Ecological relevance of associative learning in fruit fly larvae

A few invertebrate models have been used for studying neurobiological and molecular aspects of associative learning. The ecological and evolutionary aspects of associative learning in these invertebrates are not yet well understood. To further this knowledge, I tested fruit fly larvae for their ability to learn to associate odors with preferred environmental conditions. The larvae learned to avoid odors associated with predation and to prefer odors associated with high-quality food, but failed to learn about odors associated with optimal temperature. It appears that the larvae possess a general ability to evaluate a suite of environmental parameters and associate preferred conditions with relevant stimuli. Received: 6 May 1998 / Accepted after revision: 11 October 1998     

Episodic death across species of desert shrubs

Extreme events shape population and community trajectories. We report episodic mortality across common species of thousands of long-lived perennials individually tagged and monitored for 20 years in the Colorado Desert of California following severe regional drought. Demographic records from 1984 to 2004 show 15 years of virtual stasis in populations of adult shrubs and cacti, punctuated by a 55-100% die-off of six of the seven most common perennial species. In this episode, adults that experienced reduced growth in a lesser drought during 1984-1989 failed to survive the drought of 2002. The significance of this event is potentially profound because population dynamics of long-lived plants can be far more strongly affected by deaths of adults, which in deserts potentially live for centuries, than by seedling births or deaths. Differential mortality and rates of recovery during and after extreme climatic events quite likely determine the species composition of plant and associated animal communities for at least decades. The die-off recorded in this closely monitored community provides a unique window into the mechanics of this process of species decline and replacement.     

Transitioning from unstable to stable colony growth in the desert leafcutter ant Acromyrmex versicolor

Like organisms, cohesive social groups such as insect colonies grow from a few individuals to large and complex integrated systems. Growth is driven by the interplay between intrinsic growth rates and environmental factors, particularly nutritional input. Ecologically inspired population growth models assume that this relationship remains constant until maturity, but more recent models suggest that it should be less stable at small colony sizes. To test this empirically, we monitored worker population growth and fungal development in the desert leafcutter ant, Acromyrmex versicolor, over the first 6 months of colony development. As a multitrophic, symbiotic system, leafcutter colonies must balance efforts to manage both fungus production and the growth of the ants consuming it. Both ants and fungus populations grew exponentially, but the shape of this relationship transitioned at a size threshold of 89?±?9 workers. Above this size, colony mortality plummeted and colonies shifted from hypometric to hypermetric growth, with a distinct stabilization of the relationship between the worker population and fungus. Our findings suggest that developing colonies undergo key changes in organizational structure and stability as they grow, with a resulting positive transition in efficiency and robustness.     

Ant gardens: interspecific recognition in parabiotic ant species

 In French Guiana, parabiotic societies (natural mixed colonies) are frequently found in ant gardens. Crematogaster limata parabiotica (Myrmicinae), often associated with Camponotus femoratus (Formicinae), was found for the first time in parabiosis with ponerine ants: Pachycondyla goeldii and Odontomachus mayi. A detailed study of the relationships between Cr. l. parabiotica and O. mayi showed that each species is aggressive towards allospecific or conspecific individuals belonging to another colony, but tolerates allospecific individuals from the multi-species society. Studies of cuticular substances of the four ant species were made using gas chromatography. The results showed that each species, living alone or in parabiosis, possesses a specific chemical profile. Thus, the ants are able to recognise nestmate and non-nestmate individuals of the associated species, even though their cuticular profiles are different. The hypothesis that the nestmate allospecific profile is learned is suggested to explain this pattern of recognition. Received: 5 June 1996 / Accepted after revision: 17 October 1996     

Inbreeding in a lek-mating ant species, Pogonomyrmex occidentalis

In this paper we have two goals. First, we examine the effects of sample size on the statistical power to detect a given amount of inbreeding in social insect populations. The statistical power to detect a given level of inbreeding is largely a function of the number of colonies sampled. We explore two sampling schemes, one in which a single individual per colony is sampled for different sample sizes and a second sampling scheme in which constant sampling effort is maintained (the product of the number of colonies and the number of workers per colony is constant). We find that adding additional workers to a sample from a colony makes it easier to detect inbreeding in samples from given number of colonies; however, adding more colonies rather than more workers per colony always gives greater power to detect inbreeding. Because even relatively large amounts of sib-mating generate relatively small inbreeding coefficients, detection of even substantial deviations from random mating will require very large samples. Second, we look at the amount of inbreeding in a large population of the western harvest ant, Pogonomyrmex occidentalis. We find deviations from Hardy-Weinberg equilibrium equivalent to approximately 27% sib-mating in our population ( f = 0.09). Review of past studies on the population structure of other Pogonomyrmex species suggests that inbreeding may be a regular feature of the mating system of these ants. Although P. occidentalisis a swarm-mating species, there are a number of features of its population biology which suggest that the effective population size may be small. These include topographical variation that potentially breaks the population into demes, variation in the reproductive output of colonies, and variation in the size of reproductives produced by colonies. Received: 6 May 1996 / Accepted after revision: 6 October 1996     

Plasticity of social organization in a forest ant species

We manipulated availability of food and nesting sites in one population of the forest ant Myrmica punctiventris. The manipulations produced significant changes in relatedness structure, reproductive allocation, and response to hierarchical selection. Food availability appeared to have a consistently stronger influence on these aspects of social organization than did availability of nesting sites. We interpret our experimental results in light of observed differences between populations, and discuss implications for kin selection dynamics. Received: 30 July 1998 / Accepted after revision: 31 October 1998     

Individually different foraging methods in the desert ant Cataglyphis bicolor (Hymenoptera,Formicidae)

Summary Observations and field experiments on the foraging behaviour of individual workers of Cataglyphis bicolor in a Southern Tunisian shrub desert are reported. The workers search singly for their food (mostly animal carcasses) and are singleprey loaders. The individuals differ to a great extent in their persistence to re-search the place of a find on a previous foraging excursion. The differences range continuously from thoroughly researching a place to just walking by. If, in an experiment, the same reward is offered farther from the nest, each ant persists more in re-searching the place than if food is offered close to the nest. In a further experiment, some individuals persisted less in searching near the former finding site if they had collected a fly than after collecting a piece of cheese. There is, however, evidence that individuals do not differ in their food preference. Persistent individuals, which re-search the place of a former find, are faster than non-persistent ones in retrieving food that is experimentally arranged in an aggregated manner. The experiment failed to demonstrate the (reverse) superiority of non-persistent individuals foraging on homogeneously distributed food. The observations of unmanipulated foraging excursions in the field suggest such an advantage for non-persistent foragers under natural conditions where food in general occurs widely dispersed. The colony as a whole retrieves more food within the same time from an experimental lay-out that is homogeneous than from an aggregated one. The behavioural differences between individuals could be caused by a training bias of the short-lived foragers, leading to a different assessment of the profitability of a searching method which implies returning to a formerly rewarding place. Thus, each worker uses the most promising behaviour according to its individual experience. Alternatively, the individually different searching methods could mainly contribute to the welfare of the colony as a whole rather than leading to a maximal short-term efficiency of each individual. In particular, the colony, disposing of only a few highly persistent foragers, could quickly exploit occasional short-lived, but unpredictible, clumps of food within its foraging range.     

Regional variation in queen and worker aggression in incipient colonies of the desert ant Messor pergandei

The fate of queen foundress associations in ants varies across taxa: in some, lethal fighting results in survival of a single queen, while in others, queens coexist long term. One hypothesis for this difference is that selection favors fighting when group sizes are small and tolerance when groups are large. In an experiment with the ant Messor pergandei, we formed small, medium, and large groups with newly mated queens from field populations that have different mean group sizes and differ in whether multiple queens occur in older established colonies. We found that whether queens are eliminated by fighting depends upon region of origin and not group size: regardless of co-foundress number, queens from sites with single-queen adult field colonies displayed agonistic behaviors and their colonies reduced to a single queen, while queens from sites with multiple-queen colonies did not fight and co-foundresses coexisted long term. Worker aggression towards and elimination of queens were also correlated with region of origin. Where fighting occurred, queens were as likely to be killed by workers as by other queens. An aggressive display was the most common form of agonistic interaction among queens, while fighting was relatively rare. We hypothesize that queen displays evolved in response to worker attacks because they increase the probability that workers will eliminate competitor queens. Our results suggest that the evolutionary interests of workers, as well as queens, could be important in determining the evolution and maintenance of queen elimination in foundress associations.     

Worker size-related task partitioning in the foraging strategy of a seed-harvesting ant species

Messor bouvieri is a seed-harvesting ant species in which workers forage in trails from the nest to a search area. A previous observation of seed transfer events between workers returning to the nest suggested potential task partitioning. In this study, we describe seed transportation and analyze the role of task partitioning in the foraging strategy of this species in terms of seed intake efficiency in relation to costs and benefits based on transport speed and task reliability. We assess the harvesting efficiency of task partitioning by comparing cooperative seed transport (CST) and individual seed transport (IST) events. Our results show task partitioning in the form of a sequence of transfer events among workers going from the search area to the nest. Importantly, and despite the weak worker polymorphism of this species, this sequence involved workers of different sizes, with seeds usually being passed along from smaller to larger workers. In addition, we show that small workers are better at finding seeds (spend less time finding a seed), and large workers are better at transporting them (were faster when walking back to the nest and lost fewer seeds). However, we failed to demonstrate that workers of different sizes are specialized in performing the task in which they excel. Overall, sequential CST in M. bouvieri results in a greater seed intake because seed search time decreases and task reliability increases, compared to IST. The determinants and adaptive benefits of CST are discussed.     

Behavioral and antennal electrophysiological responses of a predator ant to the pygidial gland secretions of two species of Neotropical dung roller beetles

The pygidial gland secretions of Neotropical dung roller beetles appear to function as defensive agents against predator ants. Behavioral and electrophysiological tests were used to evaluate the responses of the ant Camponotus sericeiventris to pygidial compounds secreted by the dung roller beetles Canthon cyanellus cyanellus and Canthon femoralis femoralis (Coleoptera: Scarabaeidae). Our behavioral investigation revealed a lower aggressive response of C. sericeiventris to the dung roller beetles compared to their response to the fly Anastrepha ludens. Additionally, C. sericeiventris workers attacked C. f. femoralis more often than C. c. cyanellus. The behavioral aggression of C. sericeiventris was greater toward male than female C. c. cyanellus. Electroantennography showed that the antennae of C. sericeiventris react to pygidial extracts from C. c. cyanellus and C. f. femoralis. Three compounds from male and female C. f. femoralis and C. c. cyanellus elicited coupled gas chromatography-electroantennogram (GC-EAD) activity in the antennae of C. sericeiventris. Geraniol and an unknown compound (A) were specific for C. f. femoralis, whereas guaiacol and another unknown compound (B) were specific to C. c. cyanellus; phenol was shared by both beetle species. This is the first report confirming that the pygidial gland secretions of dung roller beetles play a role in their defense against predators.     

宁夏荒漠草原恢复演替过程中物种多样性与生产力的变化

荒漠草原一直是生态学者关注的焦点。文章以宁夏灵武的荒漠草原区为研究对象,对围栏条件下退化群落的物种多样性、生产力和群落演替及其生态机制进行探讨,为荒漠草原的植被恢复和重建提供理论基础。结果表明:群落生产力除受物种多样性的影响外,也受物种本身特征和环境资源的影响。在荒漠草原中功能群盖度与群落初级生产力无显著的相关关系。荒漠植物群落物种多样性指数在时间尺度上的变化表现为随着围栏时间的延续逐渐增加,而生态优势度变化趋势则相反。在不同年份年降水量直接影响植物群落物种数的多少、丰富度和生产力。     

Individual experience-based foraging can generate community territorial structure for competing ant species

To gain additional territory while defending existing territory, animals must acquire and use information regarding resource characteristics and competitive pressure. For social organisms like ants, individual workers have experiences to acquire information, but territory establishment is a colony level behavior. Colony behavior, in turn, affects community structure. Here, I investigate how an individual ant’s previous experience affects its future foraging behavior and how individual behaviors can scale up to community territorial structure for two coexisting Formica species. To do this, I combine a field survey, a multi-agent computer simulation, and a manipulation experiment. The field survey shows that workers of both species co-occur on many trees early in the season, but ants on trees become segregated by species as the season progresses. The simulation demonstrates how this segregated spatial distribution can result from ants using a foraging strategy in which individuals show a preference for foraging sites based on previous experience. The experiment suggests that these ants are indeed capable of experience-based foraging behavior; ants preferentially return to sites where they have had positive experiences and avoid sites where they have had negative experiences. Results from this study suggest that spatially explicit information can be collected and stored by individuals to facilitate colony territorial structure, and that future investigations of community territory formation should include effects of individual previous experience.