Varanid combat: Perspectives from game theory

Amphibia Reptilia

Volumn 23, Issue 4, 2002, Pages 469-485

  Earley, Ryan L ^a   Attum, Omar ^b   Eason, Perri ^b  

^a GEORGIA STATE UNIVERSITY   (United States)

^b UNIVERSITY OF LOUISVILLE   (United States)

Author keywords

[No Author keywords available]

Indexed keywords

AGGRESSION; CONTEST COMPETITION; GAME THEORY; LIZARD;

AMPHIBIA; ANIMALIA; REPTILIA; SQUAMATA;

EID: 0037004422     PISSN: 01735373     EISSN: None     Source Type: Journal    
DOI: 10.1163/15685380260462374     Document Type: Article

References (93)

1. Ali, S. (1944): The 'courtship' of the monitor lizard (Varanus monitor). J. Bombay Nat. Hist. Soc. 44: 479-480.
2. Andersen, I.L., Andenaes, H., Boe, K.E., Jensen, P., Bakken, M. (2000): The effects of weight asymmetry and resource distribution on aggression in groups of unacquainted pigs. Appl. Anim. Behav. Sci. 68: 107-120.
3. Attum, O., Earley, R.L., Bayless, M., Eason, P. (2000): The agonistic behaviour of Bose's monitor (Varanus exanthematicus Bose, 1792) in captivity. Herp. Bull. 73: 22-26.
4. Auffenberg, W. (1978): Social and feeding behaviour in Varanus komodoensis. In: Behaviour and Neurology of Lizards: An Interdisciplinary Colloquium, p. 301-331. Greenberg, N., MacLean, P.D., Eds, Rockville. Maryland, National Institute of Mental Health.
5. Auffenberg, W. (1979): Intersexual differences in behaviour of captive Varanus bengalensis (Reptilia, Lacertilia, Varanidae). J. Herp. 13: 313-315.
6. Auffenberg, W. (1981a): The Behavioural Ecology of the Komodo Monitor. Gainesville, University of Florida Press.
7. Auffenberg, W. (1981b): Combat behaviour in Varanus bengalensis (Sauria: Varanidae). J. Bombay Nat. Hist. Soc. 78: 54-72.
8. Auffenberg, W. (1983): The burrows of Varanus bengalensis: characteristics and use. Rec. Zool. Surv. India 80: 375-385.
9. Auffenberg, W. (1988): Gray's Monitor Lizard. Gainesville, University of Florida Press.
10. Auffenberg, W. (1994): The Bengal Monitor. Gainesville, University of Florida Press.
11. Bayless, M. (1994): Zur Fortpflanzungbiologie des Steppenwarans (Varanus exanthematicus). Salamandra 30: 109-118.
12. Bels, V.L., Gasc, J.-P., Goosse, V., Renous, S., Vernet, R. (1995): Functional analysis of the throat display in the sand goanna Varanus griseus (Reptilia: Squamata: Varanidae). J. Zool. Lond. 235: 95-116.
13. Bennett, A.F. (1982): The energetics of reptilian activity. In: Biology of the Reptilia. Volume 13 (Physiology), p. 155-199. Gans, C., Pough, F.H., Eds. New York. Academic Press.
14. Bennett, D. (1998): Monitor Lizards: Natural History, Biology and Husbandry. Edition Chimaira, Frankfurt am Main, Meckenheim, Germany.
15. Bishop, A.T., Cannings, C. (1978): A generalized war of attrition. J. theor. Biol. 70: 85-125.
16. Branch, W.R. (1991): The Regenia registers of 'Gogga' Brown (1869-1909): "Memoranda on a species of monitor or varan". Mertensiella 2: 57-110.
17. Bridge, A.P., Elwood, R.W., Dick, J.T.A. (2000): Imperfect assessment and limited information preclude optimal strategies in male-male fights in the orb-weaving spider Metellina mengei. Proc. R. Soc. Lond. B 267: 273-279.
18. Briffa, M., Elwood, R.W. (2000): Cumulative or sequential assessment during hermit crab shell fights: effects of oxygen on decision rules. Proc. R. Soc. Lond. B 267: 2445-2452.
19. Carpenter, C.C., Ferguson, G.W. (1977): Variation and evolution of stereotyped behaviour in reptiles. In: Biology of the Reptilia. Volume 7A (Ecology and Behaviour), p. 138-208. Gans, C., Tinkle, D.W., Eds, New York. Academic Press.
20. Carpenter, C.C., Gillingham, J.C., Murphy, J.B., Mitchell, L.A. (1976): A further analysis of the combat ritual of the pygmy mulga monitor, Varanus gilleni (Reptilia: Varanidae). Herpetologica 32: 35-40.
21. Christian, K., Weavers, B. (1994): Analysis of the activity and energetics of the lizard Varanus rosenbergi. Copeia 1994: 289-295.
22. Clements, B.N. (1968): Lizard Love. Lammergeyer 9: 53.
23. Daltry, J. (1991): The social hierarchy of the water monitor, Varanus salvator. Hamadryad 16: 10-20.
24. Davis, R., Darling, R., Darlington, A. (1986): Ritualized combat in captive Dumeril's monitors, Varanus dumerili. Herp Rev. 17: 85-88.
25. Delean, S. (1981): Notes on aggressive behaviour by Gould's goannas (Varanus gouldii) in captivity. Herpetofauna 12: 31.
26. Deraniyagala, R.Y. (1958): Pseudo-combat of the monitor lizard Varanus bengalensis (Daudin). Spolia Zeylandica 28: 11-13.
27. Dugatkin, L.A., Biederman, L. (1991): Balancing asymmetries in resource holding power and resource value in the pumpkinseed sunfish. Anim. Behav. 42: 691-692.
28. Dugatkin, L.A., Ohlsen, S.R. (1990): Contrasting asymmetries in value expectation and resource holding power: effects on attack behaviour and dominance in the pumpkinseed sunfish. Lepomis gibbosus. Anim. Behav. 39: 802-804.
29. Dugatkin, L.A., Reeve, H.K. (1998): Game Theory and Animal Behaviour. New York, Oxford University Press.
30. Earley, R.L., Hsu, Y., Wolf, L.L. (2000): The use of standard aggression testing methods to predict combat behaviour and contest outcome in Rivulus marmoratus dyads (Teleostei: Cyprinodontidae). Ethology 106: 743-761.
31. Englund, G., Olsson, T.I. (1990): Fighting and assessment in the net-spinning caddis larva Arctopsyche ladogensis: a test of the sequential assessment game. Anim. Behav. 39: 55-62.
32. Enquist, M., Leimar, O. (1983): Evolution of fighting behaviour: Decision rules and assessment of relative strength. J. theor. Biol. 102: 387-410.
33. Enquist, M., Leimar, O. (1987): Evolution of fighting behaviour: The effect of variation in resource value. J. theor. Biol. 127: 187-205.
34. Enquist, M., Leimar, O., Ljundberg, T., Mallner, Y., Segerdahl, N. (1990): A test of the sequential assessment game: fighting in the cichlid fish Nannacara anomala. Anim. Behav. 40: 1-14.
35. Gleeson, T.T. (1991): Patterns of metabolic recovery from exercise in amphibians and reptiles. J. Exper. Biol. 160: 187-207.
36. Grafen, A. (1990): Biological signals as handicaps. J. theor. Biol. 144: 517-546.
37. Hagen, H., Horn, H.-G., Hagen, W. (1995): Zu Balz-und Kopulationsverhalten des Nilwarans (Varanus n. niloticus) - Cine fotografische Dokumentation. Herpetofauna 17: 29-33.
38. Haller, J. (1995): Biochemical background for an analysis of cost-benefit interrelations in aggression. Neurosci. Biobehav. Rev. 19: 599-604.
39. Hammerstein, P. (1981): The role of asymmetries in animal contests. Anim. behav. 29: 193-205.
40. Hicks, J.W., Wang, T., Bennett, A.F. (2000): Patterns of cardiovascular and ventilatory response to elevated metabolic states in the lizard Varanus exanthematicus. J. Exper. Biol. 203: 2437-2445.
41. Hofmann, H.A., Schildberger, K. (2001): Assessment of strength and willingness to fight during aggressive encounters in crickets. Anim. Behav. 62: 337-348.
42. Honneger, R.E., Heusser, H. (1969): Beiträge zum Verhaltensinventar des Bindenwarans (varanus salvator). Zool. Garten NF 36: 251-260.
43. Horn, H.-G. (1985): Beiträge zum Verhalten von Waranen: Die Ritualkämpfe von Varanus komodoensis Ouwens. 1912 und V. semiremex Peters 1869 sowie die Imponierphasen der Ritualkämpfe von V. timorensis timorensis (Gray, 1831) und V.t. similis Mertens, 1958 (Sauria: Varanidae). Salamandra 21: 169-179.
44. Horn, H.-G. (1994): Der Ritualkampf von Varanus salvator cumingi. Herpetofauna 16: 27-30.
45. Horn, H.-G., Gaulke, M., Böhme, W. (1994): New data on ritualized combats in monitor lizards (Sauria: Varanidae), with remarks on their function and phylogenetic implications. Zool. Garten NF 64: 265-280.
46. Hsu, Y., Wolf, L.L. (1999): The winner and loser effect: integrating multiple experiences. Anim. Behav. 57: 903-910.
47. James, C.D. (1996): Ecology of the pygmy goanna (Varanus brevicauda) in spinifex grasslands of Central Australia. Aust. J. Zool. 44: 177-192.
48. Jensen, P., Yngvesson, J. (1998): Aggression between unacquainted pigs - sequential assessment and effects of familiarity and weight. Appl. Anim. Behav. Sci. 58: 49-61.
49. Kaufman, J.D., Burghardt, G.M., Phillips, J.A. (1994): Density-dependent foraging strategy of a large carnivorous lizard, the savanna monitor (Varanus albigularis). J. Comp. Psychol. 108: 381-384.
50. Kaufman, J.D., Burghardt, G.M., Phillips, J.A. (1996): Sensory cues and foraging decisions in a large carnivorous lizard, Varanus albigularis. Anim. Behav. 52: 727-736.
51. Keeley, E.R., Grant, J.W.A. (1993): Visual information, resource value, and sequential assessment in convict cichlid (Cichlasoma nigrofasciatum) contests. Behav. Ecol. 4: 345-349.
52. Knapp, R., Moore, M.C. (1996): Male morphs in tree lizards, Urosaurus ornatus, have different delayed hormonal responses to aggressive encounters. Anim. Behav. 52: 1045-1055.
53. Koops, M.A., Grant, J.W.A. (1993): Weight asymmetry and sequential assessment in convict cichlid contests. Can. J. Zool. 71: 475-479.
54. Lederer, G. (1929): Ein zahmer Bindenwaran. Waschr. Aquar. Terrar. Kunde 26: 19-20.
55. Leimar, O., Austad, S., Enquist, M. (1991): A test of the sequential assessment game: fighting in the bowl and doily spider Frontinella pyramitela. Evolution 45: 862-874.
56. Leimar, O., Enquist, M. (1984): Effects of asymmetries in owner-intruder interactions, J. theor. Biol. 111: 475-491.
57. Marden, J.H., Waage, J.K. (1990): Escalated damselfly territorial contests are energetic wars of attrition. Anim. Behav. 39: 954-959.
58. Marden, J.H., Rollins, R.A. (1994): Assessment of energy reserves by damselfies engaged in aerial contests for mating territories. Anim. Behav. 48: 1023-1030.
59. Maynard Smith, J. (1982): Evolution and The Theory of Games. Cambridge, Cambridge University Press.
60. Maynard Smith, J., Parker, G.A. (1976): The logic of asymmetric contests. Anim. Behav. 24: 159-175.
61. Maynard Smith, J., Price, G.R. (1973): The logic of animal conflict. Nature 246: 15-18.
62. McCoid, M.J., Hensley, R.A. (1991): Mating and combat in Varanus indicus. Herp Rev. 22: 16-17.
63. McMann, S. (1993): Contextual signalling and the structure of dyadic encounters in Anolis carolinensis. Anim. Behav. 46: 657-668.
64. Molina-Borja, M., Padron-Fumero, M., Alfonso-Martin, T. (1998): Morphological and behavioural traits affecting intensity and outcome of male contests in Gallotia galloti galloti (family Lacertidae). Ethology 104: 314-322.
65. Moya-Larano, J., Wise, D.H. (2000): Survival regression analysis: a powerful tool for evaluating fighting and assessment. Anim. Behav. 60: 307-313.
66. Murphy, J.B., Mitchell, L.A. (1974): Ritualized combat behaviour of the pygmy mulga monitor lizard, Varanus gilleni (Sauria: Varanidae). Herpetologica 30: 90-97.
67. Neat, F.C., Taylor, A.C., Huntingford, F.A. (1998): Proximate costs of fighting in male cichlid fish: the role of injuries and energy metabolism. Anim. Behav. 55: 875-882.
68. Olsson, M. (1992): Contest success in relation to size and residency in male sand lizards. Lacerta agilis. Anim. Behav. 44: 386-388.
69. Olsson, M., Shine, R. (2000): Ownership influences the outcome of male-male contests in the scincid lizard, Niveoscincus microlepidotus. Behav. Ecol. 11: 587-590.
70. Parker, G.A. (1974): Assessment strategy and the evolution of fighting behaviour, J. theor. Biol. 47: 223-243.
71. Parker, G.A., Rubenstein, D.I. (1981): Role assessment, reserve strategy, and acquisition of information in asymmetric animal conflicts. Anim. Behav. 29: 221-240.
72. Payne, R.J.H. (1998): Gradually escalating fights and displays: the cumulative assessment model. Anim. Behav. 56: 651-662.
73. Payne, R.J.H., Pagel, M. (1997): Why do animals repeat displays? Anim. Behav. 54: 109-119.
74. Phillips, J.A. (1995): Movement patterns and density of Varanus albigularis. J. Herp. 29: 407-416.
75. Phillips, J.A., Millar, R.P. (1998): Reproductive biology of the white-throated savanna monitor Varanus albigularis. J. Herp. 32: 366-377.
76. Pianka, E.R. (1968): Notes on the biology of Varanus eremius. W. Aust. Nat. 11: 39-44.
77. Pianka, E.R. (1969): Notes on the biology of Varanus caudolineatus and Varanus gilleni. W. Aust. Nat. 11: 76-82.
78. Pianka, E.R. (1970): Notes on the biology of Varanus gouldii flavirufus. W. Aust. Nat. 11: 141-144.
79. Schuett, G.W. (1997): Body size and agonistic experience affect dominance and mating success in male copperheads. Anim. Behav. 54: 213-224.
80. Schuett, G.W., Grober, M.S. (2000): Post-fight levels of plasma lactate and corticosterone in male copperheads, Agkistrodon contortrix (Serpentes, Viperidae): differences between winners and losers, Physiol. Behav. 71: 335-341.
81. Sneddon, L.U., Taylor, A.C., Huntingford, F.A. (1999): Metabolic consequences of agonistic behaviour: crab fights in declining oxygen tensions. Anim. Behav. 57: 353-363.
82. Stamps, J.A., Krishnan, V.V. (1994): Territory acquisition in lizards: II. Establishing social and spatial relationships. Anim. Behav. 47: 745-753.
83. Sterling, E.C. (1912): Observations on the habits of the large Central Australian monitor (Varanus giganteus). with a note on the 'fat bodies' of this species. Trans. Proc. Soc. (South Australia) 36: 26-33.
84. Thompson, G.G., Withers, P.C. (1997): Standard and maximal metabolic rates of goannas (Squamata: Varanidae). Physiol. Zool. 70: 307-323.
85. Thompson, G.G., Withers, P.C. Thompson, S.A. (1992): The combat ritual of two monitor lizards. Varanus caudolineatus and Varanus gouldii. W. Aust. Nat. 19: 21-24.
86. Tsellarius, A.Y. (1994): Behaviour and mode of life of the monitor in sand desert. Priroda 5: 26-35.
87. Tsellarius, A.Y., Men'shikov, Y.G. (1994): Indirect communications and its role in the formation of social structure in Varanus griseus (Sauria). Russian J. Herp. 1: 121-132.
88. Tsellarius, A.Y., Tsellarius, E.Y. (1997a): Behaviour of Varanus griseus during encounters with conspecifics. Asiatic Herp. Res. 7: 108-130.
89. Tsellarius, A.Y., Tsellarius, E.Y. (1997b): Thermal conditions of Varanus griseus (Reptilia, Sauria) activity. Zool. Zh. 76: 206-211.
90. Turner, G.F., Huntingford, F.A. (1986): A problem for game theory analysis: assessment and intention in male mouthbrooder contests. Anim. Behav. 34: 961-970.
91. Twigg, L.E. (1988): A note on agonistic behaviour in lace monitors Varanus varius. Herpetofauna 18: 23-25.
92. Vogel, P. (1979): Innerartliche Auseinandersetzungen bei freilebenden Bindenwaranen (Varanus salvator) (Reptilia: Sauria: Varanidae). Salamandra 15: 65-83.
93. Zahavi, A., Zahavi, A. (1997): The Handicap Principle: a Missing Piece of Darwin's Puzzle. New York. Oxford University Press.