Ant Foraging and Climate Change July 2010 Ashley Jackson, John Collins, and Ngoc Thai One of the most prominent questions reasons, it is imperative that we consider we, as scientists and civilians, have is: ants as an important part in future research “What could happen to the world if the in climate change. temperature rises overall during the next century?” What about the next decade? If we study the behavior of ant species throughout the country, we may discover an answer. Ants have been labeled by researchers as an indicator species because Fig. 1 The Lotka-Volterra competition equations are simple model equations that show in a constant environment, “complete competitors cannot coexist” (Hardin, Science, 1960). These equations were one of the first proposed models of competition between species. they are more susceptible to disturbance and climatic change (Chapman and Bourke, EL, 2001). An indicator species may act as an early warning to certain changes in the environment, whether natural or unnatural. One key trait that makes ants a strong indicator species is their short life cycle. It is much less time consuming to observe changes in a species that does not take years to start new generations. In addition to being named as an indicator species, ants have also been called a keystone species. A keystone species is a species that contributes to its environment in a way such that if it died, it would affect many different other species, including plants. Ants are important for soil turnover, litter processing and decomposition (Holec and Frouz, Pedobiologia, 2005), and seed dispersal (Beattie, Cambridge Univ., 1985). For these However, there are many species that live in the same habitat and are able to coexist. For example, ants have this ability. Within ant communities, ant species compete for certain resources and niches. The idea of occupying niches (Grinnell, AUK, 1917) is one of the reasons competing ant species are able to coexist in habitats. Another reason for this coexistence is tradeoffs between species. Many plants coexist through a competition/colonization trade-off (Hardin, Science, 1960). An example of trade-offs between ants is the fact that one species may be able to find food more quickly, but another species may be better at defending and displacing the other species once they have found the food. Furthermore, one species may be able to forage at a higher range of temperatures than another. This 1 Ant Foraging and Climate Change July 2010 Ashley Jackson, John Collins, and Ngoc Thai particular trade-off is a major concern when the issue of climate change arises. Climate change may favor one species over another. If this happens, certain niches may be left unoccupied. If the newly unoccupied niche is one that impacts humans or closely enough, we may be affected in ways that could change our everyday comfort in life. If the extinction of certain species is drastic Fig. 3 A dominance-thermal tolerance trade-off curve enough, it could lead to partial loss of (Lessard, Dunn, and Sanders, IS, 2009). biodiversity. Biodiversity is important to maintain a natural balance in nature. In the figures below, each one represents a tradeoff curve. There is also a graph that shows loss of biodiversity by means of a species dying out. Fig. 4 This graph is merely a cartoon of what could happen over time. It is in no way an actual representation of any proven data. “No studies, to our knowledge, have experimentally examined how projected temperature increases will affect ant Fig. 2 This shows a dominance-discovery trade-off curve (Lebrun, JAE, 2007). communities and the functions they provide,” (Sanders et al., in press). However, at the end of the Ice Age, we were left with the extinctions of the woolly 2 Ant Foraging and Climate Change July 2010 Ashley Jackson, John Collins, and Ngoc Thai mammoth and the saber-tooth tiger, as well movement variation. A constant number of as many other species. These losses have food pieces are dropped every morning. been accredited to the drastic climatic Food is also dropped throughout the day changes that occurred. At the same time, with a 0.1 chance every time step. The some species thrived during the Ice Age. It probability that an ant will wait during a may prove beneficial to study which species time step depends on both temperature and will be favored in the event that the foraging success. If an ant has taken more temperature does increase. We may even be than 10 consecutive moves without finding a able to provide theories and suggestions to piece of food, then the ant has a chance of help maintain as much biodiversity as waiting depending on species and possible. temperature. Ants will resume their normal There are many questions we could attempt to answer dealing with ants and climate change. Our question, in particular: How will climate change affect the competition between hot-tolerant and coldtolerant ants? We decided to use a multiscale, particularly a two-scale, model to attempt to address this issue. Our model consists of two parts, a small-scale and a large-scale. The small-scale model contains localized rules that are easily added and changed and is run using NetLogo. It is an agent-based model with two species of ants competing for the same resources. It can test different foraging rules such as ants waiting under certain conditions. Rules include running speed, food specialization, and moving pattern when they find food. Temperature and foraging rules affect both foraging success and foraging opportunities on a daily basis. The large-scale model contains more complicated rules and is run using MATLAB. Climate and food haul affect both colony success and reproduction on a monthly basis. Both of these models have their advantages. The small-scale model is mainly used for gathering statistics and data points to construct various functions that are then used in the largescale model. The large-scale model will be useful in gathering data for longer periods of time, for instance hundreds of years, given enough time. Within both models, we have used only two different species for simplicity and time constraints; however, it 3 Ant Foraging and Climate Change July 2010 Ashley Jackson, John Collins, and Ngoc Thai is tractable to add more species to create a more realistic model. Fig. 7 A gradual change in yearly temperature tips the Fig. 5 This shows the waiting probability of each ant balance in favor of the hot tolerant ant species. species dependent upon temperature ranges. We managed to gather and plot the population data for constant yearly temperature, gradually changing yearly temperature, and rapidly changing yearly temperature. The following figures represent those plots, respectively. Fig. 8 Rapidly changing yearly temperature first significantly favors the cold tolerant ants. After the temperature reaches about 30 degrees, there isn’t a very significant difference in each species. The data gathered was the result of the following functional forms that appeared in our MATLAB model. Fig. 6 With no change in yearly average temperature, both species stay proportional. Fig. 9 This is the function that represents the death that occurs in the model. 4 Ant Foraging and Climate Change July 2010 Ashley Jackson, John Collins, and Ngoc Thai Fig. 10 This function represents how much energy it costs to move, metabolism (Lighton and Bartholomew, PNAS, 1988). Fig. 11 This function represents the reproduction and when it occurs. Fig. 5 The haul function was a direct result of the data gathered from our small scale NetLogo model. Fig. 12 This is the number of food pieces stored by species All of the above are functions that were A (hot-tolerant) at month i. determined by analyzing the statistics we collected while doing runs using the small- Fig. 13 The number of food pieces stored by species B scale model. These functions were then (cold-tolerant) at month i. placed into the large-scale model to graph surface plots. Using the different foraging Fig. 14 The number of ants in species A at month i. rules, we were able to manipulate circumstances that favored one species over Fig. 15 The number of ants in species B at month i. We did many runs to gather the data in the graphs. The runs were done over the course of time that we spent once we had perfected the large scale model. From the data gathered from the small scale model, we were able to produce a Haul Function, which is graphed below. the other. This is how we expect to propose theories as to what climate change may do to ant communities. Our future goals for this project include scaling the model up to 3000 ants, adding sensitivity analyses for parameters and initial conditions, and adding stability analysis. During the 8 week program at NIMBioS, we were able to determine a 5 Ant Foraging and Climate Change July 2010 Ashley Jackson, John Collins, and Ngoc Thai problem, draw up a model, and propose the conservation biology of social possible outcomes for our problem. This insects. Ecology Letters, 4: 650–662. was done with the help of our advisors, Dr. Holec, M. and Frouz, J. (2005), Ant Sharon Bewick and Dr. Steven Wise. (Hymenoptera: Formicidae) Though, during the beginning, we met with communities in reclaimed and our advisors 3-4 times a week, we quickly unreclaimed brown coal mining spoil grew independent and began to work dumps in the Czech Republic. amongst ourselves much more often. This Pedobiologia, 49: 345-357. program taught us many useful skills. Beattie, A. J. (1985), The Different educational backgrounds meant evolutionary ecology of ant-plant that mutualisms. Cambridge University each moment, you could learn something new that you did not come into the program knowing. We all learned from each other. This interdisciplinary made and for diverse a Press. truly Exclusion Principle learning experience that will not soon be forgotten. Hardin, G. (1960), The Competitive Science 131 (3409), 1292. Grinnell, J. (1917), The NicheRelationships of the California Acknowledgements: Thrasher. The Auk, 34: 427-433. We would like to thank the National Institute for Mathematical and Biological Synthesis, the LEBRUN, E. G. and FEENER, D. H. (2007), When trade-offs interact: University of Tennessee at Knoxville, and our balance of terror enforces dominance advisors, Sharon Bewick and Steven Wise. Each discovery trade-off in a local ant party played a critical role in the success of our assemblage. Journal of Animal summer research program. Ecology, 76: 58–64. References: Chapman, R. E. and Bourke, A. F. G. (2001), The influence of sociality on Lessard, J. –P., Dunn, R. R., and Sanders, N. J., (2009), Temperaturemediated coexistence in temperate forest ant communities. Insectes Sociaux, 56: 149-156. 6 Ant Foraging and Climate Change July 2010 Ashley Jackson, John Collins, and Ngoc Thai Lighton, J. R. B. and Bartholomew, G. A., (1988), Standard energy metabolism of a desert harvester ant, Pogonomyrmex rugosus: Effects of temperature, body mass, group size, and humidity. PNAS 85 (13): 47654769. 7