild plant that grows in southern Peru and produces copious, red floral nectar. Unmanipulated flowers do not set fruit in a pollinator-free greenhouse, demonstrating lack of autogamy, but self-compatibility was demonstrated by manual self-pollinations leading to fruit-set. Anther dehiscence is staggered with the anthers of a flower dehiscing over hours on the same day. The corolla and nectar are UV-absorptive. Flowers last 4–10 days, are usually protogynous during the first day the corolla is open, and do not close for the night.     

Bumblebee death associated with Tilia x europaea L

For more than six decades, bumblebee death, which mainly occurs in August, has been assumed to be associated with intake of presumed toxic nectar from linden trees (Tilia spp.), a hypothesis which has been sustained by observations of a significant number of dead bumblebees under these trees during their flowering season. Several theories exist in current literature to account for these observations. The nectar has been assumed to contain compound(s) toxic to the bumblebees including the monosaccharide mannose, which cannot be metabolized by bumblebees. The presence of toxic compounds such as the alkaloid nicotine or pesticides of anthropogenic origin has also been indicated. However, none of the above suggested compounds have hitherto been properly characterized from the nectar. In the current paper we report on characterization of the composition of nectar of linden trees, under which a significant number of dead bumblebees were observed. The structure determinations were performed by selective 1D TOCSY NMR and extensive applications of 2D NMR spectroscopy. The nectar of the linden flowers was readily available in significant amounts during the entire period when dead bumblebees were observed under these trees. The nectar only contained non-toxic sugars such as α-glucopyranose, β-glucopyranose, sucrose, fructose and minor amounts of xylose. The nectar did not contain mannose, nor alkaloids or pesticides. Initial toxicity studies using brine shrimp lethality assay showed that the nectar did not exhibit any toxic effects even at concentrations higher than 1 mg/ml dry weight, providing disproving evidence against the assumption of the nectar's toxic character.     

Floral rewards in someScrophularia species (Scrophulariaceae) from the Iberian Peninsula and the Balearic Islands

The genusScrophularia in the Iberian Peninsula and Balearic Islands comprises two sections,Scrophularia andCanina G. Don. Analyses were carried out on flower production, flower duration and their sexual phases, pollen and nectar production together with observations on their pollinators. Nectar production is correlated with corolla size and pollen production with anther size. The taxa of sect.Scrophularia show greater nectar and pollen production than those of sect.Canina. Also, those of the first section produce more ovules per ovary than those of the second group, production being correlated with the ovary size.Some observations on floral and reproductive biology in some species ofScrophularia from the Iberian Peninsula and the Balearic Islands. I.     

Assessing risks and benefits of floral supplements in conservation biological control

The use of flowering field margins is often proposed as a method to support biological control in agro-ecosystems. In addition to beneficial insects, many herbivores depend on floral food as well. The indiscriminate use of flowering species in field margins can therefore lead to higher pest numbers. Based on results from field observations and laboratory experiments we assessed risks as well as benefits associated with the provision of nectar plants in field margins, using Brussels sprouts as a model system. Results show that Brussels sprouts bordered by nectar plants suitable for the cabbage white Pieris rapae L., suffered higher infestation levels by this herbivore. In contrast, nectar plants providing accessible nectar for the diamondback moth Plutella xylostella L., did not raise densities of P. xylostella larvae in the Brassica crop. Margins with Anethum graveolens L., selected on the basis of its suitability as nectar plant for parasitoids, significantly increased the number of adult Diadegma semiclausum Hellén in the crop. This didn’t translate into enhanced parasitism rates, as parasitism of P. xylostella by D. semiclausum exceeded 65 % in all treatments, irrespective of the plants in the field margin. Our findings emphasize the importance of taking a multitrophic approach when choosing flowering field margin plants for biocontrol or other ecosystem services.     

Feeding activity in captive New Zealand lesser short‐tailed bats (Mystacina tuberculata)

Abstract

Lesser short‐tailed bats (Mystacina tuberculata) have been reported as commonly feeding on the ground, but few direct observations of the diet and foraging behaviour of these rare and secretive bats have been published. Here, we describe the feeding behaviour of six captive M. tuberculata at Wellington Zoological Gardens, in order to experimentally clarify and validate some of the feeding behaviours previously reported from anecdotal observations. In particular, we focused on food type choice and dependence on nectar. The bats emerged every night 80 ± 30 (mean ± SE) min after sunset, irrespective of the weather, and spent most of their time foraging in leaf litter on the ground (when available) and on the branches of trees. Larvae of meal worms (Tenebrio molitor) and cerambycid beetles (Prionoplus reticularis) were favoured in the diet, followed in diminishing order by adult meal worms, tree weta (Hemideina crassidens) and crickets (Teleogryllus commodus), some nectars, sugar solutions, and finally water. Nectar from Eucalyptus sp. was preferred, followed by nectar of pohutukawa (Metrosideros excelsa), wood rose (Dactylanthus taylorii), and kakabeak (Clianthus puniceus). Nectar of Acacia sp., Hebe sp., and synthetic Dactylanthus nectar were not taken. The bats visited 50 and 25% sugar solutions significantly more frequently (91% of visits) than 12.5 and 0% sugar solutions. Various meats (carrion) and fruit were not eaten. Faecal content fairly represented the invertebrates consumed the previous night, but the volumes of invertebrates consumed could not be reliably estimated from faecal analysis.     

Geographic variation in a facultative mutualism: consequences for local arthropod composition and diversity

Geographic variation in the outcome of interspecific interactions may influence not only the evolutionary trajectories of species but also the structure of local communities. We investigated this community consequence of geographic variation for a facultative mutualism between ants and wild cotton (Gossypium thurberi). Ants consume wild cotton extrafloral nectar and can protect plants from herbivores. We chose three sites that differed in interaction outcome, including a mutualism (ants provided the greatest benefits to plant fitness and responded to manipulations of extrafloral nectar), a potential commensalism (ants increased plant fitness but were unresponsive to extrafloral nectar), and a neutral interaction (ants neither affected plant fitness nor responded to extrafloral nectar). At all sites, we manipulated ants and extrafloral nectar in a factorial design and monitored the abundance, diversity, and composition of other arthropods occurring on wild cotton plants. We predicted that the effects of ants and extrafloral nectar on arthropods would be largest in the location with the mutualism and weakest where the interaction was neutral. A non-metric multidimensional scaling analysis revealed that the presence of ants altered arthropod composition, but only at the two sites in which ants increased plant fitness. At the site with the mutualism, ants also suppressed detritivore/scavenger abundance and increased aphids. The presence of extrafloral nectar increased arthropod abundance where mutual benefits were the strongest, whereas both arthropod abundance and morphospecies richness declined with extrafloral nectar availability at the site with the weakest ant–plant interaction. Some responses were geographically invariable: total arthropod richness and evenness declined by approximately 20% on plants with ants, and extrafloral nectar reduced carnivore abundance when ants were excluded from plants. These results demonstrate that a facultative ant–plant mutualism can alter the composition of arthropod assemblages on plants and that these community-level consequences vary across the landscape.     

Odor-based recognition of nectar in cursorial spiders

Carnivorous arthropods are known to rely on non‐prey foods, such as honeydew, pollen, and nectar. Consumption of plant‐based nutrients by spiders also appears to be widespread, especially in cursorial species. This is not surprising, as studies have shown that these spiders’ activity levels, survivorship, and reproduction are increased when their diet includes plant‐based nutrients, especially under conditions of prey scarcity. However, the sensory and behavioral means by which they recognize and locate non‐prey food is unknown. Here we show that immatures of a nectarivorous spider [Hibana futilis Banks (Araneae: Anyphaenidae)] can recognize and remember particular chemical stimuli associated with nectar. Following ingestion of minute amounts of sugar, these spiders exhibited counterturning and other local searching behaviors that increased their chances of finding more nectar. When placed on test arenas, spiders that were naïve with respect to nectar aroma located artificial nectaries composed of diluted honey significantly faster than unscented nectaries composed of 1 m sucrose solution. These results indicate that H. futilis is neurophysiologically and behaviorally adapted for recognizing olfactory stimuli. Interestingly, only spiders that ingested sugar and were engaged in local search responded to nectar aroma, suggesting that stimulation into local search is necessary to prime olfactory responses. We found that H. futilis could be conditioned to associate the presence of nectar with a novel aroma, in this case vanilla, and remember this aroma over the course of several hours. In arenas with vanilla‐scented nectaries, spiders that had previous experience feeding on vanilla‐scented sucrose droplets located the nectaries significantly faster than did vanilla‐naïve spiders. The capacity to remember specific aromas could enhance the spiders’ ability to find nectar, either when moving between different parts of the same plant or among different plant species. The results here indicate that nectarivorous spiders possess the sensory capabilities and programed behaviors necessary for efficient detection, recognition, and location of nectar sources.     

Competition for nectar resources does not affect bee foraging tactic constancy

1. Competition alters animal foraging, including promoting the use of alternative resources. It may also impact how animals feed when they are able to handle the same food with more than one tactic. Competition likely impacts both consumers and their resources through its effects on food handling, but this topic has received little attention. 2. Bees often use two tactics for extracting nectar from flowers: they can visit at the flower opening, or rob nectar from holes at the base of flowers. Exploitative competition for nectar is thought to promote nectar robbing. If so, higher competition among floral visitors should reduce constancy to a single foraging tactic as foragers will seek food using all possible tactics. To test this prediction, field observations and two experiments involving bumble bees visiting three montane Colorado plant species (Mertensia ciliata, Linaria vulgaris, Corydalis caseana) were used under various levels of inter- and intra-specific competition for nectar. 3. In general, individual bumble bees remained constant to a single foraging tactic, independent of competition levels. However, bees that visited M. ciliata in field observations decreased their constancy and increased nectar robbing rates as visitation rates by co-visitors increased. 4. While tactic constancy was high overall regardless of competition intensity, this study highlights some intriguing instances in which competition and tactic constancy may be linked. Further studies investigating the cognitive underpinnings of tactic constancy should provide insight on the ways in which animals use alternative foraging tactics to exploit resources.