Modelling interactions between scavenger behaviour and farming practices: Impacts on scavenger population and ecosystem service efficiency

Long term conservation of ecosystem services requires a deep understanding of their basic processes and the development of tools to assess the effects of human practices on their efficiency. As an example of recycling service, we focus on the consumption of livestock carcasses by obligate scavengers, taking the case of vultures which face a dramatic decline worldwide. We investigate whether maintaining this recycling service through individual feeding stations called light feeding stations in contrast to aggregated resource on heavy feeding stations can meet the double objectives of vulture conservation and service efficiency. We built a spatially explicit multi-agent model to investigate the long term effects of livestock farmers practices on both the recycling service efficiency for farmers and vulture population carrying capacity including: (i) carcass disposal practices that drive the quantity and spatial distribution of resources for vultures and (ii) temporal distribution of livestock mortality due to economic choices that drive the dynamics of resources for vultures. In addition, we examine various scenarios related to vulture feeding behaviour - i.e. central place vs. random foraging and contest vs. scramble intraspecific competition - that may play a role in the vulture carrying capacity and recycling service efficiency. When accounting for vulture central place foraging, we found that favouring the use of light feeding stations instead of heavy feeding station does not affect vulture population carrying capacity and increases the number of farmer for which vulture service is optimal. The increase of light feeding station users poorly affects the number of farmers for which vulture service is optimal in contrast to results obtained with a vulture random foraging behaviour. Both of the recycling service efficiency and the vulture carrying capacity vary with the kind of intraspecific competition and with the seasonal distribution of livestock mortality. Livestock mortality distributions with a seasonal peak of mortality result in a mismatch between vulture food requirement and farmers needs for carcass removal. Finally we raise several points on the relevance of light feeding stations as a sustainable management for vulture conservation and for the recycling service and discuss the potential implications of seasonal scarcity of resources due to farming economic constraints.     

Effects of vulture declines on facultative scavengers and potential implications for mammalian disease transmission

Vultures (Accipitridae and Cathartidae) are the only known obligate scavengers. They feed on rotting carcasses and are the most threatened avian functional group in the world. Possible effects of vulture declines include longer persistence of carcasses and increasing abundance of and contact between facultative scavengers at these carcasses. These changes could increase rates of transmission of infectious diseases, with carcasses serving as hubs of infection. To evaluate these possibilities, we conducted a series of observations and experimental tests of the effects of vulture extirpation on decomposition rates of livestock carcasses and mammalian scavengers in Kenya. We examined whether the absence of vultures changed carcass decomposition time, number of mammalian scavengers visiting carcasses, time spent by mammals at carcasses, and potential for disease transmission at carcasses (measured by changes in intraspecific contact rates). In the absence of vultures, mean carcass decomposition rates nearly tripled. Furthermore, the mean number of mammals at carcasses increased 3-fold (from 1.5 to 4.4 individuals/carcass), and the average time spent by mammals at carcasses increased almost 3-fold (from 55 min to 143 min). There was a nearly 3-fold increase in the mean number of contacts between mammalian scavengers at carcasses without vultures. These results highlight the role of vultures in carcass decomposition and level of contact among mammalian scavengers. In combination, our findings lead us to hypothesize that changes in vulture abundance may affect patterns of disease transmission among mammalian carnivores.     

Modeling the impact of feeding stations on vulture scavenging service efficiency

Vultures, the only vertebrate obligate scavengers, are currently facing a dramatic worldwide decline with over half of vulture species now classified as threatened. To address this widespread decline, the use of feeding stations has been widely advocated in recovery programs. However, providing food that is more predictable in time and space than natural resources could disrupt the ecological scavenging service provided by vultures. In this study, we build a multi-agent system (MAS), which describes the social foraging behavior of Gyps vultures in order to investigate how resource predictability in space and resource management affect scavenging service efficiency. We study the possibility that vultures take into account feeding station location in addition of using local enhancement. Our results show that the efficiency of the scavenging service is heavily affected by the way resources are distributed between feeding stations and natural areas. Nevertheless, it appears possible to minimize a loss in service efficiency by increasing the number of feeding stations while keeping the total amount of resources available constant, thus reducing the predictability of the resources located on the stations. We illustrate our work in the case of European feeding stations that provide supplementary resources in areas where natural ones are scarce. Moreover, we discuss the implications worldwide, in particular in the case of India, where feeding stations for vultures are intended to lure individuals away from potentially harmful natural carrion.     

Turkey vultures foraging at experimental food patches: a test of information transfer at communal roosts

Summary Experimental food patches were used to assess the importance of food information transfer in communally roosting turkey vultures (Cathartes aura) in southern Ontario. Feeding trials failed to provide evidence of recruitment due to information transfer. Overall, fewer birds arrived at novel food patches on the days following discovery than was expected, had information transfer been operating. Earlier arrivals on second days were more likely due to local enhancement rather than information transfer since the size of groups arriving at food patches did not differ between the 2 days. These results indicate that turkey vulture roosts in Ontario do not operate as centers for food information transfer. Intraspecific competition, preferential selection of small-sized carrion, and a low degree of kin association make the proposed benefits of information transfer much less applicable to turkey vultures than has been found for other scavenging species.Offprint requests to: P.J. Weatherhead     

Suspected flunixin poisoning of a wild Eurasian Griffon Vulture from Spain

Exposure to residues of the nonsteroidal anti‐inflammatory drug (NSAID) diclofenac present in livestock carcasses has caused extensive declines in 3 Gyps vulture species across Asia. The carcass of a wild Eurasian Griffon Vulture (Gyps fulvus) was found in 2012 on an Andalucian (Spain) game hunting reserve and examined forensically. The bird had severe visceral gout, a finding consistent with Gyps vultures from Asia that have been poisoned by diclofenac. Liver and kidney samples from this Eurasian Griffon Vulture contained elevated flunixin (an NSAID) levels (median = 2.70 and 6.50 mg/kg, respectively). This is the first reported case of a wild vulture being exposed to and apparently killed by an NSAID outside Asia. It is also the first reported instance of mortality in the wild resulting from environmental exposure to an NSAID other than diclofenac. Caso de Sospecha de Envenenamiento por Flunixin de un Buitre Leonado en España     

Social dominance in migrant and resident turkey vultures at carcasses: evidence for a despotic distribution?

We placed carcasses in three different vegetation types in the heterogeneous savannas of central Venezuela to investigate the role of social dominance in habitat use by flocking migrant and resident turkey vultures (Cathartes aura meridionalis and C. a. ruficollis). Migrants foraged primarily in savanna habitats while residents foraged almost exclusively in gallery forest. In the gallery forest residents discovered carrion first significantly more often than migrants, despite there being equal densities of residents and migrants foraging over this habitat. Because residents fed in smaller groups than migrants at carcasses they had higher feeding rates. There was also a negative relationship between group sizes of residents and migrants. The feeding rate of residents declined in response to increased group size of migrants, but group size of residents had no effect. Migrant group size also had a greater effect on resident feeding rates than king vulture presence or absence. When the effect of migrant and resident group size on feeding rates in migrants was compared, the most significant factor was migrant group size. A second analysis showed that both resident group size and presence or absence of king vultures had a significant effect on feeding rates in migrants. Rates of agonistic encounters in migrant and resident turkey vultures increased weakly in relation to group size. However, there was an increase in residents' encounter rate with migrants in relation to increased migrant group size; there was no difference in resident encounter rates with other residents in relation to resident group size. Migrants dominated residents in almost all agonistic interactions over carcasses. We suggest that savanna habitats were less attractive to residents for foraging because they held larger groups of migrants.     

Priority areas for conservation of Old World vultures

The prosperity and well-being of human societies relies on healthy ecosystems and the services they provide. However, the biodiversity crisis is undermining ecosystems services and functions. Vultures are among the most imperiled taxonomic groups on Earth, yet they have a fundamental ecosystem function. These obligate scavengers rapidly consume large amounts of carrion and human waste, a service that may aid in both disease prevention and control of mammalian scavengers, including feral dogs, which in turn threaten humans. We combined information about the distribution of all 15 vulture species found in Europe, Asia, and Africa with their threats and used detailed expert knowledge on threat intensity to prioritize critical areas for conserving vultures in Africa and Eurasia. Threats we identified included poisoning, mortality due to collision with wind energy infrastructures, and other anthropogenic activities related to human land use and influence. Areas important for vulture conservation were concentrated in southern and eastern Africa, South Asia, and the Iberian Peninsula, and over 80% of these areas were unprotected. Some vulture species required larger areas for protection than others. Finally, countries that had the largest share of all identified important priority areas for vulture conservation were those with the largest expenditures related to rabies burden (e.g., India, China, and Myanmar). Vulture populations have declined markedly in most of these countries. Restoring healthy vulture populations through targeted actions in the priority areas we identified may help restore the ecosystem services vultures provide, including sanitation and potentially prevention of diseases, such as rabies, a heavy burden afflicting fragile societies. Our findings may guide stakeholders to prioritize actions where they are needed most in order to achieve international goals for biodiversity conservation and sustainable development.     

Paying for information: partial loads in central place foragers

Information about food sources can be crucial to the success of a foraging animal. We predict that this will influence foraging decisions by group-living foragers, which may sacrifice short-term foraging efficiency to collect information more frequently. This result emerges from a model of a central-place forager that can potentially receive information on newly available superior food sources at the central place. Such foragers are expected to return early from food sources, even with just partial loads, if information about the presence of sufficiently valuable food sources is likely to become available. Returning with an incomplete load implies that the forager is at that point not achieving the maximum possible food delivery rate. However, such partial loading can be more than compensated for by an earlier exploitation of a superior food source. Our model does not assume cooperative foraging and could thus be used to investigate this effect for any social central-place forager. We illustrate the approach using numerical calculations for honeybees and leafcutter ants, which do forage cooperatively. For these examples, however, our results indicate that reducing load confers minimal benefits in terms of receiving information. Moreover, the hypothesis that foragers reduce load to give information more quickly (rather than to receive it) fits empirical data from social insects better. Thus, we can conclude that in these two cases of social-insect foraging, efficient distribution of information by successful foragers may be more important than efficient collection of information by unsuccessful ones.     

Experimental tests of the information-center hypothesis with black vultures (Coragypsatratus) and turkey vultures (Cathartesaura)

In field tests of the information-center hypothesis (ICH) in south Texas with black vultures (Coragyps atratus) and turkey vultures (Cathartes aura), large carcasses were provided and kept under continuous observation. The use vultures made of these bait sites and their patterns of arrival were recorded to evaluate predictions derived from the ICH. Turkey vultures discovered most bait sites (30 of 31) first, but frequently were displaced from the food by later-arriving black vultures. This competitive exclusion by black vultures limited subsequent feeding opportunities for turkey vultures sufficiently that few (27%) returned on subsequent days to bait sites they had previously visited. I found no evidence that those turkey vultures that did return to bait sites acted as leaders for groups of naive birds and led them to bait sites – knowledgeable and naive turkey vultures did not arrive at bait sites together, and groups arriving at bait sites were not larger on subsequent days than on the first days carcasses were available. In contrast, a significantly larger percentage (47%) of knowledgeable black vultures returned to bait sites they had visited on previous days, and the first groups of black vultures arriving at bait sites on subsequent days were significantly larger than the equivalent groups on first days. Nine flocks of black vultures that arrived on subsequent days at bait sites before sunrise (which suggests the birds had commuted directly from a roost) contained knowledgeable birds, and two of these flocks contained both knowledgeable and naive individuals. Overall, 10 of 54 naive tagged black vultures (18.5%) arrived at bait sites under circumstances that suggested they had followed conspecifics to the food from a roost. However, most black vultures apparently found carcasses through independent search or by using local enhancement. Therefore, I conclude that while following from roosts to food sites is a strategy used by black vultures, at this study site it is one they use relatively infrequently. Received: 20 February 1997 / Accepted after revision: 28 June 1997     

On the possible contribution of mixed species flocks to species richness in neotropical avifaunas

Summary Mixed species foraging flocks are a dominant component of the infra-structure of avian communities in neotropical forests. In Amazonia, these flocks consist of pairs of 10–20 species, many of which are permanently associated with mixed flocks. At least half of these flocking species maintain territories that correspond exactly to the flock home range. Small individuals that participate as permanent members of the flocks must adopt the large home range of the larger nucleus species. Therefore, the densities of smaller species are dependent on the availability and density of flocks rather than the availability of food resources. Single pairs of 4 small flocking species with individual body masses of 8 g occupied exclusive territories of 8–12 ha. These were the same exact territories that were defended by at least 6 other flocking species with individual body masses of up to 37 g. Because of their attachment to flocks with large territories, small species are expected to under-utilize available food resources. The under-utilization of food resources is expected to allow smaller species to coexist with greater niche overlap resulting in increased species richness. This hypothesis was tested by quantifying foraging niche in terms of foraging height, foraging maneuver, and prey substrate; and using these values in addition to body mass and bill size (length, depth and width) to determine relative niche overlap between large versus small species pairs.Smaller species had greater foraging overlap than large flocking species and particularly the three smallest species of the genus Myrmotherula; longipennis, axillaris and menetriesii had very high overlap (average foraging niche overlap for the 3 species=0.83±0.12 compared with 0.12±0.19 for all flocking species), similar body sizes (body masses differing by no more then 8%) and similar bill morphologies (maximum ratio in length=1.08, width=1.07, and depth=1.06). These results are consistent with the hypothesis that small species participating in Amazonian mixed flocks can coexist with greater niche overlap because their density is flock dependent rather than resource dependent.     

MORPH—An individual-based model to predict the effect of environmental change on foraging animal populations

This paper describes an individual-based model, MORPH, that has been designed to predict the effect of environmental change on foraging animal populations. The key assumptions of MORPH are that individuals within populations behave in order to maximise their perceived fitness, but that perceived fitness may not always be positively related to the actual chances of survival and reproduction. MORPH has been parameterised for coastal birds on several European sites and predicted the effect of environmental change, caused by factors such as habitat loss, disturbance from humans and sea-level rise, on the survival and body condition of these species. However, MORPH contains a basic framework to describe animal physiology and foraging behaviour, and the distribution and abundance of the resources required by these animals. Therefore, MORPH is not restricted to coastal birds, and is potentially applicable to a wider range of systems. To be applied to a forager system, MORPH requires parameters describing (i) the distribution of the food supply and how food quality and abundance changes through time; (ii) the rate at which foragers consume food given the abundance of food and competitors; (iii) the amount of food the forager must consume each day to survive; (iv) the distribution and seasonal changes in other factors which influence the foraging behaviour and survival of foragers. The purpose of this paper is to (i) describe MORPH, (ii) give examples of its application, (iii) describe the types of systems to which MORPH can be applied, and (iv) publish its source code and a user guide.     

To flee or not to flee: predator avoidance by cheetahs at kills

Mammalian carnivores are unusual because their primary competitors for food are often their primary predators. This relationship is most evident at persistent kills where dominant competitors are attracted to both the carcass (as a free meal) and to the killers (as potential prey). Cheetahs (Acinonyx jubatus) are frequent victims of kleptoparasitism, and cubs, and sometimes adults, are killed by lions (Panthera leo) or spotted hyenas (Crocuta crocuta). Between 1980 and 2002, we observed 639 kills made by cheetahs in Serengeti National Park, Tanzania. These kills were often visited by scavengers, including relatively innocuous species such as vultures and jackals and potentially dangerous species, like spotted hyenas and lions. We used cheetah behavior at kills to test a number of predictions about how cheetahs should minimize risk at kill sites given they face an increased risk of predation of themselves or their cubs. In particular, we examined the propensity of cheetahs of different age/sex classes to hide carcasses after making a kill, vigilance at kills, and the delay in leaving after finishing feeding with respect to ecological factors and scavenger presence. The behavior of single females at kills did not suggest that they were trying to avoid being killed, but the behavior of males, often found in groups, was in line with this hypothesis. In contrast, the behavior of mother cheetahs at kills appeared to be influenced greatly by the risk of cubs being killed. Our results suggest that cheetahs use several behavioral counterstrategies to avoid interspecific predation of self or cubs.     

A Comparative Analysis of Specialization and Extinction Risk in Temperate-Zone Bats

Abstract:  Identifying the factors that influence the extinction risk of animals is essential in conservation biology because they help identify endangered species and provide the basis for their preservation. We present a comparative study that uses data from the literature on the diet and morphological specialization of European and North American bat species to investigate the effect of specialization on extinction risk. We focused on bats because many species are endangered and their high ecological diversity makes them a good model system for our purpose. After correcting for phylogenetic inertia, we compared the influence of dietary niche breadth as a measure of food specialization and of wing morphology as a measure of foraging strategy, habitat adaptation, and migratory ability on the vulnerability of 35 insectivorous bat species. Our results do not support the hypothesis that a narrow dietary niche breadth is related to high extinction risk. Instead they suggest that habitat specialization, which is reflected in wing morphology, influences the extinction risk of bats. Our study shows that an initial risk assessment in temperate-zone bats could be based on data of wing morphology but not on dietary data obtained from fecal analyses.     

Dominance in feeding territories relates to foraging success and offspring growth in brown skuas <Emphasis Type="Italic">Catharacta antarctica lonnbergi</Emphasis>

Territoriality should lead to strict dominance, as territory holders typically control access to resources and exclude others from their use. In feeding territories, dominance should be reflected in foraging success and ultimately in reproduction differences; however, these successive links have rarely been made explicit. Therefore, we investigated a population of brown skuas Catharacta antarctica lonnbergi, in which only part of the breeding population occupied feeding territories within penguin colonies. We identified the dominance hierarchy and determined the foraging success of the participants in fights for access to penguin carcasses within the territories. Furthermore, we monitored offspring growth from parents with and without feeding territories. Our results indicated a clear dominance hierarchy with territorial birds in their own territory dominating over territorial breeders from other territories, non-territorial breeders and non-breeding birds. However, territory owners could not completely exclude others from access to food. Foraging success was positively related to dominance scores: The dominant territory owners received 63% of a carcass, whereas non-territorial pairs could get less than 10%. The link between foraging success and offspring development was less clear: Although male chicks of non-territorial parents suffered from lower growth rates and, thus, delayed fledging, there were no such differences in female chicks. Territoriality in skuas did not imply a complete occupation of food, but guaranteed optimal growth conditions for offspring. Non-territorial individuals were forced to search for alternative resources, and the restricted access to the preferred food resulted in inferior conditions for offspring development, making this foraging strategy less rewarding.     

A patch use model to separate effects of foraging costs on giving-up densities: an experiment with white-tailed deer (<Emphasis Type="Italic">Odocoileus virginianus</Emphasis>)

The giving-up density of food (GUD), the amount of food remaining in a patch when a forager ceases foraging there, can be used to compare the costs of foraging in different food patches. But, to draw inferences from GUDs, specific effects of foraging costs (predation risk, metabolic and missed opportunities costs) on GUDs have to be identified. As high predation risk, high metabolic costs and abundant food all should produce high GUDs, this does not allow us to infer directly the quality of a habitat. In order to separate the effect of each foraging cost, we developed an optimal foraging model based on food supplementation. We illustrate the use of our model in a study where we assessed the impact of a power line right-of-way in a white-tailed deer (Odocoileus virginianus) winter yard by determining whether the negative effects of cover loss outweigh the positive effects of browse regeneration.     

Bearded seal males perceive geographic variation in their trills

Geographic variation in animal vocalisations, ranging from micro- to macro-variations, has now been widely documented. These dialects can impair communication between separated groups or populations and thus may play a role in speciation processes. Although the existence of geographical variation has been already shown in some pinnipeds species, the extent to which individuals perceive it is poorly understood. Here, we studied this question in the bearded seal Erignatus barbatus, a species found across different arctic regions. As in other phocids, bearded seal males emit sophisticated acoustic displays while defending an aquatic territory during the breeding season. First, we used playbacks to demonstrate that the trill has a function in territorial defence. Second, we used synthetic trills from two distinct populations (Nunavut and Greenland), to show that bearded seal males perceived the geographic variation in the trills. Males from Greenland responded more strongly to trills from local males than those from a distant area (Nunavut). This study provides the first experimental evidence that phocid males can perceive geographic variation in their vocalisations. Acoustic analyses combined with playback experiments can help to identify distinct populations in bearded seals. This may then provide information on both strength and scale of breeding site fidelity in this and potentially other species.     

The effects of seston food quality on planktonic food web patterns

In planktonic food webs, the conversion rate of plant material to herbivore biomass is determined by a variety of factors such as seston biochemical/elemental composition, phytoplankton cell morphology, and colony architecture. Despite the overwhelming heterogeneity characterizing the plant–animal interface, plankton population models usually misrepresent the food quality constraints imposed on zooplankton growth. In this study, we reformulate the zooplankton grazing term to include seston food quality effects on zooplankton assimilation efficiency and examine its ramifications on system stability. Using different phytoplankton parameterizations with regards to growth strategies, light requirements, sinking rates, and food quality, we examined the dynamics induced in planktonic systems under varying zooplankton mortality/fish predation, light conditions, nutrient availability, and detritus food quality levels. In general, our analysis suggests that high food quality tends to stabilize the planktonic systems, whereas unforced oscillations (limit cycles) emerge with lower seston food quality. For a given phytoplankton specification and resource availability, the amplitude of the plankton oscillations is primarily modulated from zooplankton mortality and secondarily from the nutritional quality of the alternative food source (i.e., detritus). When the phytoplankton community is parameterized as a cyanobacterium-like species, conditions of high nutrient availability combined with high zooplankton mortality led to phytoplankton biomass accumulation, whereas a diatom-like parameterization resulted in relatively low phytoplankton to zooplankton biomass ratios highlighting the notion that high phytoplankton food quality allows the zooplankton community to sustain relatively high biomass and to suppress phytoplankton biomass to low levels. During nutrient and light enrichment conditions, both phytoplankton and detritus food quality determine the extent of the limit cycle region, whereas high algal food quality increases system resilience by shifting the oscillatory region towards lower light attenuation levels. Detritus food quality seems to regulate the amplitude of the dynamic oscillations following enrichment, when algal food quality is low. These results highlight the profitability of the alternative food sources for the grazer as an important predictor for the dynamic behavior of primary producer–grazer interactions in nature.     

Exploitation of carbohydrate food sources in <Emphasis Type="Italic">Polybia occidentalis</Emphasis>: social cues influence foraging decisions in swarm-founding wasps

An efficient exploitation of carbohydrate food sources would be beneficial for social wasp species that store nectar within their nest. In the swarm-founding polistine wasp Polybia occidentalis, we now demonstrate that the decisions of when and where to forage are influenced by information from conspecifics. Only when foragers had been trained to collect at artificial carbohydrate feeders did newcomers (food-source-naive individuals) continuously arrive at these feeders during 2 h of experiment. In control tests, in which no forager had been trained, not a single newcomer alighted at any of the offered carbohydrate food sources. This indicates that, during the foraging process, a nest-based input provided by successful foragers must have stimulated nestmates to search for food. Once activated, the newcomers’ choice on where to collect was strongly influenced by field-based social information. The mere visual presence of accumulated conspecifics (wasp dummies placed on one of the feeders) attracted newcomers to the food sources. Interestingly, however, visual enhancement was not the only decision-biasing factor at the feeding site. In an experimental series where searching wasps had to choose between the experimental feeder at which 3 foragers continuously collected and the control feeder with nine wasp dummies, only 40% of the wasps chose the visually enhanced feeder. This points to the existence of additional mechanisms of local enhancement. The possibility that, in social wasps, recruitment is involved in the exploitation of carbohydrate food sources is discussed.     

Ecological constraints on group size: an analysis of spider monkey and chimpanzee subgroups

The social organization of spider monkeys (Ateles geoffroyi) and chimpanzees (Pan troglodytes) appear remarkably similar. In this paper, field studies of these two species were used to (1) test a model of ecological constraints on animal group size which suggests that group size is a function of travel costs and (2) assess ecological and social factors underlying the social organization of these two species. Spider monkeys were studied over a 6-year period in Santa Rosa National Park, Costa Rica, and chimpanzees were studied for 6 years in Kibale National Park, Uganda. Adults of both species spent their time in small subgroups that frequently changed size and composition. Thus, unlike most primate species, spider monkeys and chimpanzees were not always in a spatially cohesive social group; each individual had the option of associating in subgroups of a different size or composition. Both species relied on ripe fruit from trees that could be depleted through their feeding activity. However, spider monkey food resources tended to occur at higher densities, were more common, less temporally variable, and did not reach the low levels experienced by chimpanzees. Analyses of the relationship between subgroup size and the density and distribution of their food resources suggested that travel costs limit subgroup size. However, these ecological factors did not influence all age/sex classes equally. For example, the number of adult males in a subgroup was a function of food density and travel costs. However, this was not the case for female chimpanzees, suggesting that the benefits of being in a subgroup for females did not exceed the costs, even when ecological conditions appeared to minimize subgroup foraging costs. Therefore, it seems likely that social strategies influenced the relationship between food resource variables and subgroup size.     

Dance precision of <Emphasis Type="Italic">Apis florea</Emphasis>—clues to the evolution of the honeybee dance language?

All honeybee species make use of the waggle dance to communicate the direction and distance to both food sources and potential new nest sites. When foraging, all species face an identical problem: conveying information about profitable floral patches. However, profound differences in nesting biology (some nest in cavities while others nest in the open, often on a branch or a cliff face) may mean that species have different requirements when dancing to advertise new nest sites. In cavity nesting species, nest sites are a precise location in the landscape: usually a small opening leading to a cavity in a hollow tree. Dances for cavities therefore need to be as precise as possible. In contrast, when the potential nest site comprises a tree or perhaps seven a patch of trees, precision is less necessary. Similarly, when a food patch is advertised, dances need not be very precise, as floral patches are often large, unless they are so far away that recruits need more precise information to be able to locate them. In this paper, we study the dance precision of the open-nesting red dwarf bee Apis florea. By comparing the precision of dances for food sources and nest sites, we show that A. florea workers dance with the same imprecision irrespective of context. This is in sharp contrast with the cavity-nesting Apis mellifera that increases the precision of its dance when advertising a potential new home. We suggest that our results are in accordance with the hypothesis that the honeybees’ dance communication initially evolved to convey information about new nest sites and was only later adapted for the context of foraging.