Sting, Carry and Stock: How Corpse Availability Can Regulate De-Centralized Task Allocation in a Ponerine Ant Colony

We develop a model to produce plausible patterns of task partitioning in the ponerine ant Ectatomma ruidum based on the availability of living prey and prey corpses. The model is based on the organizational capabilities of a “common stomach” through which the colony utilizes the availability of a natural (food) substance as a major communication channel to regulate the income and expenditure of the very same substance. This communication channel has also a central role in regulating task partitioning of collective hunting behavior in a supply&demand-driven manner. Our model shows that task partitioning of the collective hunting behavior in E. ruidum can be explained by regulation due to a common stomach system. The saturation of the common stomach provides accessible information to individual ants so that they can adjust their hunting behavior accordingly by engaging in or by abandoning from stinging or transporting tasks. The common stomach is able to establish and to keep stabilized an effective mix of workforce to exploit the prey population and to transport food into the nest. This system is also able to react to external perturbations in a de-centralized homeostatic way, such as to changes in the prey density or to accumulation of food in the nest. In case of stable conditions the system develops towards an equilibrium concerning colony size and prey density. Our model shows that organization of work through a common stomach system can allow Ectatomma ruidum to collectively forage for food in a robust, reactive and reliable way. The model is compared to previously published models that followed a different modeling approach. Based on our model analysis we also suggest a series of experiments for which our model gives plausible predictions. These predictions are used to formulate a set of testable hypotheses that should be investigated empirically in future experimentation.

[1]  Brian R. Johnson A Self‐Organizing Model for Task Allocation via Frequent Task Quitting and Random Walks in the Honeybee , 2009, The American Naturalist.

[2]  P. Verhulst Notice sur la loi que la population pursuit dans son accroissement , 1838 .

[3]  J. Deneubourg,et al.  Self-organized structures in a superorganism: do ants "behave" like molecules? , 2006 .

[4]  T. Schmickl,et al.  HoPoMo : A model of honeybee intracolonial population dynamics and resource management , 2007 .

[5]  Thomas Schmickl,et al.  TaskSelSim: a model of the self-organization of the division of labour in honeybees , 2008 .

[6]  G. Robinson Modulation of alarm pheromone perception in the honey bee: evidence for division of labor based on hormonall regulated response thresholds , 1987, Journal of Comparative Physiology A.

[7]  Istvan Karsai,et al.  Organization and Regulation of Nest Construction Behavior in Metapolybia Wasps , 2004, Journal of Insect Behavior.

[8]  Karl Crailsheim,et al.  Protein trophallaxis and the regulation of pollen foraging by honey bees (Apis mellifera L.) , 1998 .

[9]  J. Lachaud,et al.  Graded recruitment and hunting strategies linked to prey weight and size in the ponerine ant Ectatomma ruidum , 1997, Behavioral Ecology and Sociobiology.

[10]  J. Deneubourg,et al.  Self-organized shortcuts in the Argentine ant , 1989, Naturwissenschaften.

[11]  Jay Wright Forrester,et al.  Urban Dynamics , 1969 .

[12]  Marco Dorigo,et al.  Division of labor in a group of robots inspired by ants' foraging behavior , 2006, TAAS.

[13]  J. Lachaud,et al.  Polyethism within hunters of the ponerine ant,Ectatomma ruidum Roger (Formicidae, Ponerinae) , 1996, Insectes Sociaux.

[14]  Thomas Schmickl,et al.  Swarm-intelligent foraging in honeybees: benefits and costs of task-partitioning and environmental fluctuations , 2010, Neural Computing and Applications.

[15]  F. Ratnieks,et al.  Task partitioning in insect societies , 1999, Insectes Sociaux.

[16]  Chris Tofts Algorithms for task allocation in ants. (A study of temporal polyethism: Theory) , 1993 .

[17]  Guy Theraulaz,et al.  Task partitioning in a ponerine ant. , 2002, Journal of theoretical biology.

[18]  J. Lachaud Spatial fidelity and individual foraging specializations in the neotropical ponerine ant, ectatomma ruidum roger (hymenoptera, formicidae) , 1995 .

[19]  Christian Bierwirth,et al.  Principles of Systems , 2000 .

[20]  S. Camazine,et al.  A model of collective nectar source selection by honey bees , 1991 .

[21]  J. Lachaud,et al.  Efficiency in the exploitation of patchy environments by the ponerine antPaltothyreus tarsatus: an ecological consequence of the flexibility of prey capture behavior , 1993, Journal of Ethology.

[22]  J. Deneubourg,et al.  The self-organizing exploratory pattern of the argentine ant , 1990, Journal of Insect Behavior.

[23]  Daniel W. McShea,et al.  The complexity and hierarchical structure of tasks in insect societies , 2001, Animal Behaviour.

[24]  M. Wells,et al.  Variations and Fluctuations of the Number of Individuals in Animal Species living together , 2006 .

[25]  E. J. Collins,et al.  The hidden cost of information in collective foraging , 2005, Proceedings of the Royal Society B: Biological Sciences.

[26]  E. Bonabeau,et al.  Fixed response thresholds and the regulation of division of labor in insect societies , 1998 .

[27]  Karl Crailsheim,et al.  Trophallactic interactions in the adult honeybee , 1997 .

[28]  Thomas Schmickl,et al.  Inner nest homeostasis in a changing environment with special emphasis on honey bee brood nursing and pollen supply , 2004 .

[29]  Raphaël Jeanson,et al.  Emergence of increased division of labor as a function of group size , 2007, Behavioral Ecology and Sociobiology.

[30]  Jean-Paul Lachaud,et al.  Individual food source fidelity in the neotropical ponerine ant Ectatomma ruidum Roger (Hymenoptera Formicidae) , 1994 .

[31]  Thomas Schmickl,et al.  Trophallaxis within a robotic swarm: bio-inspired communication among robots in a swarm , 2008, Auton. Robots.

[32]  J. Lachaud,et al.  Foraging activity and diet in some Neotropical ponerine ants. 1. Ectatomma ruidum Roger (Hymenoptera, Formicidae). , 1990 .

[33]  Thomas Schmickl,et al.  Optimisation of a honeybee-colony's energetics via social learning based on queuing delays , 2008, Connect. Sci..

[34]  E. Bonabeau,et al.  Quantitative study of the fixed threshold model for the regulation of division of labour in insect societies , 1996, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[35]  E. Wilson,et al.  The Superorganism: The Beauty, Elegance, and Strangeness of Insect Societies , 2008 .

[36]  George Kampis,et al.  Connected fragmented habitats facilitate stable coexistence dynamics , 2011 .

[37]  Stephen C. Pratt,et al.  Recruitment and Other Communication Behavior in the Ponerine Ant Ectatomma ruidum , 2010 .

[38]  Jean-Louis Deneubourg,et al.  Complexity of Environment and Parsimony of Decision Rules in Insect Societies , 2002, The Biological Bulletin.

[39]  J. Deneubourg,et al.  From individual to collective behavior in social insects , 1987 .

[40]  Chris Tofts,et al.  Foraging for work: how tasks allocate workers , 1994, Animal Behaviour.

[41]  S. Pratt,et al.  A modelling framework for understanding social insect foraging , 2003, Behavioral Ecology and Sociobiology.

[42]  Serge Kernbach,et al.  Get in touch: cooperative decision making based on robot-to-robot collisions , 2009, Autonomous Agents and Multi-Agent Systems.

[43]  John Hallam,et al.  From Animals to Animats 10 , 2008 .

[44]  Thomas Schmickl,et al.  Economic Optimisation in Honeybees: Adaptive Behaviour of a Superorganism , 2006, SAB.

[45]  D. Gordon The organization of work in social insect colonies , 1996, Nature.

[46]  Division of labour between foraging workers of the ponerine antPachycondyla caffraria (Smith) (Hymenoptera: Formicidae) , 1992, Insectes Sociaux.

[47]  Karl Crailsheim,et al.  Trophallactic interactions in the adult honeybee (Apis mellifera L.) , 1998 .

[48]  Maurice Vogels,et al.  P. F. Verhulst's “notice sur la loi que la populations suit dans son accroissement” from correspondence mathematique et physique. Ghent, vol. X, 1838 , 1975 .

[49]  T. Seeley,et al.  Collective decision-making in honey bees: how colonies choose among nectar sources , 1991, Behavioral Ecology and Sociobiology.

[50]  T. Seeley The Wisdom of the Hive: The Social Physiology of Honey Bee Colonies , 1995 .

[51]  ROBERT L. JEANNE,et al.  Regulation of nest construction behaviour in Polybia occidentalis , 1996, Animal Behaviour.

[52]  David Joyner,et al.  SAGE: system for algebra and geometry experimentation , 2005, SIGS.

[53]  R. Bernstein,et al.  FORAGING STRATEGIES OF ANTS IN RESPONSE TO VARIABLE FOOD DENSITY , 1975 .

[54]  I. Karsaia,et al.  Regulation of task partitioning by a ‘ ‘ common stomach ’ ’ : a model of nest construction in social wasps , 2011 .

[55]  P. Verhulst,et al.  Notice sur la loi que la population suit dans son accroissement. Correspondance Mathematique et Physique Publiee par A , 1838 .

[56]  Thomas Schmickl,et al.  Analysing honeybees' division of labour in broodcare by a multi-agent model , 2008, ALIFE.

[57]  R. Matthews,et al.  Ants. , 1898, Science.

[58]  A. Dornhaus,et al.  Benefits of recruitment in honey bees: effects of ecology and colony size in an individual-based model , 2006 .

[59]  G. Theraulaz,et al.  Response threshold reinforcements and division of labour in insect societies , 1998, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[60]  D. Fresneau,et al.  Social regulation in ponerine ants , 1987 .

[61]  L. Keller Social evolution in ants , 1996 .

[62]  Brian R. Johnson Global information sampling in the honey bee , 2008, Naturwissenschaften.

[63]  D. Gordon The Organization of Work in Social Insect , 2003 .

[64]  J. Lachaud,et al.  Predatory behavior of a seed‐eating ant: Brachyponera senaarensis , 1994 .

[65]  Marco Dorigo,et al.  Ant colony optimization theory: A survey , 2005, Theor. Comput. Sci..

[66]  Robert L. Eberlein,et al.  Understanding models with Vensim , 1992 .

[67]  Thomas Schmickl,et al.  Time Delay Implies Cost on Task Switching: A Model to Investigate the Efficiency of Task Partitioning , 2013, Bulletin of mathematical biology.

[68]  Thomas Schmickl,et al.  Two different approaches to a macroscopic model of a bio-inspired robotic swarm , 2009, Robotics Auton. Syst..

[69]  J. Lachaud,et al.  Étude des stratégies d'approvisionnement chez 3 espèces de fourmis ponérines (Hymenoptera, Formicidae) , 1984 .

[70]  M. Gilpin,et al.  Global models of growth and competition. , 1973, Proceedings of the National Academy of Sciences of the United States of America.

[71]  Walter R. Tschinkel,et al.  Task selection by workers of the fire ant Solenopsis invicta , 1999, Behavioral Ecology and Sociobiology.

[72]  Thomas Schmickl,et al.  The "Dance or Work" Problem: Why Do not all Honeybees Dance with Maximum Intensity , 2005, CEEMAS.

[73]  Vito Volterra,et al.  Leçons sur la théorie mathématique de la lutte pour la vie , 1931 .

[74]  Istvan Karsai,et al.  Regulation of task differentiation in wasp societies: a bottom-up model of the "common stomach". , 2012, Journal of theoretical biology.

[75]  G. Robinson Regulation of division of labor in insect societies. , 1992, Annual review of entomology.

[76]  Thievery, home ranges, and nestmate recognition inEctatomma ruidum , 1990, Oecologia.

[77]  Manuel López-Ibáñez,et al.  Ant colony optimization , 2010, GECCO '10.

[78]  D. Sumpter,et al.  From nonlinearity to optimality: pheromone trail foraging by ants , 2003, Animal Behaviour.

[79]  Gábor Balázsi,et al.  Organization of work via a natural substance: regulation of nest construction in social wasps. , 2002, Journal of theoretical biology.

[80]  Patrick Brézillon,et al.  Lecture Notes in Artificial Intelligence , 1999 .

[81]  J. Lachaud Recruitment by selective activation: an archaic type of mass recruitment in the ponerine ant (Ectatomma ruidum) , 1985 .

[82]  John N. Warfield,et al.  World dynamics , 1973 .

[83]  Thomas Schmickl,et al.  Collective Perception in a Robot Swarm , 2006, Swarm Robotics.

[84]  Guy Theraulaz,et al.  The formation of spatial patterns in social insects: from simple behaviours to complex structures , 2003, Philosophical Transactions of the Royal Society of London. Series A: Mathematical, Physical and Engineering Sciences.

[85]  I. Karsai,et al.  Productivity, individual-level and colony-level flexibility, and organization of work as consequences of colony size. , 1998, Proceedings of the National Academy of Sciences of the United States of America.

[86]  Phil Husbands,et al.  Artificial Life IX: Proceedings of the Ninth International Conference on the Simulation and Synthesis of Living Systems , 2004 .

[87]  Horst F. Wedde,et al.  A comprehensive review of nature inspired routing algorithms for fixed telecommunication networks , 2006, J. Syst. Archit..

[88]  E. Wilson,et al.  Caste and ecology in the social insects. , 1979, Monographs in population biology.

[89]  N. Franks,et al.  Teams in animal societies , 2001 .