Complexity of searching movement in the European harvester ant Messor wasmanni: effect of temperature and body size

Research Article


Individual search behavior is an important feature of ant ecology that can be influenced by many factors including environmental constraints, competitive interactions, and morphology. In this study, I investigated the effects of temperature, body size, and leg allometry on search movement in the polymorphic European harvester ant, Messor wasmanni. Temperature is known to increase movement speed, but how it interacts with morphology to influence path sinuosity is unknown. Ants in this study were grouped according to dimension (small, medium, and large) and were filmed moving individually in a neutral arena at three different temperatures: 10, 20, and 30 °C. Movements were analyzed by evaluating mean speed, path length, and path complexity using 5 different indexes to survey tortuosity: Mean Square Displacement (MSD), Straightness (ST), Intensity of Habitat Use (IU), Sinuosity (SI), and Fractal Dimension (Fractal D). Search movement seemed not to be strongly affected by size, even though extreme temperatures faintly diversified the path complexity among castes. Finally, the relationship between body weight and leg length was evaluated to test the ‘size-grain’ hypothesis among dimensional castes. Mean speed was strongly affected by temperature, but not by the size of the ants. Complexity indexes revealed different outcomes through their relationship to speed or path length. Interestingly, leg length and body size were negatively correlated, indicating that legs are proportionally shorter in large rather than in small ants, causing rejection of the ‘size-grain’ hypothesis for this species. Thus, individuals experience a similar roughness of soil surface irrespective of their size.


Ant movement Searching behavior Harvester ants Tortuosity Size-grain 



I would like to thank Alberto Masoni for his help during laboratory experiments, Giacomo Santini and Elena Ricevuto for their help during the review of the manuscript. Moreover, I would thank the anonymous reviewers that strongly contributed to improving the consistency of the manuscript. This project was funded by the University of Florence and does not conflict with other interests.


  1. Abramoff MD, Magalhães PJ, Ram SJ (2004) Image processing with ImageJ. Biophoton Int 11:36–42Google Scholar
  2. Adler FR, Gordon DM (2003) Optimization, conflict, and nonoverlapping foraging ranges in ants. Am Nat 162:529–543CrossRefPubMedGoogle Scholar
  3. Alexander R, Jayes AS (1983) A dynamic similarity hypothesis for the gaits of quadrupedal mammals. J Zool 201:135–152CrossRefGoogle Scholar
  4. Alexander RM (2005) Models and the scaling of energy costs for locomotion. J Exp Biol 208:1645–1652CrossRefPubMedGoogle Scholar
  5. Almeida PJ, Vieira MV, Kajin M, Forero-Medina G, Cerqueira R (2010) Indices of movement behaviour: conceptual background, effects of scale and location errors. Zoologia 27:674–680CrossRefGoogle Scholar
  6. Angilletta MJ, Roth II, Wilson RS, Niehaus AC, Ribeiro PL (2008) The fast and the fractalous: speed and tortuosity trade off in running ants. Funct Ecol 22:78–83Google Scholar
  7. Azcárate FM, Kovacs E, Peco B (2007) Microclimatic conditions regulate surface activity in harvester ants Messor barbarus. J Insect Behav 20:315–329CrossRefGoogle Scholar
  8. Baraibar B, Torra J, Westerman PR (2011) Harvester ant (Messor barbarus (L.)) density as related to soil properties, topography and management in semi-arid cereals. Appl Soil Ecol 51:60–65CrossRefGoogle Scholar
  9. Bartumeus F, da Luz MGE, Viswanathan GM, Catalan J (2005) Animal search strategies: a quantitative random-walk analysis. Ecology 86:3078–3087CrossRefGoogle Scholar
  10. Batschelet E (1981) Circular statistic in Biology. Academic Press, LondonGoogle Scholar
  11. Bell W (1990) Searching Behavior: the Behavioral Ecology of Finding Resources. Chapman and Hall, New YorkCrossRefGoogle Scholar
  12. Bernadou A, Fourcassié V (2008) Does substrate coarseness matter for foraging ants? An experiment with Lasius niger (Hymenoptera; Formicidae). J Insect Physiol 54:534–542CrossRefPubMedGoogle Scholar
  13. Benhamou S (2004) How to reliably estimate the tortuosity of an animal’s path: straightness, sinuosity, or fractal dimension? J Theor Biol 229:209–220CrossRefPubMedGoogle Scholar
  14. Benhamou S (2014) Of scales and stationarity in animal movements. Ecol Lett 17:261–272CrossRefPubMedGoogle Scholar
  15. Bennett AF (1984) Thermal dependence of muscle function. Am J Physiol 247:217–229CrossRefGoogle Scholar
  16. Beverly BD, McLendon H, Nacu S, Holmes S, Gordon DM (2009) How site fidelity leads to individual differences in the foraging activity of harvester ants. Behav Ecol 20:633–638CrossRefGoogle Scholar
  17. Bovet P, Benhamou S (1991) Optimal sinuosity in central place foraging movements. Anim Behav 42:57–62CrossRefGoogle Scholar
  18. Bovet P, Benhamou S (1988) Spatial analysis of animals’ movements using a correlated random walk model. J Theor Biol 131:419–433CrossRefGoogle Scholar
  19. Bowler DE, Benton TG (2005) Causes and consequences of animal dispersal strategies: relating individual behaviour to spatial dynamics. Biol Rev 80:205–225CrossRefPubMedGoogle Scholar
  20. Baraibar B, Torra J, Westerman PR (2011) Harvester ant (Messor barbarus (L.)) density as related to soil properties, topography and management in semi-arid cereals. Appl Soil Ecol 51:60–65CrossRefGoogle Scholar
  21. Burnham KP, Anderson DR (2002) Model selection and multimodel inference: a practical information-theoretic approach. Springer-Verlag, New YorkGoogle Scholar
  22. Cammeraat LH, Willott SJ, Compton SG, Incoll LD (2002) The effects of ants’ nests on the physical, chemical and hydrological properties of a rangeland soil in semi-arid Spain. Geoderma 105:1–20CrossRefGoogle Scholar
  23. Cerdá X, Retana J, Manzaneda A (1998) The role of competition by dominants and temperature in the foraging of subordinate species in Mediterranean ant communities. Oecologia 117:404–412CrossRefPubMedGoogle Scholar
  24. Clusella-Trullas S, Terblanche JS, Chown SL (2010) Phenotypic plasticity of locomotion performance in the seed harvester Messor capensis (Formicidae). Physiol Biochem Zool 83:519–530CrossRefPubMedGoogle Scholar
  25. Collignon B, Detrain C (2014) Accuracy of leadership and control of the audience in the pavement ant Tetramorium caespitum. Anim Behav 92:159–165CrossRefGoogle Scholar
  26. Crist TO, MacMahon JA (1991) Individual foraging components of harvester ants: movement patterns and seed patch fidelity. Insect Soc 38:379–396CrossRefGoogle Scholar
  27. Davidson DW (1978) Size variability in the worker caste of a social insect (Veromessor pergandei Mayr) as a function of the competitive environment. Am Nat 112:523–532CrossRefGoogle Scholar
  28. Detrain C, Tasse O, Versaen M, Pasteels JM (2000) A field assessment of optimal foraging in ants: trail patterns and seed retrieval by the European harvester ant Messor barbarus. Insect Soc 47:56–62CrossRefGoogle Scholar
  29. Edelstein-Keshet L, Watmough J, Ermentrout GB (1995) Trail following in ants: individual properties determine population behaviour. Behav Ecol Sociobiol 36:119–133CrossRefGoogle Scholar
  30. Espadaler X, Gómez C (2001) Formicine ants comply with the size-grain hypothesis. Funct Ecol 15:136–138CrossRefGoogle Scholar
  31. Farji-Brener AG, Barrantes G, Ruggiero A (2004) Environmental rugosity, body size and access to food: a test of the size-grain hypothesis in tropical litter ants. Oikos 104:165–171CrossRefGoogle Scholar
  32. Fourcassié V, Bredard C, Volpatti K, Theraulaz G (2003) Dispersion movements in ants: spatial structuring and density-dependent effects. Behav Process 63:33–43CrossRefGoogle Scholar
  33. Franks NR, Richardson T (2006) Teaching in tandem-running ants. Nature 439:153–153CrossRefPubMedGoogle Scholar
  34. Frizzi F, Bartalesi V, Santini G (2017) Combined effects of temperature and interspecific competition on the mortality of the invasive garden ant, Lasius neglectus: A laboratory study. J Therm Biol 65:76–81CrossRefPubMedGoogle Scholar
  35. Frizzi F, Ciofi C, Dapporto L, Natali C, Chelazzi G, Turillazzi S, Santini G (2015) The rules of aggression: how genetic, chemical and spatial factors affect intercolony fights in a dominant species, the mediterranean acrobat ant Crematogaster scutellaris. PLoS One 10:e0137919CrossRefPubMedPubMedCentralGoogle Scholar
  36. Fukushi T, Wehner R (2004) Navigation in wood ants Formica japonica: context dependent use of landmarks. J Exp Biol 207:3431–3439CrossRefPubMedGoogle Scholar
  37. Gillooly JF, Brown JH, West GB, Savage VM, Charnov EL (2001) Effects of size and temperature on metabolic rate. Science 293:2248–2251CrossRefPubMedGoogle Scholar
  38. Gordon DM (2002) The regulation of foraging activity in red harvester ant colonies. Am Nat 159:509–518CrossRefPubMedGoogle Scholar
  39. Gordon DM (1995) The expandable network of ant exploration. Anim Behav 50-995-1007Google Scholar
  40. Gordon DM (1989) Dynamics of task switching in harvester ants. Anim Behav 38-194-204Google Scholar
  41. Grasso DA, Sledge MF, Le Moli F, Mori A, Turillazzi S (2005) Nest-area marking with faeces: a chemical signature that allows colony-level recognition in seed harvesting ants (Hymenoptera, Formicidae). Insect Soc 52:36–44CrossRefGoogle Scholar
  42. Grime JP (2001) Plant strategies, vegetation processes, and ecosystem properties. Wiley, ChichesterGoogle Scholar
  43. Haefner JW, Crist TO (1994) Spatial model of movement and foraging in harvester ants (Pogonomyrmex)(I): The roles of memory and communication. J Theor Biol 166:299–313CrossRefGoogle Scholar
  44. Hahn M, Maschwitz U (1985) Foraging strategies and recruitment behaviour in the European harvester ant Messor rufitarsis (F.). Oecologia 68:45–51CrossRefPubMedGoogle Scholar
  45. Hailey A, Coulson IM (1996) Differential scaling of home-range area to daily movement distance in two African tortoises. Can J Zool 74:97–102CrossRefGoogle Scholar
  46. Heredia A, Detrain C (2005) Influence of seed size and seed nature on recruitment in the polymorphic harvester ant Messor barbarus. Behav Process 70:289–300CrossRefGoogle Scholar
  47. Hölldobler B (1976) Recruitment behavior, home range orientation and territoriality in harvester ants, Pogonomyrmex. Behav Ecol Sociobiol 1:3–44CrossRefGoogle Scholar
  48. Hölldobler B, Wilson EO (1990) The ants. Harvard University Press, CambridgeCrossRefGoogle Scholar
  49. Hothorn T, Bretz F, Westfall P (2008) Simultaneous Inference in General Parametric Models. Biometrical J 50:346–363CrossRefGoogle Scholar
  50. Hsu J (1996) Multiple comparisons: theory and methods. Chapman & Hall, LondonCrossRefGoogle Scholar
  51. Hurlbert AH, Ballantyne F, Powell S (2008) Shaking a leg and hot to trot: the effects of body size and temperature on running speed in ants. Ecological Entomology 33:144–154CrossRefGoogle Scholar
  52. Hurvich CM, Tsai CL (1991) Bias of the corrected AIC criterion for underfitted regression and time series models. Biometrika 78:499–509Google Scholar
  53. Johnson RA (1992) Soil texture as an influence on the distribution of the desert seed-harvester ants Pogonomyrmex rugosus and Messor pergandei. Oecologia 89:118–124CrossRefPubMedGoogle Scholar
  54. Kaspari M (1996) Worker size and seed size selection by harvester ants in a Neotropical forest. Oecologia 105:397–404CrossRefPubMedGoogle Scholar
  55. Kaspari M, Weiser MD (1999) The size–grain hypothesis and interspecific scaling in ants. Funct Ecol 13:530–538CrossRefGoogle Scholar
  56. Kaspari M, Weiser M (2007) The size–grain hypothesis: do macroarthropods see a fractal world? Ecological Entomology 32:279–282CrossRefGoogle Scholar
  57. Leonard JG, Herbers JM (1986) Foraging tempo in two woodland and species. Anim Behav 34:1172–1181CrossRefGoogle Scholar
  58. Lighton JRB, Bartholomew GA (1988) Standard energy metabolism of a desert harvester ant, Pogonomyrmex rugosus: effects of temperature, body mass, group size, and humidity. Proc Natl Acad Sci USA 85:4765–4769CrossRefPubMedPubMedCentralGoogle Scholar
  59. Longino JT, Nadkarni NM (1990) A comparison of ground and canopy leaf litter ants (Hymenoptera: Formicidae) in a neotropical montane forest. Psyche 97:81–93CrossRefGoogle Scholar
  60. Loretto D, Vieira MV (2005) The effects of reproductive and climatic seasons on movements in the black-eared opossum (Didelphis aurita Wied-Neuwied, 1826). J Mammal 86:287–293CrossRefGoogle Scholar
  61. Mandelbrot B (1984) Les objets fractals: Forme, hasard et dimension. Flammarion, ParisGoogle Scholar
  62. Marsh RL (1990) Deactivation rate and shortening velocity as determinants of contractile frequency. Am J Physiol 259:223–230Google Scholar
  63. Mazerolle MJ (2016) AICcmodavg: Model selection and multimodel inference based on (Q)AIC(c). Accessed 3 August 2017
  64. Morales JM, Ellner SP (2002) Scaling up animal movements in heterogeneous landscapes: the importance of behavior. Ecology 83:2240–2247CrossRefGoogle Scholar
  65. Morehead SA, Feener DH Jr (1998) Foraging behavior and morphology: seed selection in the harvester ant genus. Pogonomyrmex Oecologia 114:548–555CrossRefPubMedGoogle Scholar
  66. Nams VO (2006) Detecting oriented movement of animals. Anim Behav 72:1197–1203CrossRefGoogle Scholar
  67. Nams VO (1996) The VFractal: a new estimator for fractal dimension of animal movement paths. Landscape Ecol 11:289–297CrossRefGoogle Scholar
  68. Pearce-Duvet JMC, Elemans CPH, Feener DH (2011a) Walking the line: search behavior and foraging success in ant species. Behav Ecol 22:501–509CrossRefGoogle Scholar
  69. Pearce-Duvet JMC, Moyano M, Adler FR, Feener DH (2011b) Fast food in ant communities: how competing species find resources. Oecologia 167:229–240CrossRefPubMedGoogle Scholar
  70. Petit O, Bon R (2010) Decision-making processes: the case of collective movements. Behav Process 84:635–647CrossRefGoogle Scholar
  71. Pinter-Wollman N, Gordon DM, Holmes S (2012) Nest site and weather affect the personality of harvester ant colonies. Behav Ecol 23:1022–1029CrossRefPubMedPubMedCentralGoogle Scholar
  72. Powell S, Franks NR (2006) Ecology and the evolution of worker morphological diversity: a comparative analysis with Eciton army ants. Funct Ecol 20:1105–1114CrossRefGoogle Scholar
  73. Prevedello JA, Forero-Medina G, Vieira MV (2010) Movement behaviour within and beyond perceptual ranges in three small mammals: effects of matrix type and body mass. J Anim Ecol 79:1315–1323CrossRefPubMedGoogle Scholar
  74. Prothero J (1992) Scaling of bodily proportions in adult terrestrial mammals. Am J Physiol 262:492–503Google Scholar
  75. Reynolds AM, Rhodes CJ (2009) The Lévy flight paradigm: random search patterns and mechanisms. Ecology 90:877–887CrossRefPubMedGoogle Scholar
  76. Rissing SW (1982) Foraging velocity of seed-harvester ants, Veromessor pergandei (Hymenoptera: Formicidae). Environ Entomol 11:905–907CrossRefGoogle Scholar
  77. Rissing SW, Pollock GB (1984) Worker size variability and foraging efficiency in Veromessor pergandei (Hymenoptera: Formicidae). Behav Ecol Sociobiol 15:121–126CrossRefGoogle Scholar
  78. Robinson GE (1992) Regulation of division of labor in insect societies. Annu Rev Entomol 37:637–665CrossRefPubMedGoogle Scholar
  79. Rome LC, Lindstedt SL (1998) The Quest for Speed: Muscles Built for High-Frequency Contractions. News Physiol Sci 13:261–268PubMedGoogle Scholar
  80. Rosengren R, Fortelius W (1986) Ortstreue in foraging ants of the Formica rufa group - Hierarchy of orienting cues and long-term memory. Insect Soc 33-306-337Google Scholar
  81. Rowcliffe MJ, Carbone C, Kays R, Kranstauber B, Jansen PA (2012) Bias in estimating animal travel distance: the effect of sampling frequency. Methods Ecology Evol 3:653–662CrossRefGoogle Scholar
  82. Santini G, Tucci L, Ottonetti L, Frizzi F (2007) Competition trade-offs in the organisation of a Mediterranean ant assemblage. Ecol Entomol 32:319–326CrossRefGoogle Scholar
  83. Sarty M, Abbott KL, Lester PJ (2006) Habitat complexity facilitates coexistence in a tropical ant community. Oecologia 149:465–473CrossRefPubMedGoogle Scholar
  84. Schmidt-Nielsen K (1984) Scaling: why is animal size so important? Cambridge University Press, CambridgeCrossRefGoogle Scholar
  85. Schoener TW (1981) An empirically based estimate of home range. Theor Popul Biol 20:281–325CrossRefGoogle Scholar
  86. Schwander T, Rosset H, Chapuisat M (2005) Division of labour and worker size polymorphism in ant colonies: the impact of social and genetic factors. Behav Ecol Sociobiol 59:215–221CrossRefGoogle Scholar
  87. Solida L, Grasso DA, Celant A, Fanfani A, Mori A, Le Moli F (2007) Foraging activity in two species of Messor harvester ants: preliminary data on size-matching and diet breadth. Redia 90:71–73Google Scholar
  88. Swihart RK, Slade NA (1985) Testing for independence of observations in animal movements. Ecology 66:1176–1184CrossRefGoogle Scholar
  89. Swingland IR, Greenwood PJ (1983) The Ecology of Animal Movement. Oxford University Press, OxfordGoogle Scholar
  90. Torres-Contreras H, Canals M (2010) Effect of colony, patch distance, and trajectory sense on movement complexity in foraging ants. J Insect Behav 23:319–328CrossRefGoogle Scholar
  91. Traniello JF (1989) Foraging strategies of ants. Annu Rev Entomol 34:191–210CrossRefGoogle Scholar
  92. Walter H (1971) Ecology of tropical and subtropical vegetation. Oliver & Boyd, EdinburghGoogle Scholar
  93. Viswanathan GM, Buldyrev SV, Havlin S, Da Luz MGE, Raposo EP, Stanley HE (1999) Optimizing the success of random searches. Nature 401:911–914CrossRefPubMedGoogle Scholar
  94. Vogt JT, Appel AG (1999) Standard metabolic rate of the fire ant, Solenopsis invicta Buren: effects of temperature, mass, and caste. J Insect Physiol 45:655–666CrossRefPubMedGoogle Scholar
  95. Wehner R, Srinivasan MV (1981) Searching behaviour of desert ants, genus Cataglyphis (Formicidae, Hymenoptera). J Comp Physiol 142:315–338CrossRefGoogle Scholar
  96. Weier JA, Feener DH (1995) Foraging in the seed-harvester ant genus Pogonomyrmex: are energy costs important? Behav Ecol Sociobiol 36:291–300CrossRefGoogle Scholar
  97. Weiser MD, Kaspari M (2006) Ecological morphospace of New World ants. Ecol Entomol 31:131–142CrossRefGoogle Scholar
  98. Wiens JA, Crist TO, With KA, Milne BT (1995) Fractal patterns of insect movement in microlandscape mosaics. Ecology 76:663–666CrossRefGoogle Scholar
  99. Wiescher PT, Pearce-Duvet JM, Feener DH (2012) Assembling an ant community: species functional traits reflect environmental filtering. Oecologia 169:1063–1074CrossRefPubMedGoogle Scholar
  100. Willott SJ, Compton SG, Incoll LD (2000) Foraging, food selection and worker size in the seed harvesting ant Messor bouvieri. Oecologia 125:35–44CrossRefPubMedGoogle Scholar
  101. Wilson EO (1978) Division of labor in fire ants based on physical castes (Hymenoptera: Formicidae: Solenopsis). J Kansas Entomol Soc 51:615–636Google Scholar
  102. Wittlinger M, Wehner R, Wolf H (2006) The ant odometer: stepping on stilts and stumps. Science 312:1965–1967CrossRefPubMedGoogle Scholar
  103. Zollikofer C (1994) Stepping patterns in ants-influence of speed and curvature. J Exp Biol 192:95–106PubMedGoogle Scholar

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© International Union for the Study of Social Insects (IUSSI) 2018

Authors and Affiliations

  1. 1.Department of BiologyUniversity of FlorenceSesto FiorentinoItaly

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