@phdthesis{Roemer2014,
  author    = {R{\"o}mer, Daniela},
  title     = {Where and how to build? Influence of social and environmental cues on nest building behavior in leaf-cutting ants},
  url       = {http://nbn-resolving.de/urn:nbn:de:bvb:20-opus-109409},
  school      = {Universit{\"a}t W{\"u}rzburg},
  year      = {2014},
  abstract  = {This thesis explores the influence of social and environmental cues on the nest building behavior of leaf-cutting ants. Especially, the investigations are aimed at evaluating the mechanisms of nest building and how the nest environment can spatially guide building responses that lead to an adaptive nest architecture. The emergence of nest chambers in the nest of the leaf-cutting ant Acromyrmex lundi were evaluated. Rather than excavating nest chambers in advance, at places where workers encounter suitable environmental conditions for brood and fungus rearing, these items have to be present at a site. When presented in the laboratory with a choice between two otherwise identical digging sites, offering suitable environmental conditions, but one containing brood, the workers displayed a higher excavation activity at the site where they encountered the putative content of a chamber. The shape of the excavated cavity was also more round and chamber-like. It is concluded that leaf-cutting ants respond to social cues during nest building. Excavation is a costly process and colonies have to spend a part of their energy stores on nest building, so that regulatory responses for the control of nest excavation are expected to occur. Worker density at the beginning of the digging process influenced digging activity while the presence of in-nest stores did not. Stored brood and fungus did however influence the architecture of the excavated nest, leading to the excavation of larger chambers and smaller tunnels. While self-organized mechanisms appear to be involved in the nest building process, the social cues of the ants' environment during building clearly influence the nest architecture and lead to an adjustment of the nest size to the current space needs of the colony. Workers secondarily regulated nest size by the opportunistic refilling of unused space with excavated soil pellets. As the ants should provide suitable conditions for brood and fungus rearing, they should show a behavioral response to CO2 concentrations, as the gas is known to hinder fungus respiration. Workers of A. lundi did indeed avoid high CO2-levels for fungus rearing but actually preferred CO2-values in the range encountered close to the soil surface, where this species excavates their nests. However, different CO2-levels did not affect their excavation behavior. While fungus chambers make up part of a leaf-cutting ant nest, most leaf-cutting ants of the genus Atta also spent part of the colony's energy on excavating large, voluminous chambers for waste disposal, rather than scattering the material aboveground. It is expected that leaf-cutting ants also show environmental preferences for waste management. In experiments Atta laevigata workers preferred deposition in a warm and dry environment and showed no preference for specific CO2-levels. The continued accumulation of waste particles in a waste chamber seems to be based on the use of volatiles. These originate from the waste itself, and seem to be used as an orientation cue by workers relocating the material. The ensuing large accumulation of waste at one site should result in the emergence of more voluminous chambers for waste disposal.},
  subject      = {Nestbau},
  language  = {en}
}
@phdthesis{Pielstroem2013,
  author    = {Pielstr{\"o}m, Steffen},
  title     = {On the Role of Local Information in the Spatial Organisation of Collective Nest Digging in the Leaf-Cutting Ant Atta vollenweideri (Forel, 1893)},
  url       = {http://nbn-resolving.de/urn:nbn:de:bvb:20-opus-79118},
  school      = {Universit{\"a}t W{\"u}rzburg},
  year      = {2013},
  abstract  = {Many ant species excavate underground nests. One of the most impressive examples is the Chaco leaf-cutting ant Atta vollenweideri from the Gran Chaco region in South America. The nests excavated by the workers of that species are among the largest insect-built structures on the planet. They are ecavated over years possibly involving millions of working individuals. However, the mechanisms underlying the organisation of collective nest digging in ants remain largely unknown. Considering the sheer dimensions of the nest in comparison to the size and presumably limited perceptual and cognitive abilities of the single worker, the assumption can be made that organising mechanisms are mostly based on responses of individuals to local stimuli within their perceptual range. Among these local stimuli that guide nest digging we can expect environmental variables, stimuli that relate to the requirements of the colony, and stimuli related to the spatial coordination of collective effort. The present thesis investigates the role of local stimuli from these three categories in the organisation of collective digging behaviour in the Chaco leaf-cutting ant. It describes experiments on (1) how workers respond in the context of digging to differences in soil moisture, which comprises an important environmental variable; (2) how available nest space influences nest enlargement; (3) and how the spatial coordination of excavating workers is implemented by responding to stimuli arising from nest mates while engaged in digging behaviour. The experiments on soil water content show that workers prefer to dig in moist materials that allow for fast excavation and transport rates. Accordingly, an unequal distribution of water in the soil around a nest can influence how the nest shape develops. On the other hand, results also indicate that workers strongly avoid excavating in extremely moist materials. Regarding the abundant occurrence of flooding events in the Gran Chaco region, the latter can be interpreted as an adaptation to avoid water inflow into the nest. In the experiments on the effect of nest space, the ants excavated less when presented with larger nests. When a large amount of space was suddenly added to the nest during the digging process, excavation rates decreased according to the new volume. These observations confirm the hypothesis that digging activity is regulated according to space requirements, possibly because crowding conditions inside the nest influence excavation behaviour. However, observations also indicate an intrinsic decrease of digging motivation with time. Moreover, excavation rates correlate with nest size only when comparing nests of similar shape. Distributing a similar nest volume to three smaller chambers, instead of one, resulted in drastically decreased digging rates. A possible explanation for that observation lies in the distribution of workers inside the nest that may vary according to nest geometry: a different distribution of individuals can lead to in different local crowding conditions in similar nest volumes. Furthermore, two different stimuli are described that are used in the spatial coordination of collective digging effort. First, fresh soil pellets deposited close to the digging site on their way from the surface increase the probability that arriving workers join excavation efforts at the same site. The deposition of pellets on the way is a consequence of sequential task partitioning during soil transport. The pellets are carried in transport chains that closely resemble the modalities of leaf transport observed at the surface. Second, workers stridulate while digging. The short-ranged vibrational signals produced thereby also attract nest mates to excavate at the same location. Accordingly, two mutually complementing mechanisms are described that allow to concentrate excavators at one location. In both cases, a local stimulus that is generated by current close-by excavation activity increases the probability of the stimulus receiver to dig close to other excavators. In an environment otherwise poor in digging stimuli, these mechanisms can be especially important to give collective digging efforts a common direction. As a consequence it can be argued that the spatial organisation of collective digging is based on choice copying. Individuals copy nest mate decisions on where to excavate by responding to local stimuli provided by nest mate digging activity. Taken together, responses to local stimuli can determine the direction of nest growth, aid in preventing the inflow of surface water into the nest, guide the adjustment of nest size to colony requirements and spatially coordinate collective digging efforts. Even though it cannot be ruled out that digging responses based e.g. on spatial memory or long-term experience exist, the results presented here clearly demonstrate that responses to local information account for many important aspects of nest development.},
  subject      = {Blattschneiderameisen},
  language  = {en}
}
@article{BollazziRoces2011,
  author    = {Bollazzi, Martin and Roces, Flavio},
  title     = {Information Needs at the Beginning of Foraging: Grass-Cutting Ants Trade Off Load Size for a Faster Return to the Nest},
  url       = {http://nbn-resolving.de/urn:nbn:de:bvb:20-opus-68940},
  year      = {2011},
  abstract  = {Background: Acquisition of information about food sources is essential for animals that forage collectively like social insects. Foragers deliver two commodities to the nest, food and information, and they may favor the delivery of one at the expenses of the other. We predict that information needs should be particularly high at the beginning of foraging: the decision to return faster to the nest will motivate a grass-cutting ant worker to reduce its loading time, and so to leave the source with a partial load. Principal Findings: Field results showed that at the initial foraging phase, most grass-cutting ant foragers (Acromyrmex heyeri) returned unladen to the nest, and experienced head-on encounters with outgoing workers. Ant encounters were not simply collisions in a probabilistic sense: outgoing workers contacted in average 70\% of the returning foragers at the initial foraging phase, and only 20\% at the established phase. At the initial foraging phase, workers cut fragments that were shorter, narrower, lighter and tenderer than those harvested at the established one. Foragers walked at the initial phase significantly faster than expected for the observed temperatures, yet not at the established phase. Moreover, when controlling for differences in the fragment-size carried, workers still walked faster at the initial phase. Despite the higher speed, their individual transport rate of vegetable tissue was lower than that of similarly-sized workers foraging later at the same patch. Conclusions/Significance: At the initial foraging phase, workers compromised their individual transport rates of material in order to return faster to the colony. We suggest that the observed flexible cutting rules and the selection of partial loads at the beginning of foraging are driven by the need of information transfer, crucial for the establishment and maintenance of a foraging process to monopolize a discovered resource.},
  subject      = {Blattschneiderameisen},
  language  = {en}
}
@phdthesis{Saverschek2010,
  author    = {Saverschek, Nicole},
  title     = {The influence of the symbiotic fungus on foraging decisions in leaf-cutting ants - Individual behavior and collective patterns},
  url       = {http://nbn-resolving.de/urn:nbn:de:bvb:20-opus-52087},
  school      = {Universit{\"a}t W{\"u}rzburg},
  year      = {2010},
  abstract  = {Foraging behavior is a particularly fascinating topic within the studies of social insects. Decisions made by individuals have effects not only on the individual level, but on the colony level as well. Social information available through foraging in a group modulates individual preferences and shapes the foraging pattern of a colony. Identifying parameters influencing foraging behavior in leaf-cutting ants is especially intriguing because they do not harvest for themselves, but for their symbiotic fungus which in turn influences their plant preferences after the incorporation of the substrate. To learn about the substrates' unsuitability for the fungus, ants need to be able to identify the incorporated substrate and associate it with detrimental effects on the fungus. Odor is an important plant characteristic known to be used as recognition key outside the nest in the context of foraging. Chapter 1 shows that foragers are able to recall information about the unsuitability of a substrate through odor alone and consequently reject the substrate, which leads to the conclusion that inside the nest, odor might be enough to indentify incorporated substrate. Identification of plant species is a key factor in the foraging success of leaf-cutting ants as they harvest a multitude of different plant species in a diverse environment and host plant availability and suitability changes throughout the year. Fixed plant preferences of individuals through innate tendencies are therefore only one factor influencing foraging decisions. On the individual as well as the colony level, foraging patterns are flexible and a result of an intricate interplay between the different members involved in the harvesting process: foragers, gardeners and the symbiotic fungus. In chapter 2 I identified several conditions necessary for na{\"i}ve foragers to learn about the unsuitability of substrate inside the nest. In order to exchange of information about the unsuitability of a substrate, the plant in question must be present in the fungus garden. Foragers can learn without own foraging experience and even without experiencing the effects of the substrate on the fungus, solely through the presence of experienced gardeners. The presence of experienced foragers alone on the other hand is not enough to lower the acceptance of substrate by na{\"i}ve foragers in the presence of na{\"i}ve gardeners, even if experienced foragers make up the majority of the workforce inside the nest. Experienced foragers are also able to reverse their previous negative experience and start accepting the substrate again. The individual behavior of foragers and gardeners with different experiential backgrounds in the presence of suitable or unsuitable substrate inside the fungus chamber was investigated in chapter 3 to shed some light on possible mechanisms involved in the flow of information about substrate suitability from the fungus to the ants. Gardeners as well as foragers are involved in the leaf processing and treatment of the applied leaf patches on the fungus. If the plant material is unsuitable, significantly more ants treat the plant patches, but foragers are less active overall. Contacts between workers initiated by either gardeners or foragers occur significantly more frequent and last longer if the substrate is unsuitable. Even though experienced gardeners increase na{\"i}ve foragers' contact rates and duration with other workers in the presence of suitable plant patches, na{\"i}ve foragers show no differences in the handling of the plant patches. This suggests that foragers gain information about plant suitability not only indirectly through the gardening workers, but might also be able to directly evaluate the effects of the substrate on the fungus themselves. Outside the nest, foragers influence each other the trail (chapter 4). Foraging in a group and the presence of social information is a decisive factor in the substrate choice of the individual and leads to a distinct and consentaneous colony response when encountering unfamiliar or unsuitable substrates. As leaf-cutting ants harvest different plant species simultaneously on several trails, foragers gain individual experiences concerning potential host plants. Preferences might vary among individuals of the same colony to the degree that foragers on the same trail perceive a certain substrate as either suitable or unsuitable. If the majority of foragers on the trail perceives one of the currently harvested substrates as unsuitable, na{\"i}ve foragers lower their acceptance within 4 hours. In the absence of a cue in the fungus, na{\"i}ve foragers harvesting by themselves still eventually (within 6 hours) reject the substrate as they encounter experienced gardeners during visits to the nest within foraging bouts. As foraging trails can be up to 100 m long and foragers spend a considerable amount of time away from the nest, learning indirectly from experienced foragers on the trail accelerates the distribution of information about substrate suitability. The level of rejection of a formerly unsuitable substrate after eight hours of foraging by na{\"i}ve foragers correlates with the average percentage of unladen experienced foragers active on the trail. This suggests that unladen experienced foragers might actively contact laden na{\"i}ve workers transmitting information about the unsuitability of the load they carry. Results from experiments were I observed individual laden foragers on their way back to the nest backed up this assumption as individuals were antennated and received bites into the leaf disk they carried. Individuals were contacted significantly more often by nestmates that perceived the carried leaf disk as unsuitable due to previous experience than by nestmates without this experience (chapter 6). Leaf-cutting ants constantly evaluate, learn and re-evaluate the suitability of harvested substrate and adjust their foraging activity accordingly. The importance of the different sources of information within the colony and their effect on the foraging pattern of the colony depend on the presence or absence of each of them as e.g. experienced foragers have a bigger influence on the plant preferences of na{\"i}ve foragers in the absence of a cue in the fungus garden.},
  subject      = {Blattschneiderameisen},
  language  = {en}
}
@phdthesis{BollazziSosa2008,
  author    = {Bollazzi Sosa, Leonardo Martin},
  title     = {Building behaviour and the control of nest climate in Acromyrmex leaf-cutting ants},
  url       = {http://nbn-resolving.de/urn:nbn:de:bvb:20-opus-27610},
  school      = {Universit{\"a}t W{\"u}rzburg},
  year      = {2008},
  abstract  = {This work was aimed at experimentally studying whether climatic variables act as environmental cues for workers' building behaviour in leaf-cutting ants of the genus Acromyrmex, and to what extent building responses account for the maintenance of nest climate in a proper range for the inhabiting colony. Specifically, this work presents independent analysis in different Acromyrmex species with disparate ecology and nesting habits, aimed at understanding to what extent: i) temperature and humidity act as cues for workers' building behaviour, ii) inter- and intraspecific differences in the nesting habits observed in South American Acromyrmex are based on distinct building behaviours and on the variation in regional climate across continent, iii) differences in nest architecture account for the maintenance of nest climate in a proper range for colony members and, iv) climatic variables trigger building responses aimed at controlling short-term changes in nest climate. It is first experimentally shown that soil temperature acts as a cue for workers' digging behaviour. Acromyrmex lundi workers were observed to respond to both soil temperature as well as its changes, and to decide accordingly where to start or whether to stop digging. The soil temperature range preferred by workers to dig, between 20°C and maximally 30.6°C, matches the range at which colony growth is expected to be maximized. Temperature-sensitive digging might therefore lead to the establishment of the fungus chambers in soil layers with a proper range of temperatures for colony growth. Based on that, it was hypothesized that nest depth in Acromyrmex largely depends on the depth at which this temperature range is located across the soil profile, i.e., the higher the temperature in the superficial soil layers, the deeper the nest location, since soil temperature decreases with increasing depth. A bibliographic survey on nesting habits of 21 South American Acromyrmex species confirmed that the warmer the soil temperature at 50 cm depth throughout the South American continent, the higher the number of species presenting subterranean nests, compared with those inhabiting superficial nests. Temperature-sensitive digging in Acromyrmex would therefore explain the geographical distribution of nesting habits observed for this genus in the South American continent, i.e., subterranean in the northern tropical regions, and superficial in the southern temperate ones. In addition, results showed that Acromyrmex colonies from temperate regions indeed achieve thermoregulatory benefits through the determination of nest depth based on thermoregulatory needs. In sympatrically-occurring colonies of the grass-cutting ant A. heyeri, temperature inside superficial thatched nests was higher, and more suitable for colony growth, than that inside subterranean nests. This temperature surplus was even higher in spring, at the time of production of sexual brood, than in winter or summer. It was demonstrated that such temperature surplus was brought about by the low thermal diffusivity of the nest thatch, which prevents diurnal nest overheating by the incoming solar radiation, and avoids losses of the accumulated daily heat into the cold air during night, thus leading to high average nest temperatures. Although highly advantageous for colonies in terms of nest temperature, the determination of nest depth based on thermoregulatory needs may differentially affect nest ventilation and humidity depending on how nest exposition influences the exchange of nest air with the outside air. For instance, colonies with a superficial nesting habit might benefit from improved nest ventilation, but be at risk of desiccation due to their exposition and the consequent humidity losses into the dry outside air. Results demonstrated that in two Acromyrmex species, short-term regulatory building responses triggered and spatially organized by climatic variables occur, and may counteract undesired changes in internal nest humidity. Workers of the thatching grass-cutting ant A. heyeri, for instance, closed a number of nest-thatch openings as a response to desiccation of the outside air, even at a nest temperature that otherwise triggered the response of opening them so as to reduce nest temperature. In the leaf-cutting ant A. ambiguus, the direction of the airflow inside nest tunnels was shown to act as a cue for spatially guiding the building behaviour of plugging nest entrances. However, workers only responded if the humidity content of the circulating air was low, trading therefore nest ventilation for humidity maintenance.},
  subject      = {Verhaltens{\"o}kologie},
  language  = {en}
}
@phdthesis{Kleineidam1999,
  author    = {Kleineidam, Christoph},
  title     = {Sensory Ecology of Carbon Dioxide Perception in Leaf-cutting Ants},
  url       = {http://nbn-resolving.de/urn:nbn:de:bvb:20-opus-1562},
  school      = {Universit{\"a}t W{\"u}rzburg},
  year      = {1999},
  abstract  = {The study examines the sensory ecology of CO2 perception in leaf-cutting ants. It begins with the ecological role of CO2 for leaf-cutting ants. Inside the subterranean nests of Atta vollenweideri large amounts of CO2 are produced by the ants and their symbiotic fungus. Measurements in field nest at different depths revealed that CO2 concentrations do not exceed 2 per cent in mature nests. These findings indicate effective ventilation even at depths of 2 m. Small colonies often face the situation of reduced ventilation when they close their nest openings as a measure against flooding. A simulation of this situation in the field as well as in the laboratory revealed increasing CO2 concentrations causing reduced colony respiration which ultimately might limit colony success. Wind-induced ventilation is the predominant ventilation mechanism of the nests of Atta vollenweideri, shown by an analysis of external wind and airflow in the channels. The mound architecture promotes nest ventilation. Outflow channels have their openings in the upper, central region and inflow channels had their openings in the lower, peripheral region of the nest mound. Air is sucked out through the central channels, followed by a delayed inflow of air through the peripheral channels. The findings support the idea that the nest ventilation mechanism used by Atta vollenweideri resembles the use of Bernoulli's principle in Venturi Tubes and Viscous Entrainment. CO2 is important in a second context besides microclimatic control. A laboratory experiment with Atta sexdens demonstrated that leaf-cutting ants are able to orientate in a CO2 gradient. Foragers chose places with higher CO2 concentration when returning to the nest. This effect was found in all homing foragers, but it was pronounced for workers carrying leaf fragments compared to workers without leaf fragments. The findings support the hypothesis that CO2 gradients are used as orientation cue inside the (dark) nest to find suited fungus chambers for unloading of the leaf fragments. After the importance of CO2 in the natural history of the ants has thus been demonstrated, the study identifies for the first time in Hymenoptera type and location of the sensory organ for CO2 perception. In Atta sexdens a single neuron associated with the sensilla ampullacea was found to respond to CO2. Since it is the only neuron of this sensillum, the sensillum characters can be assumed to be adapted for CO2 perception. A detailed description of the morphology and the ultrastructure allows a comparison with sensilla for CO2 perception found in other insects and provides more information about sensillum characters and their functional relevance. The CO2 receptor cells respond to increased CO2 with increased neural activity. The frequency of action potentials generated by the receptor cell shows a phasic-tonic time course during CO2 stimulation and a reduced activity after stimulation. Phasic response accomplished with a reduced activity after stimulation results in contrast enhancement and the ability to track fast fluctuations in CO2 concentration. The neurons have a working range of 0 to 10 per cent CO2 and thus are able to respond to the highest concentrations the ants might encounter in their natural environment. The most exciting finding concerning the receptor cells is that the CO2 neurons of the leaf-cutting ants do not adapt to continuous stimulation. This enables the ants to continuously monitor the actual CO2 concentration of their surroundings. Thus, the sensilla ampullacea provide the ants with the information necessary to orientate in a CO2 gradient (tracking of fluctuations) as well as with the necessary information for microclimatic control (measuring of absolute concentrations).},
  subject      = {Blattschneiderameisen},
  language  = {en}
}