The influence of nectar secretion rates on the responses of bumblebees (Bombus spp.) to previously visited flowers

Bumblebees can avoid recently depleted flowers by responding to repellent scent-marks deposited on flower corollas by previous visitors. It has previously been suggested that avoidance of visited flowers for a fixed period would be a poor strategy, since different plant species vary greatly in the rate at which they replenish floral rewards. In this study, we examined the duration of flower repellency after an initial bumblebee visit, using wild bumblebees (Bombus lapidarius, B. pascuorum and B. terrestris) foraging on four different plant species (Lotus corniculatus, Melilotus officinalis, Phacelia tanacetifolia and Symphytum officinale). We constructed a model to predict flower visitation following an initial visit, based on the nectar secretion pattern of the different plant species, the insect visitation rate per flower, and the search and handling times of bumblebees foraging on the plant species in question. The model predicts an optimal duration of flower avoidance which maximises the rate of reward acquisition for all bees. However, this optimum may be open to cheating. For two plant species, the evolutionary stable strategy (ESS) is a shorter duration of flower avoidance than the optimum. We found the duration of flower avoidance was markedly different among flower species and was inversely related to nectar secretion rates. The predicted ESSs for each plant species were close to those observed, suggesting that the key parameters influencing bumblebee behaviour are those included in the model. We discuss how bees may alter the duration of their response to repellent scents, and other factors that affect flower re-visitation.     

The importance of experience in the interpretation of conspecific chemical signals

Foraging bumblebees scent mark flowers with hydrocarbon secretions. Several studies have found these scent marks act as a repellent to bee foragers. This was thought to minimize the risk of visiting recently depleted flowers. Some studies, however, have found a reverse, attractive effect of scent marks left on flowers. Do bees mark flowers with different scents, or could the same scent be interpreted differently depending on the bees’ previous experience with reward levels in flowers? We use a simple experimental design to investigate if the scent marks can become attractive when bees forage on artificial flowers that remain rewarding upon the bees’ return after having depleted them. We contrast this with bees trained in the more natural scenario where revisits to recently emptied flowers are unrewarding. The bees’ association between scent mark and reward value was tested with flowers scent marked from the same source. We find that the bees’ experience with the level of reward determines how the scent mark is interpreted: the same scent can act as both an attractant and a repellent. How experience and learning influence the interpretation of the meaning of chemical signals deposited by animals for communication has rarely been investigated.     

Patch departure rules in Bumblebees: evidence of a decremental motivational mechanism

The patch living rules of a pollinator, the bumblebee Bombus terrestris L., are studied here in the framework of motivational models widely used for parasitoids: The rewarding events found during the foraging process are supposed to increase or decrease suddenly the tendency of the insect to stay in the current patch and therefore to adjust the patch residence time to the patch profitability. The foraging behaviour of these pollinators was observed in two environment types to determine their patch-leaving decisions. The rich environment was composed of male-fertile flowers, offering pollen and nectar, and the poor one of male-sterile flowers, offering little nectar and no pollen. The experimental design consisted of a patch system in which inflorescences were evenly arranged in two rows (1 m distance). Residence times of foragers inside inflorescences and rows were analysed by a Cox proportional hazards model, taking into account recent and past experience acquired during the foraging bout. Most of the results showed a decremental motivational mechanism, that is, a reduction in the residence time on the inflorescence or in the row related to exploitation of flowers within inflorescences and inflorescences within rows These results indicate that bumblebees tend to leave the patch using departure rules similar to those found in parasitoids. The results also provide information on the memory, learning and evaluating capabilities of bumblebees especially when rich and poor environments were compared. The patch-leaving mechanism suggested by this study is consistent with the central place foraging theory.     

The role of internal and external information in foraging decisions of Melipona workers (Hymenoptera: Meliponinae)

Social insect foragers have to make foraging decisions based on information that may come from two different sources: information learned and memorised through their own experience (“internal” information) and information communicated by nest mates or directly obtained from their environment (“external” information). The role of these sources of information in decision-making by foragers was studied observationally and experimentally in stingless bees of the genus Melipona. Once a Melipona forager had started its food-collecting career, its decisions to initiate, continue or stop its daily collecting activity were mainly based upon previous experience (activity on previous days, the time at which foraging was initiated the day(s) before, and, during the day, the success of the last foraging flights) and mediated through direct interaction with the food source (load size harvested and time to collect a load). External information provided by returning foragers advanced the start of foraging of experienced bees. Most inexperienced bees initiated their foraging day after successful foragers had returned to the hive. The start of foraging by other inexperienced bees was stimulated by high waste-removal activity of nest mates. By experimentally controlling the entries of foragers (hence external information input) it was shown that very low levels of external information input had large effect on the departure of experienced foragers. After the return of a single successful forager, or five foragers together, the rate of forager exits increased dramatically for 15 min. Only the first and second entry events had large effect; later entries influenced forager exit patterns only slightly. The results show that Melipona foragers make decisions based upon their own experience and that communication stimulates these foragers if it concerns the previously visited source. We discuss the organisation of individual foraging in Melipona and Apis mellifera and are led to the conclusion that these species behave very similarly and that an information-integration model (derived from Fig. 1) could be a starting point for future research on social insect foraging. Received: 16 April 1997 / Accepted after revision: 30 August 1997     

Lévy flight patterns are predicted to be an emergent property of a bumblebees’ foraging strategy

Bumblebees forage uninterrupted for long periods of time because they are not distracted by sex or territorial defense and have few predators. This has led to a long running debate about whether bumblebees forage optimally. This debate has been enriched by the possibility that bumblebees foraging within clover patches have flight patterns that can be approximated by Lévy flights. Such flight patterns optimise the success of random searches. Bumblebees foraging within a flower patch tend to approach the nearest flower but then often depart without landing or probing it if it has been visited previously; unvisited flowers are not rejected in this manner. Here, this foraging behaviour has been replicated in numerical simulations. Lévy flight patterns are found to be an inconsequential emergent property of a bumblebees’ foraging behaviour. Lévy flights are predicted to emerge when bees reject at least 99% of previously visited flowers. A foraging bumblebee can certainly empty a clover flower head of nectar in one visit, but lower rates of rejection are observed for many other flowers. These findings suggest that Lévy flight patterns in foraging bumblebees are rare and specific to a few flower species and that if they exist, then they are not part of an innate, evolved optimal searching strategy.     

The role of a scent beacon in the communication of food location by the stingless bee, Melipona panamica

Melipona panamica foragers can deposit a scent beacon that influences the orientation of foragers near a food source. In misdirection experiments, newcomers (bees from the same colony as trained foragers) consistently preferred the feeder at which trained foragers had fed (training feeder) over an identical feeder at which bees had never fed (control feeder) even when the training feeder was placed at a site where experienced foragers had never foraged. Through similar misdirection experiments, the effective radius of the scent beacon was determined to be greater than 6 and less than 12 m. Foragers may deposit this beacon during a sequence of departure behaviors performed at the feeder. Prior to leaving the feeder with a load of sugar solution, bees tended to perform the following sequence of behaviors: (1) spinning, (2) grooming, (3) abdomen dragging, (4) excreting anal droplets, and (5) producing sounds, although not all behaviors were performed prior to each departure or at all sucrose concentrations (0.5–2.5 m). As sucrose concentration increased, the number of newcomers significantly increased, and the number of experienced foragers producing sounds and spinning on the feeder increased. The exact source of the scent beacon remains a mystery. However, three important sources have been excluded. When choosing between identical paired feeders, foragers were not attracted to the feeders with (1) anal droplets, (2) extracts of sucrose solution at which foragers had fed, or (3) mandibular gland extracts. They were indifferent to the first two preparations and exhibited only typical alarm behavior towards the mandibular gland (MG) extract: they oriented towards the feeder with MG extract but consistently landed on the feeder with no MG extract. Other authors have suggested that Melipona foragers deposit anal droplets to attract recruits, however the frequency of anal droplet production and the mass of anal droplets produced by M. panamica foragers are negatively correlated with sucrose concentration. Thus the scent beacon is evidently not deposited with anal droplets, infused into the feeder solution, or produced by mandibular glands. Received: 2 September 1997 / Accepted after revision: 30 January 1998     

Regulation of reproduction by dominant workers in bumblebee (Bombus terrestris) queenright colonies

The mechanisms of regulating worker reproduction in bumblebees were studied by direct behavioral observations and by measuring ovarian development and juvenile hormone (JH) biosynthesis rates in workers under different social conditions. Workers in the last stage of Bombus terrestris colony development (the competition phase) had the lowest ovarian development and JH biosynthesis rates. Callows introduced into colonies immediately after queen removal (dequeened colonies) demonstrated a significant increase in ovarian development before, but not during, the competition phase. These findings differ from the higher ovarian development in colonies during the competition phase predicted by the prevailing hypothesis that worker reproduction starts in response to a decrease in queen inhibition. Reproduction of callows housed with dominant workers in small queenless groups was inhibited as in queenright colonies. This suggests that the reduced ovarian development and JH biosynthesis rates observed in dequeened and normally developing colonies during the competition phase also reflect inhibition by dominant workers. Thus, two distinct stages of inhibition of reproduction seem to exist: (1) before the competition phase, when the queen slows down worker ovarian development and prevents oviposition; (2) during the competition phase, when dominant workers inhibit ovarian development of other workers. Between these stages there seems to be a temporal “window” of enhanced worker reproductive development. The queen's typical switch to haploid egg production was not associated with changes in worker ovarian development or JH biosynthesis rates. These findings suggest that regulation of worker reproduction in B. terrestris is not determined by simple changes in the queen's inhibition capacity or by the sex of offspring and that the worker's role is more important than previously believed. Received: 18 March 1998 / Accepted after revision: 18 July 1998     

The use of waggle dance information by honey bees throughout their foraging careers

We studied the extent to which worker honey bees acquire information from waggle dances throughout their careers as foragers. Small groups of foragers were monitored from time of orientation flights to time of death and all in-hive behaviors relating to foraging were recorded. In the context of a novice forager finding her first food source, 60% of the bees relied, at least in part, on acquiring information from waggle dances (being recruited) rather than searching independently (scouting). In the context of an experienced forager whose foraging has been interrupted, 37% of the time the bees resumed foraging by following waggle dances (being reactivated) rather than examining the food source on their own (inspecting). And in the context of an experienced forager engaged in foraging, 17% of the time the bees initiated a foraging trip by following a waggle dance. Such dance following was observed much more often after an unsuccessful than after a successful foraging trip. Successful foragers often followed dances just briefly, perhaps to confirm that the kind of flowers they had been visiting were still yielding forage. Overall, waggle dance following for food discovery accounted for 12–25% of all interactions with dancers (9% by novice foragers and 3–16% by experienced foragers) whereas dance following for reactivation and confirmation accounted for the other 75–88% (26% for reactivation and 49–62% for confirmation). We conclude that foragers make extensive use of the waggle dance not only to start work at new, unfamiliar food sources but also to resume work at old, familiar food sources.     

A complex social structure with fission–fusion properties can emerge from a simple foraging model

Precisely how ecological factors influence animal social structure is far from clear. We explore this question using an agent-based model inspired by the fission–fusion society of spider monkeys (Ateles spp). Our model introduces a realistic, complex foraging environment composed of many resource patches with size varying as an inverse power law frequency distribution with exponent β. Foragers do not interact among them and start from random initial locations. They have either a complete or a partial knowledge of the environment and maximize the ratio between the size of the next visited patch and the distance traveled to it, ignoring previously visited patches. At intermediate values of β, when large patches are neither too scarce nor too abundant, foragers form groups (coincide at the same patch) with a similar size frequency distribution as the spider monkey’s subgroups. Fission–fusion events create a network of associations that contains weak bonds among foragers that meet only rarely and strong bonds among those that repeat associations more frequently than would be expected by chance. The latter form subnetworks with the highest number of bonds and a high clustering coefficient at intermediate values of β. The weak bonds enable the whole social network to percolate. Some of our results are similar to those found in long-term field studies of spider monkeys and other fission–fusion species. We conclude that hypotheses about the ecological causes of fission–fusion and the origin of complex social structures should consider the heterogeneity and complexity of the environment in which social animals live.     

Floral scent variation in two <Emphasis Type="Italic">Antirrhinum majus</Emphasis> subspecies influences the choice of naïve bumblebees

Two wild subspecies of snapdragon, Antirrhinum majus, subspecies pseudomajus and striatum, differ in floral color and can be visually discriminated by insect visitors. The extent to which olfactory cues derived from floral scents contribute to discrimination between snapdragon subspecies is however unknown. We tested whether these two subspecies differ in floral scent and whether these olfactory differences are used by bumblebees (Bombus terrestris) to discriminate between them. We grew individuals of both subspecies, collected from a total of seven wild populations, under controlled conditions. We quantified the volatile organic compounds (VOCs) emitted by the flowers using gas-chromatography/mass-spectrometry/flame-ionization-detection. We studied antennal detection of VOCs by bumblebees, by means of electroantennogram study (EAG). We also performed behavioral experiments in a Y-maze to determine the innate response of bumblebees to the main floral VOCs emitted by our snapdragon subspecies. The floral scent of Antirrhinum majus pseudomajus contained three volatile benzenoids absent in the floral scent of Antirrhinum majus striatum. One of them, acetophenone, contributed over 69% of the absolute emissions of A. majus pseudomajus. These benzenoids elicited a significantly higher EAG response compared with other VOCs. In the Y-maze, bumblebees were significantly less attracted by acetophenone, suggesting an aversive effect of this VOC. Our findings indicate that bumblebees are able to discriminate between the two Antirrhinum majus subspecies. Differences in flower scent between these subspecies and olfactory bumblebee preferences are discussed in the light of biochemical constraints on VOCs synthesis and of the role of flower scent in the evolutionary ecology of A. majus.     

Colony energy requirements affect the foraging currency of bumble bees

Summary This study examines whether the foraging behavior of worker bumble bees (Bombus: Apidae) collecting nectar on inflorescences of seablush (Plectritis congesta: Valerianaceae) is affected by colony energetic requirements, which were experimentally manipulated either by adding sucrose solution to honey pots or by removing virtually all available nectar from the pots. The competing hypotheses tested were: (1) no change; energetic requirements do not affect behavior, since there is a single best way to collect food in a given environment; (2) energetic currency; the energetic currency maximized by foragers changes according to colony energetic condition, with nectar-depletion causing a shift from maximizing long-term productivity to maximizing immediate energetic gain, thereby de-emphasizing energetic costs; and (3) predation; foragers devalue risk of predation as risk of starvation increaes, with colony nectar-depletion causing foragers to be less predation riskaverse in order to increase immediate energetic gain. Relative to when their colony energy reserves were enhanced, foragers from nectar-depleted colonies selected smaller inflorescences, visited fewer flowers per inflorescence, probed flowers at a higher rate while on each inflorescence, and walked between inflorescences less often, thereby spending a greater proportion of their foraging trip in flight. These behaviors increased a bee's energetic costs while foraging, and should also have increased its immediate energetic gains, allowing rejection of the no change hypothesis. Predictions of the predation hypothesis were generally not supported, and our results best support the energetic currency hypothesis. Foraging currency of bumble bees therefore appears to be a function of colony energetic state. Offprint requests to: R.V. Cartar     

Colony nutritional status modulates worker responses to foraging recruitment pheromone in the bumblebee Bombus terrestris

Foraging activity in social insects should be regulated by colony nutritional status and food availability, such that both the emission of, and response to, recruitment signals depend on current conditions. Using fully automatic radio-frequency identification (RFID) technology to follow the foraging activity of tagged bumblebees (Bombus terrestris) during 16,000 foraging bouts, we tested whether the cue provided by stored food (the number of full honeypots) could modulate the response of workers to the recruitment pheromone signal. Artificial foraging pheromones were applied to colonies with varied levels of food reserves. The response to recruitment pheromones was stronger in colonies with low food, resulting in more workers becoming active and more foraging bouts being performed. In addition to previous reports showing that in colonies with low food successful foragers perform more excited runs during which they release recruitment pheromone and inactive workers are more prone to leave the nest following nectar influx, our results indicate that evolution has shaped a third pathway that modulates bumblebee foraging activity, thus preventing needless energy expenditure and exposure to risk when food stores are already high. This new feedback loop is intriguing since it involves context-dependent response to a signal. It highlights the integration of information from both forager-released pheromones (signal) and nutritional status (cue) that occurs within individual workers before making the decision to start foraging. Our results support the emerging view that responses to pheromones may be less hardwired than commonly acknowledged. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Choosing rewarding flowers; perceptual limitations and innate preferences influence decision making in bumblebees and honeybees

Flowers exhibit great intra-specific variation in the rewards they offer. At any one time, a significant proportion of flowers often contain little or no reward. Hence, foraging profitably for floral rewards is problematic and any ability to discriminate between flowers and avoid those that are less rewarding will confer great advantages. In this study, we examine discrimination by foraging bees among flowers of nasturtium, Tropaeolum majus. Bee visitors included carpenter bees, Xylocopa violacea, which were primary nectar robbers; honeybees, Apis mellifera, which either acted as secondary nectar robbers or gathered pollen legitimately and bumblebees, Bombus hortorum, which were the only bees able to gather nectar legitimately. Many flowers were damaged by phytophagous insects. Nectar volume was markedly lower in flowers with damaged petals (which were also likely to be older) and in flowers that had nectar-robbing holes. We test whether bees exhibit selectivity with regards to the individual flowers, which they approach and enter, and whether this selectivity enhances foraging efficiency. The flowers approached (within 2 cm) by A. mellifera and B. hortorum were non-random when compared to the floral population; both species selectively approached un-blemished flowers. They both approached more yellow flowers than would be expected by chance, presumably a reflection of innate colour preferences, for nectar standing crop did not vary according to flower colour. Bees were also more likely to accept (land on) un-blemished flowers. A. mellifera gathering nectar exhibited selectivity with regards to the presence of robbing holes, being more likely to land on robbed flowers (they are not able to feed on un-robbed flowers). That they frequently approached un-robbed flowers suggests that they are not able to detect robbing holes at long-range, so that foraging efficiency may be limited by visual acuity. Nevertheless, by using a combination of long-range and short-range selectivity, nectar-gathering A. mellifera and B. hortorum greatly increased the average reward from the flowers on which they landed (by 68% and 48%, respectively) compared to the average standing crop in the flower population. Overall, our results demonstrate that bees use obvious floral cues (colour and petal blemishes) at long-range, but can switch to using more subtle cues (robbing holes) at close range. They also make many mistakes and some cues used do not correlate with floral rewards.     

Chemical ecology involved in invasion success of the cuckoo bumblebee <Emphasis Type="Italic">Psithyrus vestalis</Emphasis> and in survival of workers of its host <Emphasis Type="Italic">Bombus terrestris</Emphasis>

In bumblebees all species of the subgenus Psithyrus are social parasites in the nests of their Bombus hosts. In the bumblebee B. terrestris we investigated how colony size influences survival rates of nest entering females of the social parasite Psithyrus vestalis. Furthermore, we studied whether the host worker’s dominance status and age are reflected in its individual scent and whether Psithyrus females use volatiles to selectively kill host workers. The survival rate of Psithyrus vestalis females drops from 100%, when entering colonies with five workers, to 0% for colonies containing 50 host workers. Older host workers, born before the nest invasion, were selectively killed when Psithyrus females entered the nest. In contrast, all workers born after the nest invasion survived. The host workers’ dominance status and age are reflected by their individual odours: newly emerged workers produced a significantly lower total amount of secretions than 4-day-old workers. In chemical analyses of female groups we identified saturated and unsaturated hydrocarbons, aldehydes, and unsaturated wax-type esters of fatty acids. In a discriminant function analysis different worker groups were mainly separated by their bouquets of hydrocarbons. Killed workers release significantly more scent and of a different chemical composition, than survivors. Survivors alter scent production and increase it beyond the level of the killed workers within 1 day of the invasion. The Psithyrus female clearly maintains reproductive dominance utilizing these differences in the odour bouquets as criteria for killing workers that compete for reproduction.     

Potential mechanisms for the communication of height and distance by a stingless bee, Melipona panamica

This study investigates the recruitment communication mechanisms of a stingless bee, Melipona panamica, whose foragers can evidently communicate the three-dimensional location of a good food source. To determine if the bees communicate location information inside or outside the nest, we conducted removal experiments by training marked foragers to one of two identical feeders and then separating these experienced foragers from potential recruits as they left the nest. The feeders were positioned to test the communication of each dimension. The results show that recruits do not simply follow experienced foragers to the food source. Height and distance are communicated within the nest, while direction is communicated outside the nest. We then examined the pulsed sounds produced by recruiting foragers. While unloading food, recruiting foragers produced several short pulses and one or more very long pulses. On average, the longest unloading pulse per performance was 31–50% longer (P ≤ 0.018) for bees foraging on the forest floor than for bees foraging at the top of the forest canopy (40 m high). While dancing, recruiting foragers produced sound pulses whose duration was positively correlated with the distance to the food source (P < 0.001). Dancing recruiters also produced several short sound pulses followed by one or more long pulses. The longest dance pulse per performance was 291 ± 194 ms for a feeder 25 m from the nest and 1858 ± 923 ms for a feeder 360 m away from the nest. The mechanism of directional communication remains a mystery. However, the direction removal experiment demonstrates that newcomers cannot use forager-deposited scent marks for long-distance orientation (>100 m from the nest). Received: 25 September 1997 / Accepted after revision: 31 May 1998     

Brood pheromone stimulates pollen foraging in honey bees (Apis mellifera)

Foraging and the mechanisms that regulate the quantity of food collected are important evolutionary and ecological attributes for all organisms. The decision to collect pollen by honey bee foragers depends on the number of larvae (brood), amount of stored pollen in the colony, as well as forager genotype and available resources in the environment. Here we describe how brood pheromone (whole hexane extracts of larvae) influenced honey bee pollen foraging and test the predictions of two foraging-regulation hypotheses: the indirect or brood-food mechanism and the direct mechanism of pollen-foraging regulation. Hexane extracts of larvae containing brood pheromone stimulated pollen foraging. Colonies were provided with extracts of 1000 larvae (brood pheromone), 1000 larvae (brood), or no brood or pheromone. Colonies with brood pheromone and brood had similar numbers of pollen foragers, while those colonies without brood or pheromone had significantly fewer pollen foragers. The number of pollen foragers increased more than 2.5-fold when colonies were provided with extracts of 2000 larvae as a supplement to the 1000 larvae they already had. Within 1 h of presenting colonies with brood pheromone, pollen foragers responded to the stimulus. The results from this study demonstrate some important aspects of pollen foraging in honey bee colonies: (1) pollen foragers appear to be directly affected by brood pheromone, (2) pollen foraging can be stimulated with brood pheromone in colonies provided with pollen but no larvae, and (3) pollen forager numbers increase with brood pheromone as a supplement to brood without increasing the number of larvae in the colony. These results support the direct-stimulus hypothesis for pollen foraging and do not support the indirect-inhibitor, brood-food hypothesis for pollen-foraging regulation. Received: 5 March 1998 / Accepted after revision: 29 August 1998     

Chemical signals in bumble bee foraging

Summary Foraging bumblebees (Bombus vosnesenskii) deposit a substance on rewarding flowers which assists in discrimination between rewarding and nonrewarding flowers in a controlled laboratory environment. Discrimination occurs while the bee is on a flower; workers probe rewarding flowers as well as empty ones that have rewarded in the recent past, but they do not probe flowers that have had no reward. Recognition is not the result of honey contamination left on the flower by the bee during feeding. The deposit is only slightly soluble in water or ethyl alcohol but is very soluble in pentane.     

Optimal diets: simultaneous search and handling of multiple-prey loads by salamander larvae

Summary A key assumption of conventional opitmal diet theory is that foragers cannot search for prey while already handling one prey item. Some foragers, however, can handle multiple-prey loads; i.e., they can search for, attack and handle further prey when already handling one or more prey. We examined diet selection by small-mouthed salamander larvae, Ambystoma texanum, that can search while handling up to two prey at a time. We gave A. texanum larvae a choice between two size classes of Daphnia pulex at two prey densities. Larval A. texanum diet choice did not fit the predictions of conventional optimal diet theory, but fit very well with the predictions of a multiple-prey model. At low prey density, A. texanum larvae were nonselective. At high prey density, larvae were non-selective when their mouths were empty, but showed a strong preference for larger, more valuable prey when larvae already had prey in their mouths. In 16 out of 18 instances, foragers either accepted or rejected small prey in keeping with a multiple-prey model's predictions.     

Chemical resistance of gorgonian corals against fungal infections

The frequency and impact of diseases affecting corals throughout the Caribbean have been increasing but little is known about the factors promoting the emergence and outbreak of disease. A disease caused by a fungal pathogen [Aspergillus sydowii (Thom et Church)] which affects Caribbean sea fan corals provided an opportunity to examine the efficacy of coral crude extracts in disease resistance. Minimum inhibitory concentration (MIC) assays showed that of the 20 common gorgonian species in the Florida Keys, extracts from 15 species had MICs < 15 mg ml⁻¹ against A. sydowii pathogenic to sea fans. Extracts from several species in two gorgonian genera (Pseudoplexaura and Pseudopterogorgia) were among the most active, with MICs < 10 mg ml⁻¹. Gorgonia ventalina L., one of two sea fan species known to be hosts to A. sydowii in the field, had an MIC < 10 mg ml⁻¹, suggesting that complete disease resistance requires more active extracts. For the antifungal compounds to be effective in situ, they must also occur in sufficiently high concentrations in living coral tissue. For example, Pseudopterogorgia americana (Gmelin) had comparatively potent extracts but did not have sufficient concentrations in the tissue to be effective. Conversely, Plexaura homomalla Esper extracts were less potent but occurred in high enough concentrations in the tissue to be effective against A. sydowii. When potency and extract concentration are considered together (i.e. potency × concentration), several other gorgonian corals emerge as likely hosts to A. sydowii. Crude extracts from the most active gorgonian species were also effective against two geographic variants of A. sydowii pathogenic to sea fans, a non-pathogenic terrestrial strain of A. sydowii, and three strains of A. flavus Link known to be human, plant, and insect pathogens (MIC range, 7.5 to > 15 mg ml⁻¹). Although the potency in these assays did not attain a clinically significant level, the potency is comparable to a known antifungal agent, hygromycin B, which had an MIC ≤7.5 mg ml⁻¹ in our assays, highlighting the potential of these gorgonian corals for bioprospecting. Received: 29 May 1999 / Accepted: 22 May 2000     

An automated system for tracking and identifying individual nectar foragers at multiple feeders

Nectar-feeding animals have served as the subjects of many experimental studies and theoretical models of foraging. Their willingness to visit artificial feeders renders many species amenable to controlled experiments using mechanical “flowers” that replenish nectar automatically. However, the structural complexity of such feeders and the lack of a device for tracking the movements of multiple individuals have limited our ability to ask some specific questions related to natural foraging contexts, especially in competitive situations. To overcome such difficulties, we developed an experimental system for producing computer records of multiple foragers harvesting from simple artificial flowers with known rates of nectar secretion, using radio frequency identification (RFID) tags to identify individual animals. By using infrared detectors (light-emitting diodes and phototransistors) to activate the RFID readers momentarily when needed, our system prevents the RFID chips from heating up and disturbing the foraging behavior of focal animals. To demonstrate these advantages, we performed a preliminary experiment with a captive colony of bumble bees, Bombus impatiens. In the experiment, two bees were tagged with RFID chips (2.5 × 2.5 mm, manufactured by Hitachi-Maxell, Ltd., Tokyo, Japan) and allowed to forage on 16 artificial flowers arranged in a big flight cage. Using the resulting data set, we present details of how the bees increased their travel speed between flowers, while decreasing the average nectar crop per flower, as they gained experience. Our system provides a powerful tool to track the movement patterns, reward history, and long-term foraging performance of individual foragers at large spatial scales.