Skip to main content
Log in

Exclusive core areas and intrasexual territoriality in Eurasian red squirrels (Sciurus vulgaris) revealed by incremental cluster polygon analysis

  • Original Article
  • Published:
Ecological Research

Abstract

When animal home ranges overlap extensively in species lacking overt territorial behaviours, identifying exclusive core areas within individual ranges can be difficult. By analysing the size and overlap of successively smaller core areas among individual Eurasian red squirrels (Sciurus vulgaris), we determined exclusive areas within the home ranges of resident males and females. Possible effects of habitat composition and food supplies were explored by monitoring squirrels in different conifer forests and during years with low and high tree seed production. Using outlier-exclusive cores (OEC) revealed that the total ranges consisted of large sally zones (on average, 35% of the total minimum convex polygon [MCP] range) around home ranges with multi-nucleate cores. The mean OEC home range size did not differ between the sexes but was larger with poor food availability. Home ranges (99% incremental cluster polygons [ICP]) overlapped extensively between sexes (average overlap high food–low food: males by females 21–40%, females by males 43–45%) and among males (males by males 26–44%), while intrasexual overlap among females was low (9–10%). The overlap of inner cores among females rapidly approached zero, suggesting the intrasexual territoriality of 75% core areas. This was not the case among male squirrels, for which intrasexual overlap averaged only 4% at 50% but 18% at 75% core areas. Even the smallest inner cores had some degree of intersexual overlap, indicating that complete territoriality did not occur in this species. Female home ranges were more strongly affected by annual fluctuations in food supplies than male ranges. Females reduced the size of their food-based intrasexual territories when food availability increases. Males probably benefit from using larger home ranges and core areas, which overlap with the ranges of several females, by increasing their probability of successful mating.

This is a preview of subscription content, log in via an institution to check access.

Access this article

Price excludes VAT (USA)
Tax calculation will be finalised during checkout.

Instant access to the full article PDF.

Fig. 1
Fig. 2
Fig. 3

Similar content being viewed by others

References

  • Andrén H, Delin A (1994) Habitat selection in the Eurasian red squirrel, Sciurus vulgaris, in relation to forest fragmentation. Oikos 70:43–48

    Article  Google Scholar 

  • Andrén H, Lemnell P-A (1992) Population fluctuations and habitat selection in the Eurasian red squirrel Sciurus vulgaris. Ecography 15:303–307

    Article  Google Scholar 

  • Aragón P, López P, Martin J (2001) Chemosensory discrimination of familiar and unfamiliar conspecifics by lizards: implications of field spatial relationships between males. Behav Ecol Sociobiol 50:128–133

    Article  Google Scholar 

  • Bakker VJ (2006) Microhabitat features influence the movements of red squirrels (Tamiasciurus hudsonicus) on unfamiliar ground. J Mammal 87:124–130

    Article  Google Scholar 

  • Bakker VJ, Van Vuren DH (2004) Gap-crossing decisions by the red squirrel (Tamiasciurus hudsonicus), a forest-dependent small mammal. Conserv Biol 18:689–697

    Article  Google Scholar 

  • Bertolino S, Wauters LA, De Bruyn L, Canestri-Trotti G (2003) Prevalence of coccidia parasites (Protozoa) in red squirrels (Sciurus vulgaris): effects of host phenotype and environmental factors. Oecologia 137:286–295

    Article  PubMed  Google Scholar 

  • Brown JL, Orians GH (1970) Spacing patterns in mobile animals. Annu Rev Ecol Syst 1:239–262

    Article  Google Scholar 

  • Bryce J, Johnson PJ, MacDonald DW (2002) Can niche use in red and grey squirrels offer clues for their apparent coexistence? J Appl Ecol 39:875–887

    Article  Google Scholar 

  • Burt WH (1943) Territoriality and home range concepts as applied to mammals. J Mammal 24:346–352

    Article  Google Scholar 

  • Carpenter FL, MacMillen RE (1976) Threshold model of feeding territoriality and test with a Hawaiian honeycreeper. Science 194:639–642

    Article  PubMed  CAS  Google Scholar 

  • Clutton-Brock TH (1989) Mammalian mating systems. Proc R Soc Lond Series B 236:339–372

    CAS  Google Scholar 

  • de Solla SR, Bonduriansky R, Brooks RJ (1999) Eliminating autocorrelation reduces biological relevance of home range estimates. J Anim Ecol 68:221–234

    Article  Google Scholar 

  • Getz LL, McGuire B (1993) A comparison of living singly and in male–female pairs in the prairie vole, Microtus ochrogaster. Ethology 94:265–278

    Article  Google Scholar 

  • Gurnell J (1987) The natural history of squirrels. Christopher Helm, London, UK

    Google Scholar 

  • Hagen CA, Kenkel NC, Walker DJ, Baydack RK, Braun CE (2001) Fractal-based spatial analysis of radiotelemetry data. In: Millspaugh JJ, Marzluff JM (eds) Radio tracking and animal populations. Academic Press, San Diego, California, pp 167–187

    Chapter  Google Scholar 

  • Hemson G, Johnson P, South A, Kenward RE, Ripley R, MacDonald D (2005) Are kernels the mustard? Data from global positioning system (GPS) collars suggests problems for kernel home-range analyses with least-squares cross-validation. J Anim Ecol 74:455–463

    Article  Google Scholar 

  • Ims RA (1988) Spatial clumping of sexually receptive females induces space sharing among male voles. Nature 335:541–543

    Article  PubMed  CAS  Google Scholar 

  • Jedrzejewski W, Jedrzejewska B, Szymura L (1995) Weasel population response, home range, and predation on rodents in a deciduous forest in Poland. Ecology 76:179–195

    Article  Google Scholar 

  • Kaufmann JH (1983) On the definitions and functions of dominance and territoriality. Biol Rev 58:1–20

    Article  Google Scholar 

  • Kenward RE (2001) A manual for wildlife radio tagging. Academic Press, Sydney, Australia

    Google Scholar 

  • Kenward RE, Clarke RT, Hodder KH, Walls SS (2001) Density and linkage estimators of home range: nearest-neighbor clustering defines multinuclear cores. Ecology 82:1905–1920

    Google Scholar 

  • Kenward RE, South A, Walls S (2003) Ranges6 v1.2: for the analysis of tracking and location data. Anatrack Ltd., Wareham, UK

    Google Scholar 

  • Kernohan BJ, Gitzen RA, Millspaugh JJ (2001) Analysis of animal space use and movements. In: Millspaugh JJ, Marzluff JM (eds) Radio tracking and animal populations. Academic Press, San Diego, California, pp 125–166

    Chapter  Google Scholar 

  • Kerr GD, Bull CM (2006) Exclusive core areas in overlapping ranges of the sleepy lizard, Tiliqua rugosa. Behav Ecol 17:380–391

    Article  Google Scholar 

  • Koprowski JL (1998) Conflict between the sexes: a review of social and mating systems of the tree squirrels. In: Steele MA, Merritt JF, Zegers DA (eds) Ecology and evolutionary biology of tree squirrels. Virginia Museum of Natural History, Special Publication no 6, pp 33–41

  • Lair H (1987) Estimating the location of the focal center in red squirrel home ranges. Ecology 68:1092–1101

    Article  Google Scholar 

  • Lee TH (2001) Mating behavior of the Eurasian red squirrel (Sciurus vulgaris Linnaeus, 1758) in Hokkaido, Japan. Mammalia 65:131–142

    Google Scholar 

  • Lomnicki A (1980) Regulation of population density due to individual differences and patchy environment. Oikos 35:185–193

    Article  Google Scholar 

  • Lurz PWW, Garson PJ, Rushton SP (1995) The ecology of squirrels in spruce dominated plantations: implications for forest management. For Ecol Manage 79:79–90

    Article  Google Scholar 

  • Lurz PWW, Garson PJ, Wauters LA (1997) Effects of temporal and spatial variation in habitat quality on red squirrel dispersal behaviour. Anim Behav 54:427–435

    Article  PubMed  Google Scholar 

  • Lurz PWW, Garson PJ, Wauters LA (2000) Effects of temporal and spatial variations in food supply on the space and habitat use of red squirrels, Sciurus vulgaris L. J Zool Lond 251:167–178

    Article  Google Scholar 

  • Mahady SJ, Wolff JO (2002) A field test of the Bruce effect in the monogamous prairie vole (Microtus ochrogaster). Behav Ecol Sociobiol 52:31–37

    Article  Google Scholar 

  • Maher CR, Lott DF (1995) Definitions of territoriality used in the study of variation in vertebrate spacing systems. Anim Behav 49:1581–1597

    Article  Google Scholar 

  • Maher CR, Lott DF (2000) A review of ecological determinants of territoriality within vertebrate species. Am Midl Nat 143:1–29

    Article  Google Scholar 

  • Mares MA, Lacher TE Jr (1987) Social spacing in small mammals: patterns of individual variation. Am Zool 27:293–306

    Google Scholar 

  • McLoughlin PD, Ferguson SH, Messier F (2000) Intraspecific variation in home range overlap with habitat quality: a comparison among brown bear populations. Evol Ecol 14:39–60

    Article  Google Scholar 

  • Molinari A, Wauters LA, Airoldi G, Cerinotti F, Martinoli A, Tosi G (2006) Cone selection by Eurasian red squirrels in mixed conifer forests in the Italian Alps. Acta Oecologica 30:1–10

    Article  Google Scholar 

  • Moorcroft PR, Lewis MA, Crabtree RL (1999) Home range analysis using a mechanistic home range model. Ecology 80:1656–1665

    Article  Google Scholar 

  • Münch S (1998) Populationsökologie, Aktivität, Raum- und Habitatnutzung beim Europäischen Eichhörnchen (Sciurus vulgaris L.) im Bergmischwald des Nationalparks Bayerischer Wald. PhD thesis, Friedrich-Schiller-Universität, Jena, Germany

  • Newton I (1998) Population limitation in birds. Academic Press, London, UK

    Google Scholar 

  • Ostfeld RS (1985) Limiting resources and territoriality in microtine rodents. Am Nat 126:1–15

    Article  Google Scholar 

  • Ostfeld RS (1990) The ecology of territoriality in small mammals. Trends Ecol Evol 5:411–415

    Article  Google Scholar 

  • Parker GA, Sutherland WJ (1986) Ideal free distributions when individuals differ in competitive ability: phenotype-limited ideal free models. Anim Behav 34:1222–1242

    Article  Google Scholar 

  • Samuel MD, Pierce DJ, Garton EO (1985) Identifying areas of concentrated use within the home range. J Anim Ecol 54:711–719

    Article  Google Scholar 

  • SAS (1999) SAS STAT user’s guide, version v.8.0. SAS Institute, Cary, North Carolina

  • Steele M, Wauters LA, Larsen KW (2005) Selection, predation and dispersal of seeds by tree squirrels in temperate and boreal forests: are tree squirrels keystone granivores? In: Forget J-M, Forget P-M, Lambert JE, Hulme PE, Vander Wall SB (eds) Seed fate: predation, dispersal, and seedling establishment. CAB International, Wallingford, UK, pp 205–221

    Google Scholar 

  • Steury TD, Murray DL (2003) Causes and consequences of individual variation in territory size in the American red squirrel. Oikos 101:147–156

    Article  Google Scholar 

  • Swihart RK, Slade NA (1985) Testing for independence of observations in animal movements. Ecology 66:1176–1184

    Article  Google Scholar 

  • Trizio I, Crestanello B, Galbusera P, Wauters LA, Tosi G, Matthysen E, Hauffe HC (2005) Geographical distance and physical barriers shape the genetic structure of Eurasian red squirrels (Sciurus vulgaris) in the Italian Alps. Mol Ecol 14:469–481

    Article  PubMed  CAS  Google Scholar 

  • Verbeke G, Molenberghs G (2000) Linear mixed models for longitudinal data. Springer series in statistics. Springer, New York

    Google Scholar 

  • Wauters LA, Dhondt AA, (1987) Activity budget and foraging behaviour of the red squirrel (Sciurus vulgaris Linnaeus, 1758) in a coniferous habitat. Z Säugetierkd 52:341–353

    Google Scholar 

  • Wauters LA, Dhondt AA (1989) Body weight, longevity and reproductive success in red squirrels (Sciurus vulgaris). J Anim Ecol 58:637–651

    Article  Google Scholar 

  • Wauters LA, Dhondt AA (1990) Nest-use by red squirrels (Sciurus vulgaris Linnaeus, 1758). Mammalia 54:377–389

    Article  Google Scholar 

  • Wauters LA, Dhondt AA (1992) Spacing behaviour of red squirrels, Sciurus vulgaris: variation between habitats and the sexes. Anim Behav 43:297–311

    Article  Google Scholar 

  • Wauters LA, Dhondt AA (1993) Immigration patterns and success in red squirrels. Behav Ecol Sociobiol 33:159–167

    Article  Google Scholar 

  • Wauters LA, Dhondt AA (1995) Lifetime reproductive success and its correlates in female Eurasian red squirrels. Oikos 72:402–410

    Article  Google Scholar 

  • Wauters LA, Dhondt AA (1998) Variation in spacing behavior of Eurasian red squirrels, Sciurus vulgaris: effects of density and food. In: Steele MA, Merritt JF, Zegers D (eds) Ecology and evolutionary biology of tree squirrels. Virginia Museum of Natural History, Special Publication no 6, pp 71–78

  • Wauters LA, Gurnell J (1999) The mechanism of replacement of red squirrels by grey squirrels: a test of the interference competition hypothesis. Ethology 105:1053–1071

    Article  Google Scholar 

  • Wauters LA, De Vos R, Dhondt AA (1990) Factors affecting male mating success in red squirrels (Sciurus vulgaris). Ethol Ecol Evol 2:195–204

    Article  Google Scholar 

  • Wauters LA, Swinnen C., Dhondt AA (1992) Activity budget and foraging behaviour of red squirrels (Sciurus vulgaris) in coniferous and deciduous habitats. J Zool 1:71–86

    Article  Google Scholar 

  • Wauters LA, Casale P, Dhondt AA (1994) Space use and dispersal of red squirrels in fragmented habitats. Oikos 69:140–146

    Article  Google Scholar 

  • Wauters LA, Lens L, Dhondt AA (1995) Variation in territory fidelity and territory shifts among red squirrel, Sciurus vulgaris, females. Anim Behav 49:187–193

    Article  Google Scholar 

  • Wauters LA, Lurz PWW, Gurnell J (2000) Interspecific effects of grey squirrels (Sciurus carolinensis) on the space use and population demography of red squirrels (Sciurus vulgaris) in conifer plantations. Ecol Res 15:271–284

    Article  Google Scholar 

  • Wauters LA, Gurnell J, Preatoni D, Tosi G (2001) Effects of spatial variation in food availability on spacing behaviour and demography of Eurasian red squirrels. Ecography 24:525–538

    Article  Google Scholar 

  • Wauters LA, Zaninetti M, Tosi G, Bertolino S (2004) Is coat-colour polymorphism in Eurasian red squirrels (Sciurus vulgaris L.) adaptive? Mammalia 68:37–48

    Article  Google Scholar 

  • Wauters LA, Bertolino S, Adamo M, Van Dongen S, Tosi G (2005) Food shortage disrupts social organization: the case of red squirrels in conifer forests. Evol Ecol 19:375–404

    Article  Google Scholar 

  • Wauters LA, Preatoni D, Molinari A, Tosi G (2007) Radio-tracking squirrels: performance of home range density and linkage estimators with small range and sample size. Ecol Modelling 202:333–344

    Article  Google Scholar 

  • White GC, Garrott RA (1990) Analysis of wildlife radio-tracking data. Academic Press, London, UK

    Google Scholar 

  • Zalewski A, Jedrzejewski W (2006) Spatial organisation and dynamics of the pine marten Martes martes population in Bialowieza Forest (E Poland) compared with other European woodlands. Ecography 29:31–43

    Article  Google Scholar 

  • Zscheile K (1995) Zur Ökologie des Europäischen Eichhörnchens Sciurus vulgaris L.,1758 und seiner Verbreitung in den östlichen Bundesländern Deutschlands. Diplom Arbeit, Martin-Luther Universität Halle-Wittenberg, Germany

Download references

Acknowledgements

We thank Peter Lurz for his help with the Ranges6 software. Constructive comments by the two anonymous reviewers helped to improve the manuscript. This study is part of the ASPER (Alpine Squirrel Population Ecology Research) project, funded by the province of Sondrio (Servizio Agricoltura, Caccia e Pesca, Settore Risorse Ambientali), the Stelvio National Park and the Parco Regionale delle Orobie Valtellinesi to Istituto Oikos NGO, Milan, Italy. Additional financial support was given by MIUR (Ministero dell’Istruzione, dell’Università della Ricerca, project COFIN 2003, number 2003053710-006) to the University of Insubria Varese, Italy. Further logistic support was provided by the province of Sondrio and the Stelvio National Park. The trapping and handling of squirrels complied with the current laws on animal research in Italy and were carried out under the permission of the Region of Lombardy. This is paper #10 of the ASPER project.

Author information

Authors and Affiliations

Authors

Corresponding author

Correspondence to Lucas A. Wauters.

About this article

Cite this article

Pierro, E.D., Molinari, A., Tosi, G. et al. Exclusive core areas and intrasexual territoriality in Eurasian red squirrels (Sciurus vulgaris) revealed by incremental cluster polygon analysis. Ecol Res 23, 529–542 (2008). https://doi.org/10.1007/s11284-007-0401-0

Download citation

  • Received:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI: https://doi.org/10.1007/s11284-007-0401-0

Keywords

Navigation