Journal of Insect Behavior

, Volume 32, Issue 2, pp 164–179 | Cite as

Factors Influencing Sun Basking in Red Wood Ants (Formica polyctena): a Field Experiment on Clustering and Phototaxis

  • Š. Kadochová
  • J. FrouzEmail author
  • A. Tószögyová


We monitored nest interior and surface temperatures together with two aspects of the sun-basking behavior of wood ants: aggregation (by regular monitoring of occurrence and density of clusters) and phototaxis (tendency to move from shade to sun or vice versa, recorded as a response to artificial shading during sunny weather), using ten nests of Formica polyctena near Vimperk Czechia. Dense clusters occurred only in spring; weak clusters rarely occur the rest of the year. Statistical analysis showed that timing of dense clusters is affected by both environmental parameters (light intensity, insulation status of the nest) and internal nest factors (inner nest temperature, ant population size, nest volume). Phototaxis changed from positive to negative during the year. In spring, ant workers performed the sun basking readily and when shaded moved into the sun. In summer, however, the ants avoided sun basking and aggregated in the shade. The shift from sun basking to sun avoidance was driven mainly by nest surface temperature. The switch occurred when the temperature of sun-exposed nest surface reached 42.8 °C, which is two degrees higher than the experimentally measured lethal temperature (LD50) for red wood ants. This shows that two basic components of sun basking, aggregation and sun exposure, are driven by a different set of environmental conditions and their interplay is likely to maintain balance between the needs of the colony to heat up the nest and risk for individual workers arising from overheating during prolonged sun exposure.


Red wood ants Thermoregulation Sun basking Field experiment Sunning clusters 



We thank our colleagues, namely Veronika Jílková, for help and support in our research. Our thanks also belong to Mrs. Somková for proof reading and Mr. Jonathan Riches for language correction. We also thank two anonymous reviewers and the editors of the Journal of Insect Behavior, Drs. Jeremy Allison and Ring Cardé for valuable comments and suggestions. Programs for research support at Charles University, namely the programs PRVOUK P02 and Progress P41 are thanked for financial support as well as ministry of education, youth and sports LM2015075 and EF16_013/0001782.


This study was funded by Program for research support at Charles University PRVOUK (programs P02 and P41) and MEYS grants number LM2015075 and EF16_013/0001782.

Compliance with Ethical Standards

Conflict of Interest

The authors declare that they have no conflict of interest.

Supplementary material

10905_2019_9713_Fig7_ESM.png (4.3 mb)

Scheme of the shading experiment and evaluation of nest surface temperature from IR photos. a + b = a sunning cluster before the shading experiment, a – digital image of sun basking ants, b – IR image of sun basking ants with the temperature scale indicated; c + d = situation on the nest surface at the end of the shading experiment, c – digital image of the shading experiment, d – IR image of the shading experiment; note that the shaded area is colder. The area marked by rectangle in both IR images is the area where the temperature of sun-exposed nest surface (b) and shaded nest surface (d) was measured by FLIR tools. Maximal, minimal and average temperature for the evaluated rectangles is shown in the IR picture (PNG 4362 kb)

10905_2019_9713_MOESM1_ESM.tif (5.7 mb)
High resolution image (TIF 5786 kb)
10905_2019_9713_Fig8_ESM.png (695 kb)

A detailed look on ant workers on nest surface, obtained with an infrared camera. Temperatures are depicted in false colors, the darker is the color the lower is the temperature. Note that two ant individuals are marked as Sp1 and Sp2, their body temperature is depicted in upper left corner. The cold (dark purple) ants are those, who emerged from nest interior or from shaded areas, four visible hot ants (yellow-white) came from sunning cluster. (PNG 694 kb)

10905_2019_9713_MOESM2_ESM.tif (988 kb)
High resolution image (TIF 988 kb)


  1. Anderson KE, Munger JC (2003) Effect of temperature on brood relocation in Pogonomyrmex salinus (Hymenoptera: Formicidae). West N Am Nat 63:122–128Google Scholar
  2. Bollazzi M, Roces F (2002) Thermal preference for fungus culturing and brood location by workers of the thatching grass-cutting ant Acromyrmex heyeri. Insect Soc 49:153–157CrossRefGoogle Scholar
  3. Brandt CJ (1980) The thermal diffusivity of the organic material of a mound of Formica polyctena in relation to the thermoregulation of the brood (Hymenoptera, Formicidae). Netherlands Journal of Zoology 30:326–344CrossRefGoogle Scholar
  4. Brian MV (1973) Temperature choice and its relevance to brood survival and caste determination in the ant Myrmica rubra L. Physiol Zool 46:245–252CrossRefGoogle Scholar
  5. Calabi P, Porter SD (1989) Worker longevity in the fire ant, Solenopsis invicta, ergonomic considerations of correlations between temperature size and metabolic rates. J Insect Physiol 35:643–650CrossRefGoogle Scholar
  6. Challet M, Jost C, Grimall A et al (2005) How temperature influences displacements and corpse aggregation behaviors in the ant Messor sancta. Insect Soc 52:309–315CrossRefGoogle Scholar
  7. Cherix D, Freitag A, Maeder A (2006) Fourmis des Bois. Lausanne, Switzerland: Musee de ZoologieGoogle Scholar
  8. Coenen-Stass D, Schaarschmidt B, Lamprecht I (1980) Temperature distribution and calorimetric determination of heat production in the nest of the wood ants Formica polyctena (Hymenoptera Formicidae). Ecology 61:238–244CrossRefGoogle Scholar
  9. Crawley MJ (2007) The R book. John Wiley & Sons Ltd, ChichesterCrossRefGoogle Scholar
  10. Dlusskij GM (1967) Muravji roda Formica. Nauka, MoskvaGoogle Scholar
  11. Domish T, Finér L, Neuvonen S, Niemelä P, Risch AC, Kilpeläinen J, Ohashi M, Jurgensen MF (2009) Foraging aktivity and dietary spektrum of wood ants (Formica rufa group) and their role in nutritient fluxes in boreal forests. Ecological Entomology 34:369–377CrossRefGoogle Scholar
  12. Ellis S, Robinson EJH (2014) Polydomy in red wood ants. Insect Soc 61:111–122CrossRefGoogle Scholar
  13. Frouz J (1996) The role of nest moisture in thermoregulation of ant (Formica polyctena, Hymenoptera, Formicidae) nests. Biologia 51:541–547Google Scholar
  14. Frouz J (2000) The effect of nest moisture on daily temperature regime in the nests of Formica polyctena wood ants. Insect Soc 47:229–235CrossRefGoogle Scholar
  15. Frouz J, Finer L (2007) Diurnal and seasonal fluctuations in wood ant (Formica polyctena) nest temperature in two geographically distant populations along a south-north gradient. Insect Soc 54:251–259CrossRefGoogle Scholar
  16. Frouz J, Rybníček M, Cudlín P, Chmelíková E (2008) Influence of the wood ant, Formica polyctena, on soil nutrient and the spruce tree growth. J Appl Entomol 132:281–284CrossRefGoogle Scholar
  17. Hamilton WD (1964) The genetical evolution of social behaviour. J. Theor. Biol 7:1–16CrossRefGoogle Scholar
  18. Heinrich B (1993) The hot-blooded insects. Strategies and mechanisms of thermoregulation. BerlinGoogle Scholar
  19. Higgins RJ, Lindgren BS (2012) The effect of manipulated shading on the colony abundance of two species of ants, Formica aserva and Leptothorax muscorum, in dead wood. Entomologia Experimentalis et Applicata 143:292–300CrossRefGoogle Scholar
  20. Hölldobler B, Wilson EO (1990) The ants. Belknap Press, Springer, BerlinCrossRefGoogle Scholar
  21. Horstmann K, Schmid H (1986) Temperature regulation in nests of the wood ant, Formica polyctena (Hymenoptera: Formicidae). Entomologia Generalis 11:229–236CrossRefGoogle Scholar
  22. Jílková L, Matějíček J, Frouz J (2011) Changes in the pH and other soil chemical parameters in soil surrounding wood ant (Formica polyctena) nests. Eur J Soil Biol 47:72–76CrossRefGoogle Scholar
  23. Jílková V, Cajthaml T, Frouz J (2015a) Respiration in wood ant (Formica aquilonia) nests as affected by altitudinal and seasonal changes in temperature. Soil Biol Biochem 86:50–57CrossRefGoogle Scholar
  24. Jílková V, Frouz J, Mudrák O, Vohník M (2015b) Effects of nutrient-rich substrate and ectomycorrhizal symbiosis on spruce seedling biomass in abandoned nests of the wood ant (Formica polyctena): a laboratory experiment. Geoderma 259:56–61CrossRefGoogle Scholar
  25. Jones JC, Oldroyd BP (2006) Nest thermoregulation in social insects. Advances in Insect Physiology 33:153–191CrossRefGoogle Scholar
  26. Jongepier E, Foitzik S (2016) Fitness costs of worker specialization for ant societies. Proc R Soc B 283:20152572CrossRefGoogle Scholar
  27. Kadochová Š, Frouz J (2013) Thermoregulation strategies in ants in comparison to other social insects, with a focus on Formica rufa. F1000Research 2:280. CrossRefGoogle Scholar
  28. Kadochová Š, Frouz J (2014) Red wood ants Formica polyctena switch off active thermoregulation of the nest in autumn. Insect Soc 61:297–306CrossRefGoogle Scholar
  29. Kadochová Š, Frouz J, Roces F (2017) Sun basking in red wood ants Formica polyctena (Hymenoptera, Formicidae): individual behaviour and temperature-dependent respiration rates. PLoS One.
  30. Kasimova RG, Tishin D, Obnosov YV, Dlussky GM, Baksht FB, Kacimov AR (2014) Ant mound as an optimal shape in constructal design: solar irradiation and circadian brood/fungi-warming sorties. J Theor Biol 355:21–32CrossRefGoogle Scholar
  31. Kilpeläinen J, Punttila P, Finér L, Niemelä P, Domisch T, Jurgensen MF, Neuvonen S, Ohashi M, Risch AC, Sundström L (2008) Distribution of ant species and mounds (Formica) in different-aged managed spruce stands in eastern Finland. J Appl Entomol 132:315–325CrossRefGoogle Scholar
  32. Kilpeläinen J, Finer L, Neuvonen S, Niemelä P, Domisch T, Risch AC, Jurgensen MF, Ohashi M, Sundström L (2009) Does the mutualism between wood ants (Formica rufa group) and Cinara aphids affect Norway spruce growth? For Ecol Manag 257:238–243CrossRefGoogle Scholar
  33. Kipyatkov VE, Schederova SS (1985) Seasonal changes in behavior patterns of the ant Formica polyctena in atrificial nest with temperature gradient. Zoologicheskii zhurnal 65:1847–1857Google Scholar
  34. Kipyatkov VE, Schederova SS (1990) The endogenous rhythm of queens reproductivity in red wood ants (Formica group). Zoologicheskii zhurnal 69:40–52Google Scholar
  35. Kleinhenz M, Bujok B, Fuchs S, Tautz J (2003) Hot bees in empty broodnest cells: heating from within. J Exp Biol 206:4217–4231CrossRefGoogle Scholar
  36. Kneitz G (1964) Untersuchungen zum Aufbau und zur Erhaltung des Nestwärmehaushaltes bei Formica polyctena Foerst (Hym, Formicidae). Dissertation, University of Würzburg, GermanyGoogle Scholar
  37. Korb J, Linsenmair KE (1998) The effect of temperature on the architecture and distribution of Macrotermes bellicosus (Isoptera, Macrotermitinae) mounds in different habitats of west African Guinea savanna. Insect Soc 45:51–65CrossRefGoogle Scholar
  38. Lamprecht I, Maierhofer C, Röllig M (2007) Infrared thermography and thermometry of phototropic plants. J Therm Anal Calorim 87:49–54CrossRefGoogle Scholar
  39. Lenoir L, Persson T, Bengtsson J (2001) Wood ant nests as potential hot spots for carbon and nitrogen mineralisation. Biol Fert Soils 34:235–240CrossRefGoogle Scholar
  40. Lindauer M (1954) Temperaturregulierung und Wasserhaushalt im Bienenstaat. Zeitschrift für vergleichende Physiologie 36:391–432CrossRefGoogle Scholar
  41. McGlynn TP, Dunn T, Wayman E, Romero A (2010) A thermophile in the shade: light-directed nest relocation in the Costa Rican ant Ecatomma ruidum. J Trop Ecol 26:559–562CrossRefGoogle Scholar
  42. Neven LG (2000) Physiological responses of insects to heat. Postharvest Biol Technol 21:103–111CrossRefGoogle Scholar
  43. Nowak MA, Tarnita CE, Wilson EO (2010) The evolution of eusociality. Nature 466:1057–1062CrossRefGoogle Scholar
  44. Pamilo P, Rosengren R (1983) Sex ratio strategies in Formica ants. Oikos 40:24–35CrossRefGoogle Scholar
  45. Penick CA, Tschinkel WR (2008) Thermoregulatory brood transport in the fire ant, Solenopsis invicta. Insect Soc 55:176–182CrossRefGoogle Scholar
  46. Porter SD (1988) Impact of temperature on colony growth and developmental rates of the ant Solenopsis invicta. J Insect Physiol 34:1127–1133CrossRefGoogle Scholar
  47. Porter SD, Tschinkel WR (1987) Foraging in Solenopsis invicta (Hymenoptera: Formicidae): effects of weather and season. Environ Entomol 16:802–808CrossRefGoogle Scholar
  48. Prange HD (1996) Evaporative cooling in insects. J Insect Physiol 42:493–499CrossRefGoogle Scholar
  49. Punttila P, Niemela P, Karhu K (2004) The impact of wood ants (Hymenotpera: Formicidae) on the structure of invertebrate community on mountain birch (Betula pubescens). Annales Zoological Fennici 41:429–446Google Scholar
  50. R Core Team (2014). R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria.
  51. Roces F, Núñez JA (1989) Brood translocation and circadian variation of temperature preference in the ant Campotonus mus. Oecologia 81:33–37CrossRefGoogle Scholar
  52. Rosengren R, Fortelius W, Lindström K, Luther A (1987) Phenology and causation of nest heating and thermoregulation in red wood ants of the Formica rufa group studied in coniferous forest habitats in southern Finland. Ann Zool Fenn 24:147–155Google Scholar
  53. Seeley TD (2009) The wisdom of the hive: the social physiology of honey bee colonies, vol 295. Cambridge University Press, MassatchussetsGoogle Scholar
  54. Stockan JA, Robinson EJH (2016) Wood ant ecology and conservation. Cambridge University Press, CambridgeCrossRefGoogle Scholar
  55. Wilson EO (1971) The insect societies. Belknap Press of Harvard University Press, MassachusettsGoogle Scholar
  56. Zahn M (1958) Temperatursinn, Wärmehaushat und Bauweise der role Waldameisen (Formica rufa L). Zoologische Beiträge 3:127–194Google Scholar

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Authors and Affiliations

  1. 1.Department of Ecology, Faculty of ScienceCharles UniversityPraha 2Czech Republic
  2. 2.Institute for Environmental Studies, Faculty of ScienceCharles UniversityPraha 2Czech Republic
  3. 3.Environment CentreCharles UniversityPraha 6Czech Republic

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