Skip to main content
SpringerLink
Log in

Costs of territoriality: a review of hypotheses, meta-analysis, and field study

  • Behavioral ecology–original research
  • Published:
Oecologia Aims and scope Submit manuscript

Abstract

The evolution of territoriality reflects the balance between the benefit and cost of monopolising a resource. While the benefit of territoriality is generally intuitive (improved access to resources), our understanding of its cost is less clear. This paper combines: 1. a review of hypotheses and meta-analytic benchmarking of costs across diverse taxa; and 2. a new empirical test of hypotheses using a longitudinal study of free-living male territorial lizards. The cost of territoriality was best described as a culmination of multiple factors, but especially costs resulting from the time required to maintain a territory (identified by the meta-analysis) or those exacerbated by a territory that is large in size (identified by the empirical test). The meta-analysis showed that physiological costs such as energetic expenditure or stress were largely negligible in impact on territory holders. Species that used territories to monopolise access to mates appeared to incur the greatest costs, whereas those defending food resources experienced the least. The single largest gap in our current understanding revealed by the literature review is the potential cost associated with increased predation. There is also a clear need for multiple costs to be evaluated concurrently in a single species. The empirical component of this study showcases a powerful analytical framework for evaluating a range of hypotheses using correlational data obtained in the field. More broadly, this paper highlights key factors that should be considered in any investigation that attempts to account for the evolutionary origin or ecological variation in territorial behaviour within and between species.

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

Access this article

Log in via an institution

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

Instant access to the full article PDF.

Institutional subscriptions

Fig. 1
Fig. 2
Fig. 3
Fig. 4

taken from associated path models (Table 4; see also Figure S4)

Similar content being viewed by others

Availability of data and materials

All data associated with this study is publicly available through the Dryad digital data repository (https://doi.org/10.5061/dryad.r7sqv9sd1).

Code availability

All code associated with this study is available from the author (t.ord@unsw.edu.au).

References

  • Abell AJ (2000) Costs of reproduction in male lizards, Sceloporus virgatus. Oikos 88:630–640

    Google Scholar 

  • Adams ES (2001) Approaches to the study of territory size and shape. Ann Rev Ecol Syst 32:277–303

    Google Scholar 

  • Aluja M, Perez-Staples D, Sivinski J, Sanchez A, Pinero J (2008) Effects of male condition on fitness in two tropical tephritid flies with contrasting life histories. Anim Behav 76:1997–2009

    Google Scholar 

  • Amsler SJ (2010) Energetic costs of territorial boundary patrols by wild chimpanzees. Am J Primatol 72:93–103

    PubMed  Google Scholar 

  • Ancona S, Drummond H, Zaldivar-Rae J (2010) Male whiptail lizards adjust energetically costly male guarding to male-male competition and female reproductive value. Anim Behav 79:75–82

    Google Scholar 

  • Arvidsson B, Askenmo C, Neergaard R (1997) Food supply for settling male rock pipits affects territory size. Anim Behav 54:67–72

    CAS  PubMed  Google Scholar 

  • Bachman G, Widemo F (1999) Relationships between body composition, body size and alternative reproductive tactics. Funct Ecol 13:411–416

    Google Scholar 

  • Baldo S, Mennill DJ, Guindre-Parker S, Gilchrist HG, Love OP (2015) The oxidative cost of acoustic signals: examining steroid versus aerobic activity hypotheses in a wild bird. Ethology 121:1081–1090

    Google Scholar 

  • Barrette C, Vandal D (1990) Sparring, relative antler size, and assessment in male caribou. Behav Ecol Sociobiol 26:383–387

    Google Scholar 

  • Berger-Tal O, Embar K, Kotler BP, Saltz D (2015) Everybody loses: intraspecific competition induces tragedy of the commons in Allenby’s gerbils. Ecology 96:54–61

    PubMed  Google Scholar 

  • Border SE, DeOliveira GM, Janeski HM, Piefke TJ, Brown TJ, Dijkstra PD (2019) Social rank, color morph, and social network metrics predict oxidative stress in a cichlid fish. Behav Ecol 30:490–499

    Google Scholar 

  • Borenstein M, Hedges LV, Higgins JPT, Rothstein HR (2009) Introduction to meta-analysis. John Wiley & Sons, Chichester, UK

    Google Scholar 

  • Brown DR, Sherry TW (2008) Alternative strategies of space use and response to resource change in a wintering migrant songbird. Behav Ecol 19:1314–1325

    Google Scholar 

  • Burnham KP, Anderson DR (2002) Model selection and multimodal inference: a practical information-theoretic approach, 2nd edn. Springer, New York

    Google Scholar 

  • Burnham KP, Anderson DR, Huyvaert KP (2011) AIC model selection and multimodel inference in behavioral ecology: some background, observations, and comparisons. Behav Ecol Sociobiol 65:23–35

    Google Scholar 

  • Christensen C, Radford AN (2018) Dear enemies or nasty neighbors? Causes and consequences of variation in the responses of group-living species to territorial intrusions. Behav Ecol 29:1004–1013

    Google Scholar 

  • Cleveland A (1999) Energetic costs of agonistic behavior in two herbivorous damselfishes (Stegastes). Copeia 1999:857–867

    Google Scholar 

  • Clutton-Brock TH, Albon SD (1979) The roaring of red deer and the evolution of honest advertisement. Behaviour 69:145–169

    Google Scholar 

  • Cohen J (1969) Statistical power analysis for the behavioral sciences. Academic Press, New York

    Google Scholar 

  • Contreras-Garduno J, Canales-Lazcano J, Cordoba-Aguilar A (2006) Wing pigmentation, immune ability, fat reserves and territorial status in males of the rubyspot damselfly, Hetaerina americana. J Ethology 24:165–173

    Google Scholar 

  • Cooper WE Jr (1999) Tradeoffs between courtship, fighting, and antipredatory behavior by a lizard, Eumeces laticeps. Behav Ecol Sociobiol 47:54–59

    Google Scholar 

  • Corlatti L, Bassano B, Valencak TG, Lovari S (2013) Foraging strategies associated with alternative reproductive tactics in a large mammal. J Zool 291:111–118

    Google Scholar 

  • Del Re AC (2013) compute.es: compute effect sizes. Version 0.2-4. http://cran.r-project.org/web/packages/compute.es: R package version 0.2-4

  • Del Re AC, Hoyt WT (2014) MAd: meta-analysis with mean differences. Version 0.8-2.1. https://cran.r-project.org/package=MAd: R package version 0.8-2.1

  • Dias RI, Castilho L, Macedo RH (2010) Experimental evidence that sexual displays are costly for nest survival. Ethology 116:1011–1019

    Google Scholar 

  • Diaz-Uriarte R (1999) Anti-predator behaviour changes following an aggressive encounter in the lizard Tropidurus hispidus. Proc Roy Soc Lond B 266:2457–2464

    CAS  Google Scholar 

  • Endler JA (1982) Convergent and divergent effects of natural selection on color patterns in two fish faunas. Evolution 36:178–188

    PubMed  Google Scholar 

  • Ezenwa VO, Snider MH (2016) Reciprocal relationships between behaviour and parasites suggest that negative feedback may drive flexibility in male reproductive behaviour. Proc Roy Soc Lond B 283:20160423

    Google Scholar 

  • Garamszegi LZ (2006) Comparing effect sizes across variables: generalization without the need for Bonferroni correction. Behav Ecol 17:682–687

    Google Scholar 

  • Gordon NM (2004) The effect of supplemental feeding on the territorial behavior of the green frog (Rana clamitans). Amphibia Reptilia 25:55–62

    Google Scholar 

  • Isvaran K, Jhala Y (2000) Variation in lekking costs in blackbuck (Antilope cervicapra): relationship to lek-territory location and female mating patterns. Behaviour 137:547–563

    Google Scholar 

  • Jaeger RG (1981) Dear enemy recognition and the costs of aggression between salamanders. Am Nat 117:962–974

    Google Scholar 

  • Jakobsson S, Brick O, Kullberg C (1995) Escalated fighting behaviour incurs increased predation risk. Anim Behav 49:235–239

    Google Scholar 

  • Keeley ER (2000) An experimental analysis of territory size in juvenile steelhead trout. Anim Behav 59:477–490

    CAS  PubMed  Google Scholar 

  • Kokko H, Lopez-Fernandez A, Morrell LJ (2006) From hawks and doves to self-consistent games of territorial behavior. Am Nat 167:901–912

    PubMed  Google Scholar 

  • Krams I (2001) Communication in crested tits and the risk of predation. Anim Behav 61:1065–1068

    Google Scholar 

  • Lebigre C, Alatalo RV, Siitari H (2013) Physiological costs enforce the honesty of lek display in the black grouse (Tetrao tetrix). Oecologia 172:983–993

    PubMed  Google Scholar 

  • Low M (2006) The energetic cost of mate guarding is correlated with territoria intrusions in the New Zealand stitchbird. Behav Ecol 17:270–276

    Google Scholar 

  • Maan ME, Groothuis TGG, Wittenberg J (2001) Escalated fighting despite predictors of conflict outcome: solving the paradox in a South American cichlid fish. Anim Behav 62:623–634

    Google Scholar 

  • Marchetti GM, Drton M, Sadeghi K (2020) ggm: Graphical Markov models with mixed graphs. Version 2.5. http://cran.r-project.org/web/packages/ggm. R package version 2.5

  • Marino A (2012) Indirect measures of reproductive effort in a resource-defense polygynous ungulate: territorial defense by male guanacos. J Ethol 30:83–91

    Google Scholar 

  • Martin J, Lopez P (2001) Risk of predation may explain the absence of nuptial coloration in the wall lizard, Podarcis muralis. Evol Ecol Res 3:889–898

    Google Scholar 

  • Martinez-Padilla J, Perez-Rodriguez L, Mougeot F, Ludwig S, Redpath SM (2014) Intra-sexual competition alters the relationship between testosterone and ornament expression in a wild territorial bird. Horm Behav 65:435–444

    CAS  PubMed  Google Scholar 

  • McGregor PK, Dabelsteen T, Shepherd M, Pedersen SB (1992) The signal value of matched singing in great tits: evidence from interactive playback experiments. Anim Behav 43:987–998

    Google Scholar 

  • McLeod BT, Ritchison G (2018) Effects of supplemental food on the behaviour and paternity status of male Indigo Buntings (Passerina cyanea). Avian Biol Res 11:67–73

    Google Scholar 

  • Moretz JA, Morris MR (2006) Phylogenetic analysis of the evolution of a signal of aggressive intent in northern swordtail fishes. Am Nat 168:336–349

    PubMed  Google Scholar 

  • Munguia-Steyer R, Cordoba-Aguilar A, Romo-Beltran AR (2010) Do individuals in better condition survive for longer? Field survival estimates according to male alternative reproductive tactics and sex. J Evol Biol 23:175–184

    CAS  PubMed  Google Scholar 

  • Nakagawa S (2004) A farewell to Bonferroni: the problems of low statistical power and publication bias. Behav Ecol 15:1044–1045

    Google Scholar 

  • Nakagawa S, Noble DWA, Senior AM, Lagisz M (2017) Meta-evaluation of meta-analysis: ten appraisal questions for biologists. BMC Biol. https://doi.org/10.1186/s12915-017-0357-7

    Article  PubMed  PubMed Central  Google Scholar 

  • Noel MV, Grant JWA, Carrigan JG (2005) Effects of competitor-to-resource ratio on aggression and size variation within groups of convict cichlids. Anim Behav 69:1157–1163

    Google Scholar 

  • O’Brien DM, Katsuki M, Emlen DJ (2017) Selection on an extreme weapon in the frog-legged leaf beetle (Sagra femorata). Evolution 71:2584–2598

    PubMed  Google Scholar 

  • Ord TJ, Stamps JA (2008) Alert signals enhance animal communication in “noisy” environments. Proc Nat Acad Sci USA 105:18830–18835

    CAS  PubMed  PubMed Central  Google Scholar 

  • Ord TJ, Stamps JA (2017) Why does the rate of signal production in ectotherms vary with temperature? Behav Ecol 28:1272–1282

    Google Scholar 

  • Ord TJ, Blumstein DT, Evans CS (2001) Intrasexual selection predicts the evolution of signal complexity in lizards. Proc Roy Soc Lond B 268:737–744

    CAS  Google Scholar 

  • Ord TJ, Peters RA, Clucas B, Stamps JA (2007) Lizards speed up visual displays in noisy motion habitats. Proc Roy Soc Lond B 274:1057–1062

    Google Scholar 

  • Ord TJ, Stamps JA, Losos JB (2010) Adaptation and plasticity of animal communication in fluctuating environments. Evolution 64:3134–3148

    PubMed  Google Scholar 

  • Ord TJ, King L, Young AR (2011) Contrasting theory with the empirical data of species recognition. Evolution 65:2572–2591

    PubMed  Google Scholar 

  • Ord TJ, Charles GK, Palmer M, Stamps JA (2016) Plasticity in social communication and its implications for the colonization of novel habitats. Behav Ecol 27:341–351

    Google Scholar 

  • Ord TJ, Blazek K, White TE, Das I (2021) Conspicuous animal signals can avoid the cost of predation by being intermittent or novel: confirmation in the wild using hundreds of robotic prey. Proc Roy Soc Lond B 288:20210706

    Google Scholar 

  • Packer C, Hilborn R, Mosser A, Kissui B, Borner M, Hopcraft G, Wilmshurst J, Mduma S, Sinclair ARE (2005) Ecological change, group territoriality, and population dynamics in Serengeti lions. Science 307:390–393

    CAS  PubMed  Google Scholar 

  • Pearson HC, Packard JM, Davis RW (2006) Territory quality of male sea otters in Prince William Sound, Alaska: relation to body and territory maintenance behaviors. Can J Zool 84:939–946

    Google Scholar 

  • Peixoto PE, Benson WW (2008) Body mass and not wing length predicts territorial success in a tropical satyrine butterfly. Ethology 114:1069–1077

    Google Scholar 

  • Peixoto PEC, Benson WW (2011) Fat and body mass predict residency status in two tropical satyrine butterflies. Ethology 117:722–730

    Google Scholar 

  • Peters RA, Clifford CWG, Evans CS (2002) Measuring the structure of dynamic visual signals. Anim Behav 64:131–146

    Google Scholar 

  • Port M, Schulke O, Ostner J (2017) From individual to group territoriality: competitive environments promote the evolution of sociality. Am Nat 189:E46–E57

    PubMed  Google Scholar 

  • Praw JC, Grant JWA (1999) Optimal territory size in the convict cichlid. Behaviour 136:1347–1363

    Google Scholar 

  • Rich LN, Mitchell MS, Gude JA, Sime CA, Van Vuren H (2012) Anthropogenic mortality, intraspecific competition, and prey availability influence territory sizes of wolves in Montana. J Mammal 93:722–731

    Google Scholar 

  • Riebli T, Taborsky M, Chervet N, Apolloni N, Zurcher Y, Heg D (2012) Behavioural type, status and social context affect behaviour and resource allocation in cooperatively breeding cichlids. Anim Behav 84:925–936

    Google Scholar 

  • Rimbach R, Blanc S, Zahariev A, Pillay N, Schradin C (2019) Daily energy expenditure of males following alternative reproductive tactics: solitary roamers spend more energy than group-living males. Physiol Behav 199:359–365

    CAS  PubMed  Google Scholar 

  • Romo-Beltran A, Macias-Ordonez R, Cordoba-Aguilar A (2009) Male dimorphism, territoriality and mating success in the tropical damselfly, Paraphlebia zoe Selys (Odonata: Megapodagrionidae). Evol Ecol 23:699–709

    Google Scholar 

  • Ros AFH, Groothuis TGG, Apanius V (1997) The relation among gonadal steroids, immunocompetence, body mass, and behavior in young black-headed gulls (Larus ridibundus). Am Nat 150:201–219

    CAS  PubMed  Google Scholar 

  • Rosenthal GG, Martinez TYF, de Leon FJG, Ryan MJ (2001) Shared preferences by predators and females for male ornaments in swordtails. Am Nat 158:146–154

    CAS  PubMed  Google Scholar 

  • Rusch TW, Angilletta MJ Jr (2017) Competition during thermoregulation altered the body temperatures and hormone levels of lizards. Funct Ecol 31:1519–1528

    Google Scholar 

  • Schradin C, Schmohl G, Rodel HG, Schoepf I, Treffler SM, Brenner J, Bleeker M, Schubert M, Konig B, Pillay N (2010) Female home range size is regulated by resource distribution and intraspecific competition: a long-term field study. Anim Behav 79:195–203

    Google Scholar 

  • Schutz D, Pachler G, Ripmeester E, Goffinet O, Taborsky M (2010) Reproductive investment of giants and dwarfs: specialized tactics in a cichlid fish with alternative male morphs. Funct Ecol 24:131–140

    Google Scholar 

  • Shipley B (2013) The AIC model selection method applied to path analytic models compared using a d-separation test. Ecology 94:560–564

    PubMed  Google Scholar 

  • Siracusa ER, Boutin S, Dantzer B, Lane JE, Coltman DW, McAdam AG (2021) Familiar neighbors, but not relatives, enhance fitness in a territorial mammal. Curr Biol 31:438–445

    CAS  PubMed  Google Scholar 

  • Stamps JA (1977) The function of the survey posture in Anolis lizards. Copeia 1977:756–758

    Google Scholar 

  • Stamps JA (1983) Sexual selection, sexual dimorphism, and territoriality. In: Huey RB, Pianka ER, Schoener TW (eds) Lizard ecology: studies of a model organism. Harvard University Press, Cambridge, MA, pp 169–204

    Google Scholar 

  • Stamps JA, Tanaka S (1981) The relationship between food and social behaviour in juvenile lizards (Anolis aeneus). Copeia 1981:422–434

    Google Scholar 

  • Steinberg DS, Losos JB, Schoener TW, Spiller DA, Kolbe JJ, Leal M (2014) Predation-associated modulation of movement-based signals by a Bahamian lizard. Proc Nat Acad Sci USA 111:9187–9192

    CAS  PubMed  PubMed Central  Google Scholar 

  • Stuart-Fox D, Moussalli A, Marshall NJ, Owens IPF (2003) Conspicuous males suffer higher predation risk: visual modelling and experimental evidence from lizards. Anim Behav 66:541–550

    Google Scholar 

  • Symonds MRE, Moussalli A (2011) A brief guide to model selection, multimodel inference and model averaging in behavioural ecology using Akaike’s information criterion. Behav Ecol Sociobiol 65:13–21

    Google Scholar 

  • Takeuchi T (2006) The effect of morphology and physiology on butterfly territoriality. Behaviour 143:393–403

    Google Scholar 

  • Temeles EJ (1994) The role of neighbours in territorial systems: when are they “dear enemies”? Anim Behav 47:339–350

    Google Scholar 

  • Tiebout HM (1993) Mechanisms of competition in tropical hummingbirds: metabolic costs for losers and winners. Ecology 74:405–418

    Google Scholar 

  • Tuttle MD, Ryan MJ (1981) Bat predation and the evolution of frog vocalizations in the neotropics. Science 214:677–678

    CAS  PubMed  Google Scholar 

  • Velde LV, Van Dyck H (2013) Lipid economy, flight activity and reproductive behaviour in the speckled wood butterfly: on the energetic cost of territory holding. Oikos 122:555–562

    Google Scholar 

  • Viechtbauer W (2010) Meta-analysis package for R. Version 2.1-0. http://www.wvbauer.com/

  • Viera VM, Viblanc VA, Filippi-Codaccioni O, Cote SD, Groscolas R (2011) Active territory defence at a low energy cost in a colonial seabird. Anim Behav 82:69–76

    Google Scholar 

  • Vollestad LA, Quinn TP (2003) Trade-off between growth rate and aggression in juvenile coho salmon, Oncorhynchus kisutch. Anim Behav 66:561–568

    Google Scholar 

  • von Kuerthy C, Tschirren L, Taborsky M (2015) Alternative reproductive tactics in snail shell-brooding cichlids diverge in energy reserve allocation. Ecol Evol 5:2060–2069

    Google Scholar 

  • Waas JR (1991) Do little blue penguins signal their intentions during aggressive interactions with strangers? Anim Behav 41:375–382

    Google Scholar 

  • White AJ, Rundle HD (2015) Territory defense as a condition-dependent component of male reproductive success in Drosophila serrata. Evolution 69:407–418

    PubMed  Google Scholar 

  • Wikelski M, Trillmich F (1994) Foraging strategies of the Galapagos marine iguana (Amblyrhynchus cristatus): adapting behavioral rules to ontogenetic size change. Behaviour 128:255–279

    Google Scholar 

  • Willisch CS, Ingold P (2007) Feeding or resting? The strategy of rutting male alpine chamois. Ethology 113:97–104

    Google Scholar 

  • Young AJ, Monfort SL (2009) Stress and the costs of extra-territorial movement in a social carnivore. Biol Lett 5:439–441

    PubMed  PubMed Central  Google Scholar 

Download references

Acknowledgements

I am especially grateful to Grace Charles and Meredith Palmer for their assistance in the field on the original longitudinal study of A. gundlachi territorial behaviour, and C. Tucker Pforzheimer for his tireless efforts in creating the detailed territorial maps. Three anonymous reviewers provided many thoughtfully points that prompted extensive revisions and new analyses being included that greatly improved this paper. The empirical work outlined in this paper was approved by research permits from the Caribbean National Forest and Department of Natural Resources and Environment of Puerto Rico.

Funding

The longitudinal study described in this paper was supported by a grant from the National Science Foundation (IOB-0517041/0516998).

Author information

Authors and Affiliations

  1. Evolution and Ecology Research Centre, and School of Biological, Earth and Environmental Sciences, University of New South Wales, Kensington, NSW, Australia

    Terry J. Ord

Authors
  1. Terry J. Ord
    View author publications

    You can also search for this author in PubMed Google Scholar

Contributions

TJO conceived, designed, and executed the study and wrote the manuscript. No other person is entitled to authorship.

Corresponding author

Correspondence to Terry J. Ord.

Ethics declarations

Conflict of interest

There are no conflicts or competing interests associated with this work.

Ethics approval

The empirical work outlined in this paper was covered under Animal Use and Care Protocols 05-11652/15243 from the Institutional Animal Care and Use Committee of the University of California at Davis.

Consent to participate

Not applicable.

Consent for publication

Not applicable.

Additional information

Communicated by Jean-François Le Galliard.

Supplementary Information

Below is the link to the electronic supplementary material.

Fig. S1

PRISMA flow diagram outlining the approach of reviewing and screening literature for summarising hypotheses previously investigated by empirical investigations of territorial costs and benchmarking those hypotheses using standardised effect size analysis (i.e. meta-analysis). Supplementary file1 (PPTX 45 KB)

Fig. S2

An example map of a male A. gundlachi lizard territory showing estimated territory size (in red) and perches used during the study. Neighbour positions were inferred by compass bearing and distance from the central perch.. Supplementary file2 (EPS 2022 KB)

Fig. S3

The (a) estimated population growth rate of male A. gundlachi at the study location used to compute changes in body condition (the residual value estimated for a male at the start of the study subtracted from the residual value estimated for that same male at the end of the study). Males (b) in better condition at the start of the study were typically those with the largest territories. Supplementary file3 (XLSX 21 KB)

Fig. S4

The range of plausible path models that might account for changes in body condition of male A. gundlachi lizards depending on the hypothesis supported (see Figure 1). Alternative models within hypotheses are grouped by arrow colour.. Supplementary file4 (PPTX 44 KB)

Fig. S5

Effect sizes and study degrees of freedom for the range of species included in the meta-analysis. Supplementary file5 (EPS 2341 KB)

Rights and permissions

Reprints and permissions

About this article

Check for updates. Verify currency and authenticity via CrossMark

Cite this article

Ord, T.J. Costs of territoriality: a review of hypotheses, meta-analysis, and field study. Oecologia 197, 615–631 (2021). https://doi.org/10.1007/s00442-021-05068-6

Download citation

  • Received:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI: https://doi.org/10.1007/s00442-021-05068-6

Keywords

Search

Navigation