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Professor Alex Kacelnik

Prof. Alex Kacelnik
Tel: +44 1865 271164
Fax: +44 1865 271120

I am interested in biological thinking about behaviour. I trained as a zoologist in Argentina, later worked in zoology and psychology departments in Oxford, Groningen and Cambridge, until returning to Oxford in 1990 to set up the present Behavioural Ecology Group. The common thread of my rather diversified research is the attempt to marry theories and experimental work. Most of my work includes modelling, but little is purely or even predominantly theoretical.

Current research topics can be seen in the list of publications and the group's introduction. Ecology, psychology and evolutionary theory are all present both in the selection of research questions and in the nature of the explanations submitted. Members of my group differ in their relative inclinations towards these fields, since some are true muddy-booted field workers while others are white-coated experimentalists or ragged-jeaned theoreticians, but after some time here virtually all of them are stung by the same spirit and acquire a habit of thinking about ecology, psychology and evolution in whatever topic they are dealing with. Although it is less obvious from the publication list, I have a strong interest in philosophical and social implications of ethology, and would engage gladly in any unsolvable argument raised in these areas.

Non-work, I do much the same kind of things, with a strong bias towards watching good cinema, reading good novels, cooking (good food) for people I like, and travelling.





Kacelnik A, El Mouden C (2013). Triumphs and trials of the risk paradigm. Animal Behaviour 86: 1117-1129 doi:10.1016/j.anbehav.2013.09.034

Monteiro T, Vasconcelos M, Kacelnik A (2013). Starlings uphold principles of economic rationality for delay and probability of reward. Proceedings of the Royal Society B 280: 20122386 doi:10.1098/rspb.2012.2386

Gloag, R., Fiorini, V.D., Reboreda, J-C and Kacelnik, A. (2013). The wages of violence: mobbing by mockingbirds as a frontline defence against brood-parasitic cowbirds. Animal Behaviour 86: 1023-1029 doi:10.1016/j.anbehav.2013.09.007

Vasconcelos M, Monteiro T, Kacelnik A (2013). Context-Dependent Preferences in Starlings: Linking Ecology, Foraging and Choice. PLoS ONE 8(5): e64934 doi:10.1016/j.cub.2012.09.002

Auersperg AMI, Kacelnik A, von Bayern AMP. (2013). Explorative learning and functional inferences on a five-step means-end problem in Goffin's cockatoos (Cacatua goffini). PlosOne

Gloag, R. & Kacelnik, A. (2013). Host manipulation via begging call structure in the brood parasitic shiny cowbird. Animal Behaviour86: 101-109

Auersperg, A. M. I., Szabo, B., von Bayern, A. M. P. & Kacelnik, A. (2012). Spontaneous innovation of tool use and flexible manufacture in the Goffin's cockatoo (Cacatua goffini). Current Biology 22(21):R903-904. doi:10.1371/journal.pone.0064934

Gloag, R., Fiorini, F., Reboreda, J-C. & Kacelnik, A. (2012). Brood parasite eggs enhance host egg survival in a multiply parasitized host. Proceedings of the Royal Society B 279(1734): 1831-1839. doi:10.1098/rspb.2011.2047

Gallup, A. C., Hale, J.J., Sumpter, D.J.T., Garnier, S., Kacelnik, A., Krebs, J.R. & Couzin, I.D. (2012). Visual attention and the acquisition of information in human crowds. Proceedings of the National Academy of Sciences USA 109(19): 7245-7250. doi:10.1073/pnas.1116141109

Kacelnik, A. (2012). Putting mechanisms into Behavioral Ecology. In: Evolution and the Mechanisms of Decision Making (ed. P. Hammerstein & J. R. Stevens). Strungmann Forum Report, vol. 11. Cambridge, MA: MIT Press.

Hagen, E.H., Chater, N., Gallistel, C.R., Houston, A. I., Kacelnik, A., Kalenscher, T., Nettle, D., Oppenheimer, D. & Stephens D. W. (2012). Decision making: What can evolution do for us? In: Evolution and the Mechanisms of Decision Making (ed. P. Hammerstein & J. R. Stevens). Strungmann Forum Report, vol. 11. Cambridge, MA: MIT Press.

Vasconcelos, M., Monteiro, T. & Kacelnik, A. (2012). On the flexibility of lizards' cognition: a comment on Leal and Powell (2011). Biology Letters 8(1): 42-43. doi:10.1098/rsbl.2011.0848

Aw, J., Montiero, T., Vasconceles, M. & Kacelnik, A. (2012). Cognitive mechanisms of risky choice:Is there an evaluation cost? Behavioural Processes 89(2): 95-103. doi:10.1016/j.beproc.2011.09.007

Vasconcelos, M., Hollis, K., Nowahari, E. & Kacelnik, A. (2012). Pro-sociality without empathy. Biology Letters 8(6):910-912. doi:10.1098/rsbl.2012.0554

Shapiro, M. S.,Schuck-Paim, C. & Kacelnik, A. (2012). Risk sensitivity for amounts of and delay to rewards: Adaptation for uncertainty or by-product of reward rate maximising? Behavioural Processes 89(2): 104-114. doi:10.1016/j.beproc.2011.08.016

Freidin, E. & Kacelnik, A. (2011). Rational choice, context dependence, and the value of information in European starlings (Sturnus vulgaris). Science 334(6058): 1000-1002. doi:10.1126/science.1209626

Gloag, R., Tuero, D. T., Fiorini, V. D., Reboreda, J-C. & Kacelnik, A. (2011). The economics of nestmate-killing in avian brood parasites: a provisions trade-off. Behavioral Ecology 23(1): 132-140. doi:10.1093/beheco/arr166

Auersperg, A. M. I., von Bayern, A. M. P., Gajdon, G. K., Huber, L. & Kacelnik, A. (2011). Flexibility in problem-solving and tool use of Kea and New Caledonian crows in a multi-access box paradigm. PLOS One e20231 doi:10.1371/journal.pone.0020231

Aw, J. M., Vasconcelos, M., & Kacelnik, A. (2011). How costs affect preferences: Experiments on state-dependence, hedonic state and within-trial contrast in starlings. Animal Behaviour 81(6): 1117-1128. doi:10.1016/j.anbehav.2011.02.015

Kenward, B., Schloegl, C., Rutz, C. Weir, A. S., Bugnyar, T. & Kacelnik, A. (2011). On the evolutionary and ontogenetic origins of tool-oriented behavior in New Caledonian crows (Corvus moneduloides). Biological Journal of the Linnean Society 102(4): 870-877. doi:10.1111/j.1095-8312.2011.01613.x

Wimpenny, J. H., Weir, A. A. S., & Kacelnik, A. (2011). New Caledonian crows use tools for non-foraging activities. Animal Cognition 14(3):459-464. doi:10.1007/s10071-010-0366-1

Kacelnik, A., Vasconcelos, M., Monteiro, T., & Aw, J. (2011). Darwin's Tug-of-War vs. Starlings' Horse-Racing: how adaptations for sequential encounters drive simultaneous choice. Behavioral Ecology and Sociobiology 65(3): 547-558. doi:10.1007/s00265-010-1101-2

Rutz, C., Bluff, L. A., Reed, N., Toscianko,J., Newton, J. Inger, R., Kacelnik, A., Bearhop, S. (2010). The ecological significance of tool use in New Caledonian crows. Science 329(5998): 1523-1526. doi:10.1126/science.1192053

Bluff, L.A., Kacelnik, A., & Rutz, C. (2010). Vocal culture in New Caledonian crows Corvus moneduloides. Biological Journal of the Linnean Society 101: 767-776. doi:10.1111/j.1095-8312.2011.01613.x

Bluff, L.A., Troscianko, J., Weir, A.A.S., Kacelnik, A. & Rutz, C. (2010). Tool use by wild New Caledonian crows Corvus moneduloides at natural foraging sites. Proceedings of the Royal Society B 277: 1377-1385. doi:10.1098/rspb.2009.1953.

Vasconcelos, M., Monteiro, T., Aw, J., & Kacelnik, A. (2010). Choice in multi-alternative environments: A trial-by-trial implementation of the Sequential Choice Model. Behavioural Processes 84: 435-439. doi:10.1016/j.beproc.2009.11.010

Pompilio, L. & Kacelnik, A. (2010). Context-dependent utility overrides absolute memory as a determinant of choice. Proceedings of the National Academy of Sciences USA 107: 508-512. doi:10.1073/pnas.0909468107


von Bayern, A.M.P., Heathcote, R.J.P., Rutz, C., & Kacelnik, A. (2009). The Role of Experience in Problem Solving and Innovative Tool Use in Crows. Current Biology 19(22): 1965-1968. doi:10.1016/j.cub.2009.10.037

Kacelnik, A. (2009) Tools for thought or thoughts for tools? Proceedings of the National Academy of Sciences USA 106: 10071-10072. doi:10.1073/pnas.0904735106

Wimpenny, J.H., Weir, A.A.S., Clayton, L., Rutz, C., & Kacelnik, A. (2009) Cognitive Processes Associated with Sequential Tool Use in New Caledonian Crows. PLoS ONE 4(8): e6471. doi:10.1371/journal.pone.0006471.

Aw, J.M, Holbrook, R.I., Burt de Perera, T., & Kacelnik, A. (2009) State-dependent valuation learning in fish: Banded tetras prefer stimuli associated with greater past deprivation. Behavioural Processes 81: 333-336. doi:10.1016/j.beproc.2008.09.00.

Freidin, E. Aw, J. & Kacelnik, A. (2009) Sequential and simultaneous choices: Testing the diet selection and sequential Choice models. Behavioural Processes 80:218-223.doi:10.1016/j.beproc.2008.12.001

Freidin, E, Cuello, M I, and Kacelnik, A. (2009) Successive negative contrast in a bird: starlingsí behaviour after unpredictable negative changes in food quality. Animal Behaviour 77:857-865. doi:10.1016/j.anbehav.2008.12.010

Shapiro, M. S., Siller, S., & Kacelnik, A. (2008). Simultaneous and Sequential Choice as a Function of Reward Delay and Magnitude: Normative, Descriptive and Process-Based Models Tested in the European Starling (Sturnus vulgaris). Journal of Experimental Psychology: Animal Behavior Processes, 34, 75ñ93. Journal of Experimental Psychology: Animal Behavior 34: 75-93. doi: 10.1037/0097-7403.34.1.75.

Schuck-Paim, C., and Kacelnik, A., (2007) Choice processes in multi-alternative decision making. Behavioral Ecology. Link to online source

139. Rutz, C., Bluff, L.A., Weir, A.A.S., & Kacelnik, A. (2007). Video cameras on wild birds. Science 318: 765. doi 10.1126/science.1146788. Download a [PDF] here.

Bluff, L A, Weir, A A S, Rutz, C, Wimpenny, J H & Kacelnik, A (2007). Tool-related cognition in New Caledonian crows. Comparative Cognition & Behavior Reviews 2: 1-25. Link to online source. Download a [PDF] here.

Kacelnik A., Schuck-Paim C., & Pompilio L., (2006) Inconsistency in animal and human choice. In: Is There Value in Inconsistency?, (Eds: L. Daston and C. Engel) Nomos, pp. 379-396.

Kenward, B, Rutz, C, Weir, A A S & Kacelnik, A (2006). Development of tool use in New Caledonian crows: inherited action patterns and social influence. Animal Behaviour 72: 1329-1343. doi 10.1016/j.anbehav.2006.04.007. Download a [PDF] here.

Weir, A.A.S. & Kacelnik, A. (2006). A New Caledonian crow (Corvus moneduloides) creatively re-designs tools by bending or unbending aluminium strips. Animal Cognition 9: 317-334. doi 10.1007/s10071-006-0052-5. Download a [PDF] here.

Pompilio, L., Kacelnik, A. & Behmer, S.T. (2006). State-dependent learned valuation drives choice in an invertebrate. Science 311 (5767): 1613-1615. doi 10.1126/science.1123924.

Nair-Roberts, R.G., Erichsen, J.T., Reboreda, J.C. & Kacelnik, A. (2006). Distribution of substance P reveals a novel subdivision in the hippocampus of parasitic south American cowbirds. Journal of Comparative Neurology 496 (5): 610-626. DOI 10.1002/cne.20915.

Kacelnik, A., Chappell, J., Weir, A.A.S. & Kenward, B. (2006). Cognitive adaptations for tool-related behaviour in New Caledonian Crows. In: Comparative cognition: experimental explorations of animal intelligence (eds. Wasserman, E.A. & Zentall, T.R.), pp. 515-528. Oxford University Press, Oxford.

Kacelnik, A. (2006). Meanings of rationality. In: Rational Animals? (eds. Nudds, M. & Hurley, S.), pp. 87-106. Oxford University Press, Oxford.

Kacelnik, A. (2006). Feathered friends: review of 'In the Company of Crows and Ravens' by John R. Marzluff and Tony Angell, and 'Crows: Encounters with the Wise Guys of the Avian World' by Candace Savage. Nature 439 (7079): 914-915. DOI 10.1038/439914a.

Fernández-Juricic, E., Smith, R. & Kacelnik, A. (2005). Increasing the costs of conspecific scanning in socially foraging starlings affects vigilance and foraging behaviour. Animal Behaviour 69 (1): 73-81. DOI 10.1016/j.anbehav.2004.01.019.

Kenward, B., Weir, A. A. S., Rutz, C. & Kacelnik, A. (2005). Tool manufacture by naive juvenile crows. Nature 433 (7022): 121. DOI 10.1038/433121a.

Ney-Nifle, M., Bernstein, C., Reboreda, J. C. & Kacelnik, A. (2005). Population dynamics and avian brood parasitism: persistence and invasions in a three-species system. Journal of Animal Ecology 74 (2): 274-284. DOI 10.1111/j.1365-2656.2005.00921.x.

Pompilio, L. & Kacelnik, A. (2005). State-dependent learning and suboptimal choice: when starlings prefer long over short delays to food. Animal Behaviour 70: 571-578. DOI 10.1016/j.anbehav.2004.12.009.

Chappell, J. & Kacelnik, A. (2004). Selection of tool diameter by New Caledonian crows Corvus moneduloides. Animal Cognition 7 (2): 121-127. DOI 10.1007/s10071-003-0202-y.

Fernández-Juricic, E., Erichsen, J. T. & Kacelnik, A. (2004). Visual perception and social foraging in birds. Trends in Ecology & Evolution 19 (1): 25-31. DOI 10.1007/s00265-003-0698-9.

Fernández-Juricic, E. & Kacelnik, A. (2004). Information transfer and gain in flocks: The effects of quality and quantity of social information at different neighbour distances. Behavioral Ecology and Sociobiology 55 (5): 502-511. DOI 10.1007/s00265-003-0698-9.

Fernández-Juricic, E., Siller, S. & Kacelnik, A. (2004). Flock density, social foraging, and scanning: An experiment with starlings. Behavioral Ecology 15 (3): 371-379. DOI 10.1093/beheco/arh017.

Kacelnik, A., Chappell, J., Weir, A. A. S. & Kenward, B. (2004). Tool use and manufacture in birds. In: Encyclopedia of animal behavior (ed. Bekoff, M.), pp. 1067-1069, Volume 3. Greenwood Publishing Group, Westport, CT, US.

Kenward, B., Rutz, C., Weir, A. A. S., Chappell, J. & Kacelnik, A. (2004). Morphology and sexual dimorphism of the New Caledonian Crow Corvus moneduloides, with notes on its behaviour and ecology. Ibis 146 (4): 652-660. DOI 10.1111/j.1474-919x.2004.00299.x.

Marsh, B., Schuck-Paim, C. & Kacelnik, A. (2004). Energetic state during learning affects foraging choices in starlings. Behavioral Ecology 15 (3): 396-399. DOI 10.1093/beheco/arh034.

Schuck-Paim, C., Pompilio, L. & Kacelnik, A. (2004). State-dependent decisions cause apparent violations of rationality in animal choice. PLoS Biology 2 (12): 2305-2315. DOI 10.1371/journal.pbio.0020402.

Weir, A. A. S., Kenward, B., Chappell, J. & Kacelnik, A. (2004). Lateralization of tool use in New Caledonian crows (Corvus moneduloides). Proceedings of the Royal Society of London B (Suppl.) 271 (1548 S5): S344ñS346. DOI 10.1098/rsbl.2004.0183.

Kacelnik, A. (2003). The evolution of patience. In: Time and Decision: Economic and Psychological Perspectives on Intertemporal Choice (eds. Loewenstein, G., Read, D. & Baumeister, R.), pp. 115-138. Russell Sage Foundation, New York.

Lewis, P. A., Miall, R. C., Daan, S. & Kacelnik, A. (2003). Interval timing in mice does not rely upon the circadian pacemaker. Neuroscience Letters 348 (3): 131-134. DOI 10.1016/S0304-3940(03)00521-4.

Scharlemann, J. P. W., Eckel, C. C., Kacelnik, A. & Wilson, R. K. (2003). The Value of a Smile: Game theory with a human face [reprinting of Scharlemann et al., 2001]. In: TRUST (ed. Khalil, E.L.). Critical Studies in Economic Institutions. Edward Elgar, Cheltenham, UK.

Chappell, J. & Kacelnik, A. (2002). Tool selectivity in a non-mammal, the New Caledonian crow (Corvus moneduloides). Animal Cognition 5 (2): 71-78. DOI 10.1007/s10071-002-0130-2.

Kacelnik, A. & Brunner, D. (2002). Timing and foraging: Gibbon's Scalar Expectancy Theory and Optimal Patch Exploitation. Learning and Motivation 33 (1): 177-195. DOI 10.1006/lmot.2001.1110.

Kacelnik, A. & Marsh, B. (2002). Cost can increase preference in starlings. Animal Behaviour 63: 245-250. DOI 10.1006/anbe.2001.1900.

Marsh, B. & Kacelnik, A. (2002). Framing effects and risky decisions in starlings. Proceedings of the National Academy of Sciences 99 (5): 3352-3355. DOI 10.1073/pnas.042491999.

Schuck-Paim, C. & Kacelnik, A. (2002). Rationality in risk-sensitive foraging choices by starlings. Animal Behaviour 64: 869-879. DOI 10.1006/anbe.2003.2003.

Weir, A. A. S., Chappell, J. & Kacelnik, A. (2002). Shaping of hooks in New Caledonian crows. Science 297 (5583): 981. DOI 10.1126/science.1073433.

Bautista, L M, Tinbergen, J & Kacelnik, A (2001). To walk or to fly: how birds choose among foraging modes. Proceedings of the National Academy of Sciences 98 (3): 1089-1094. Link to online source.

Goldstein, D G, Gigerenzer, G, Hogarth, R M, Kacelnik, A, Kareev, Y, Klein, G, Martignon, L, Payne, J W & Schlag, K H (2001). Group Report: why and when do simple heuristics work? In: Bounded Rationality: The Adaptive Toolbox (eds. Gigerenzer, G. & Selten, R.), pp. 173-190. (Report of the 84th Dahlem Workshop, Berlin, March 14-19 1999). MIT Press, Cambridge, Mass.

Kacelnik, A (2001). Instinto y Pulsión. Revista de Psicoanálisis 58: 197-225.

Kacelnik, A., (2001) Review of Pepperberg, I., (1999) The Alex Studies in Ibis 143:688-703.

Rodriguez-Gironés, M A & Kacelnik, A (2001). Relative importance of perceptual and mnemonic variance in human temporal bisection. Quarterly Journal of Experimental Psychology A 54 (2): 527-546. DOI 10.1080/02724980042000264.

Scharlemann, J P W, Eckel, C C, Kacelnik, A & Wilson, R K (2001). The value of a smile: game theory with a human face. Journal of Economic Psychology 22 (5): 617-640. DOI 10.1016/S0167-4870(01)00059-9.

Vásquez, R A & Kacelnik, A (2000). Foraging rate versus sociality in the starling Sturnus vulgaris. Proceedings of the Royal Society of London B 267 (1439): 157-164. Link to online source.


Brito e Abreu, F. & Kacelnik, A. (1999). Energy budgets and risk-sensitive foraging in starlings. Behavioral Ecology 8 (3): 338-345. DOI 10.1093/beheco/10.3.338.

Cotton, P. A., Wright., J & Kacelnik, A. (1999). Chick begging strategies in relation to brood hierarchies and hatching asynchrony. American Naturalist 153 (4): 412-420. DOI 10.1086/303178.

Hauber, M. E., Clayton, N. S., Kacelnik, A., Reboreda, J. C. & DeVoogd, T. J. (1999). Sexual dimorphism and species differences in HVC volumes of cowbirds. Behavioral Neuroscience 113 (5): 1095-1099. Link to online source.

Rodriguez-Gironés, M. A. & Kacelnik, A. (1999). Behavioral adjustment to modifications in the temporal parameters of the environment. Behavioural Processes 45 (1-3): 173-191. DOI 10.1016/S0376-6357(99)00017-0.

Astié, A. A., Kacelnik, A. & Reboreda, J. C. (1998). Sexual differences in memory in shiny cowbirds. Animal Cognition 1: 77-82. DOI 10.1007/s100710050011.

Bateson, M. & Kacelnik, A. (1998). Risk-sensitive foraging: Decision-making in variable environments. In: Cognitive Ecology (ed. Dukas, R.), pp. 297-341. University of Chicago Press, Chicago, IL, US.

Bautista, L. M., Tinbergen, J., Wiersma, P. & Kacelnik, A. (1998). Optimal foraging and beyond: how starlings cope with changes in food availability. American Naturalist 152 (4): 543-561. Link to online source.

Kacelnik, A. (1998). Normative and descriptive models of decision making: time discounting and risk sensitivity [reprinted from Kacelnik, 1997]. In: Rational Models of cognition (eds. Oaksford, M. & Chater, N.), pp. 54-70. Oxford University Press, Oxford.

Kacelnik, A. & Brito e Abreu, F. (1998). Risky choice and Weber's Law. Journal of Theoretical Biology 194: 289-298. DOI 10.1006/jtbi.1998.0763.

Kacelnik, A. & Norris, S. (1998). Signalling via testosterone: Communicating health and vigour. Behavioral and Brain Sciences 21 (3): 378. DOI 10.1017/S0140525X98411221.

Vásquez, R A & Kacelnik, A (1998). Animal foraging: more than met the eye. Trends in Ecology & Evolution 13 (3): 110-111. DOI 10.1016/S0169-5347(97)83353-8.

Bateson, M & Kacelnik, A (1997). Starlings' preferences for predictable and unpredictable delays to food. Animal Behaviour 53: 1129-1142. DOI 10.1006/anbe.1996.0388.

Clayton, N S, Reboreda, J C & Kacelnik, A (1997). Seasonal changes in hippocampus volume in parasitic cowbirds. Behavioural Processes 41: 237-243. DOI 10.1016/S0376-6357(97)00050-8.

Kacelnik, A (1997). Normative and descriptive models of decision making: time discounting and risk sensitivity. In: Characterizing human psychological adaptations (eds. Bock, G.R. & Cardew, G.), pp. 51-70. Ciba Foundation Symposium, Volume 208. Wiley, Chichester.

Kacelnik, A & Bateson, M (1997). Risk-sensitivity: crossroads for theories of decision-making. Trends in Cognitive Sciences 1 (8): 304-309. DOI 10.1016/S1364-6613(97)01093-0.

Kacelnik, A & Krebs, J R (1997). Yanomamö dreams and starling prey loads: The logic of optimality. In: Human nature: a critical reader (ed. Betzig, L.), pp. 21-35. Oxford University Press, Oxford.

Krebs, J. R. & Kacelnik, A. (1997). Risk: a scientific view. In: Science Policy and Risk, pp. 31-43. Discussion Meeting held at the Royal Society, 15 March 1997. The Royal Society, London.

Bateson, M. & Kacelnik, A. (1996). Rate currencies and the foraging starling: The fallacy of the averages revisited. Behavioral Ecology 7 (3): 341-352. Link to online source.

Brunner, D., Kacelnik, A. & Gibbon, J. (1996). Memory for inter-reinforcement interval variability and patch departure decisions in the starling, Sturnus vulgaris. Animal Behaviour 51: 1025-1045. DOI 10.1006/anbe.1996.0105.

Cotton, P. A., Kacelnik, A. & Wright, J. (1996). Chick begging as a signal: are nestlings honest? Behavioral Ecology 7 (2): 178-182. Link to online source.

Davies, N. B., Brooke, M. D. L. & Kacelnik, A. (1996). Recognition errors and probability of parasitism determine whether reed warblers should accept or reject mimetic cuckoo eggs. Proceedings of the Royal Society of London B 263 (1372): 925-931. Link to online source.

Kacelnik, A. (1996). Sexualidade e Biologia [Portugese translation of Kacelnik, 1994]. In: Psicanálise Hoje: Uma revoluçao do olhar (eds. Pellanda, N.M.C. & Pellanda, L.E.C.), pp. 671-689. Vozes, Petropolis.

Kacelnik, A. & Bateson, M. (1996). Risky theories: The effects of variance on foraging decisions. American Zoologist 36 (4): 402-434. Link to online source.

Reboreda, J. C., Clayton, N. S. & Kacelnik, A. (1996). Species and sex differences in hippocampus size in parasitic and non-parasitic cowbirds. NeuroReport 7 (2): 505-508.

Rodriguez-Gironés, M. A., Cotton, P. A. & Kacelnik, A. (1996). The evolution of begging: Signaling and sibling competition. Proceedings of the National Academy of Sciences 93 (25): 14637-14641. Link to online source.

Rodriguez-Gironés, M. A., Drummond, H. & Kacelnik, A. (1996). Effect of food deprivation on dominance status in blue-footed booby (Sula nebouxii) broods. Behavioral Ecology 7 (1): 82-88. Link to online source.

Bateson, M. & Kacelnik, A. (1995). Accuracy of memory for amount in the foraging starling, Sturnus vulgaris. Animal Behaviour 50: 431-443. DOI 10.1006/anbe.1995.0257.

Bateson, M. & Kacelnik, A. (1995). Preferences for fixed and variable food sources: variability in amount and delay. Journal of the Experimental Analysis of Behavior 63 (3): 313-329. DOI 10.1901/jeab.1995.63-313.

Driessen, G., Bernstein, C., Vanalphen, J. J. M. & Kacelnik, A. (1995). A count-down mechanism for host search in the parasitoid Venturia canescens. Journal of Animal Ecology 64 (1): 117-125. Link to online source.

Kacelnik, A. (1995). Sexualidad y Biología [reprinted from Kacelnik, 1994]. Anuario de Sexología [Journal of Asociación Estatal de Profesionales de la Sexología] 1: 99-113.

Kacelnik, A., Cotton, P. A., Stirling, L. & Wright, J. (1995). Food allocation among nestling starlings: Sibling competition and the scope of parental choice. Proceedings of the Royal Society of London B 259 (1356): 259-263. Link to online source.

Rodriguez-Gironés, M. A. & Kacelnik, A. (1995). Interval bisection with and without reference memory. In: Time and the dynamic control of behavior (eds. Richelle, M., DeKeyser, V., d'Ydewalle, G. & Vandierendonck, A.), pp. 47-62. Interuniversity Pole of Attraction, Universite de Liege, Liege, Belgium.

Cassini, M. H. & Kacelnik, A. (1994). Patch choice in guinea pigs: is patch recognition important? Behavioural Processes 31 (2-3): 145-156.

Cuthill, I., Haccou, P. & Kacelnik, A. (1994). Starlings (Sturnus vulgaris) exploiting patches: response to long-term changes in travel time. Behavioral Ecology 5: 81-90. Link to online source.

Kacelnik, A. (1994). Leaf-cutting ants may be optimal foragers - reply. Trends in Ecology & Evolution 9 (2): 63.

Kacelnik, A. (1994). Sexualidad y Biología. Psicoanálisis 16 (2).

Kacelnik, A. & Bernstein, C. (1994). Modelos de optimalidad en etología. In: Etología. Introducción a la Ciencia del Comportamiento (ed. Carranza, J.), pp. 153-180, Volume 8. Universidad de Extremadura, Cáceres.

Cassini, M. H., Lichtenstein, G., Ongay, J. P. & Kacelnik, A. (1993). Foraging behavior in guinea-pigs: further tests of the marginal value theorem. Behavioural Processes 29 (1-2): 99-112.

Kacelnik, A. (1993). Leaf-cutting ants tease optimal foraging theorists. Trends in Ecology & Evolution 8 (10): 346-348.

Reboreda, J. C. & Kacelnik, A. (1993). The role of autoshaping in cooperative two-player games between starlings. Journal of the Experimental Analysis of Behavior 60 (1): 67-83. DOI 10.1901/jeab.1993.60-67.

Todd, I. A. & Kacelnik, A. (1993). Psychological mechanisms and the Marginal Value Theorem: dynamics of scalar memory for travel time. Animal Behaviour 46 (4): 765-775. DOI 10.1006/anbe.1993.1254.

Brunner, D., Kacelnik, A. & Gibbon, J. (1992). Optimal foraging and timing processes in the starling, Sturnus vulgaris: Effect of inter-capture interval. Animal Behaviour 44 (4): 597-613.

Caraco, T., Kacelnik, A., Mesnik, N. & Smulewitz, M. (1992). Short-term rate maximization when rewards and delays covary. Animal Behaviour 44: 441-447.

Kacelnik, A., Bernstein, C. & Krebs, J. R. (1992). Habitat selection and predator prey dynamics - reply. Trends in Ecology & Evolution 7 (9): 313-314.

Kacelnik, A., Bernstein, C. & Krebs, J. R. (1992). The ideal free distribution and predator-prey interactions. Trends in Ecology & Evolution 7: 50-55.

Kacelnik, A. & Todd, I. A. (1992). Psychological mechanisms and the marginal value theorem effect of variability in travel time on patch exploitation. Animal Behaviour 43 (2): 313-322.

Beauchamp, G., Ens, B. J. & Kacelnik, A. (1991). A dynamic model of food allocation to starling Sturnus vulgaris nestlings. Behavioral Ecology 2 (1): 21-37. Link to online source.

Beauchamp, G. & Kacelnik, A. (1991). Effects of the knowledge of partners on learning rates in zebra finches Taeniopygia guttata. Animal Behaviour 41: 247-254. DOI 10.1016/S0003-3472(05)80476-2.

Bernstein, C., Kacelnik, A. & Krebs, J. R. (1991). Individual decisions and the distribution of predators in a patchy environment II: The influence of travel costs and structure of the environment. Journal of Animal Ecology 60 (1): 205-226. Link to online source.

Bernstein, C, Krebs, J R & Kacelnik, A (1991). Distribution of birds among habitats: Theory and practice. In: Bird Population Studies: Relevance to Conservation and Management (eds. Perrins, C.M., Lebreton, J.-D. & Hirons, G.J.M.), pp. 317-345. Oxford University Press.

Krebs, J R & Kacelnik, A (1991). Decision making. In: Behavioural Ecology: An Evolutionary Approach (eds. Krebs, J.R. & Davies, N.B.), pp. 105-137. Blackwell Scientific Publishers, Oxford.

Reboreda, J C & Kacelnik, A (1991). Risk sensitivity in starlings: variability in food amount and food delay. Behavioral Ecology 2 (4): 301-308. Link to online source.

Beauchamp, G & Kacelnik, A (1990). On the fitness functions relating parental care to reproductive value. Journal of Theoretical Biology 146 (4): 513-522.

Cassini, M H, Kacelnik, A & Segura, E T (1990). The tale of the screaming hairy armadillo, the guinea-pig and the marginal value theorem. Animal Behaviour 39: 1030-1050. DOI 10.1016/S0003-3472(05)80776-6.

Cuthill, I & Kacelnik, A (1990). Central place foraging: a reappraisal of the 'loading effect'. Animal Behaviour 40: 1087-1101. DOI 10.1016/S0003-3472(05)80175-7.

Cuthill, I., Kacelnik, A., Krebs, J. R., Haccou, P. & Iwasa, Y. (1990). Starlings exploiting patches: the effect of recent experience on foraging decisions. Animal Behaviour 40: 625-640. DOI 10.1016/S0003-3472(05)80692-X.

Kacelnik, A., Brunner, D. & Gibbon, J. (1990). Timing mechanisms in optimal foraging: some applications of Scalar Expectancy Theory. In: Behavioural Mechanisms of Food Selection (ed. Hughes, R.N.), pp. 61-82. NATO ASI Series G: Ecological Sciences, Volume 20. Springer-Verlag, Heidelberg, NY.

Kacelnik, A. & Cuthill, I. (1990). Central place foraging in starlings Sturnus vulgaris II: Food allocation to chicks. Journal of Animal Ecology 59 (2): 655-674. Link to online source.

Reboreda, J. C. & Kacelnik, A. (1990). On cooperation, tit-for-tat and mirrors. Animal Behaviour 40: 1188-1189. DOI 10.1016/S0003-3472(05)80188-5.


Kacelnik, A., (1989) Review of Mangel, M., & Clark, C.W., (1989) Dynamic modelling in behavioral ecology, Princeton University Press appeared in Nature, 339: 674.

Kacelnik, A., (1989) Review of: Holland, J.H., Holyoak, J., Nisbett, R.E.,& Thagard, P.R., (1986) Induction: processes of inference, learning and discovery, MIT Press, in Ethology, 80: 347-348.

Bernstein, C., Kacelnik, A. & Krebs, J. R. (1988). Individual decisions and the distribution of predators in a patchy environment. Journal of Animal Ecology 57 (3): 1007-1026. Link to online source.

Gibbon, J., Church, R. M., Fairhurst, S. & Kacelnik, A. (1988). Scalar Expectancy Theory and choice between delayed rewards. Psychological Review 95: 102-114. Link to online source.

Houston, A., Schmid-Hempel, P. & Kacelnik, A. (1988). Foraging strategy, worker mortality, and the growth of the colony in social insects. American Naturalist 131 (1): 107-114. Link to online source.

Kacelnik, A. (1988). Short-term adjustments of parental effort in starlings. In: Acta XIX Congressus Internationalis Ornithologici, Vol II (ed. Ouellet, H.), pp. 1843-1856. University of Ottawa Press.

Kacelnik, A. & Bernstein, C. (1988). Optimal foraging and arbitrary food distributions: patch models gain a lease of life. Trends in Ecology & Evolution 3 (10): 251-253.

Commons, M. L., Kacelnik, A. & Shettleworth, S. J. (Eds). (1987). Harvard Symposium on Quantitative Analyses of Behaviour, Volume VI: Foraging. Lawrence Erlbaum and Associates.

Kacelnik, A. (1987). Information primacy or preference for familiar foraging techniques? A critique of Inglis & Ferguson. Animal Behaviour 35 (3): 925-926. DOI 10.1016/S0003-3472(87)80130-6.

Kacelnik, A. & Bloem, G (1987). Optimalisatie van Fourageergedrag bij Spreeuwen. Vakblad voor Bioloogen (The Netherlands) 67: 117-121.

Kacelnik, A. & Cuthill, I. C. (1987). Starlings and optimal foraging theory: modelling in a fractal world. In: Foraging Theory (eds. Kamil, A.C., Krebs, J.R. & Pulliam, H.R.), pp. 303-333. Plenum Press, New York.

Kacelnik, A., Krebs, J. R. & Ens, B. (1987). Foraging in a changing environment: an experiment with starlings (Sturnus vulgaris). In: Quantitative analyses of behavior, Vol. VI: Foraging (eds. Commons, M.L., Kacelnik, A. & Shettleworth, S.J.), pp. 63-87. Lawrence Erlbaum and Associates, Hillsdale, NJ.

Hunter, M. L., Kacelnik, A., Roberts, J. & Vuillermoz, M. (1986). Directionality of avian vocalizations: a laboratory study. Condor 88 (3): 371-375. Link to online source.

Kacelnik, A., Houston, A. I. & Schmidt-Hempel, P. (1986). Central place foraging in honey bees: the effect of travel time and nectar flow on crop filling. Behavioral Ecology and Sociobiology 19: 19-24. DOI 10.1007/BF00303838.

Cheverton, J., Kacelnik, A. & Krebs, J. R. (1985). Optimal foraging: constraints and currencies. Fortschritte Der Zoologie 31: 109-126.

Kacelnik, A. & Krebs, J. R. (1985). Learning to exploit patchily distributed food. In: Behavioural ecology: ecological implications of adaptive behaviour (eds. Sibly, R.M. & Smith, R.), pp. 189-206. British Ecological Society Symposium. Blackwell Scientific Publications, Oxford.

Kacelnik, A. & Krebs, J. R. (1985). Rate of reinforcement matters in optimal foraging theory. Behavioral and Brain Sciences 8 (2): 340-340.

Schmid-Hempel, P., Kacelnik, A. & Houston, A. I. (1985). Honeybees maximize efficiency by not filling their crop. Behavioral Ecology and Sociobiology 17: 61-66. DOI 10.1007/BF00299430.

Thomas, G., Kacelnik, A. & Vandermeulen, J. (1985). The three-spined stickleback and the two-armed bandit. Behaviour 93: 227-240.

Hunter, M. L., Kacelnik, A., Roberts, J. & Vuillermoz, M. (1984). Directionality of avian vocalizations: a laboratory study. Proceedings of the Institute of Acoustics: 177-184.

Kacelnik, A. (1984). Central place foraging in starlings (Sturnus vulgaris) I: Patch residence time. Journal of Animal Ecology 53: 283-299. Link to online source.

Kacelnik, A. & Houston, A. (1984). The use of evolutionary analogies and the rejection of state variables by B F Skinner. Behavioral and Brain Sciences 7 (4): 691-692.

Kacelnik, A. & Houston, A. I. (1984). Some effects of energy costs on foraging strategies. Animal Behaviour 32: 609-614. DOI 10.1016/S0003-3472(84)80298-5.

Krebs, J. R. & Kacelnik, A. (1984). Evolutionarily stable learning strategies. A comment on Maynard-Smith. Behavioral and Brain Sciences 7: 109-110.

Krebs, J. R. & Kacelnik, A. (1984). Time horizons of foraging animals. In: Timing and Time Perception (eds. Gibbon, J. & Allan, L.), pp. 278-291. Annals of the New York Academy of Sciences, Volume 423.

Kacelnik, A. & Krebs, J. R. (1983). The dawn chorus in the great tit (Parus major): proximate and ultimate causes. Behaviour 83: 287-309.

Kacelnik, A. (1984) Review of: Aspy, W.P., & Lustick, S., (1983) Behavioral Energetics: the cost of survival in vertebrates. Ohio State University Press appeared in Ibis 126(2): 261.

Kacelnik, A. (1985) Review of: Escalante, R. (1983) Catalogo de las Aves del Uruguayin, Intendencia Municipal de Montevideo appeared in Ibis, 127: 126.

Houston, A. I., Kacelnik, A. & McNamara, J. (1982). Some learning rules for acquiring information. In: Functional Ontogeny (ed. McFarland, D.J.), pp. 140-191. Pitman, Boston.

Kacelnik, A. (1982). Interacción etología - psicoanálisis: el punto de visita etológico. Revista Argentina de Psicología 13 (32): 9-57.

Kacelnik, A., Houston, A. I. & Krebs, J. R. (1981). Optimal foraging and territorial defense in the great tit (Parus major). Behavioral Ecology and Sociobiology 8 (1): 35-40. DOI 10.1007/BF00302841.

Roberts, J. P., Hunter, M. L. & Kacelnik, A. (1981). The ground effect and acoustic communication - reply. Animal Behaviour 29: 633-634.

Roberts, J. P., Kacelnik, A. & Hunter, M. L. (1980). Some consequences of sound interference patterns of bird song. Acoustic Letters 3: 141-146.


Kacelnik, A. (1979). The foraging efficiency of great tits (Parus major L) in relation to light intensity. Animal Behaviour 27: 237-242.

Roberts, J. P., Kacelnik, A. & Hunter, M. L. (1979). A model of sound interference in relation to acoustic communication. Animal Behaviour 27: 1271-1273.

Krebs, J. R., Kacelnik, A. & Taylor, P. (1978). Tests of optimal sampling by foraging great tits. Nature 275: 27-31. DOI 10.1038/275027a0.

Segura, E. T. & Kacelnik, A. (1977). Cardiorespiratory and electroencephalographic responses to stimulation of mesencephalic tegmentum in toads, lizards, and rats. Experimental Neurology 57 (2): 364-373.

Kacelnik, A. & Segura, E. T. (1975). Effects of sodium pentobarbital upon cardiovascular-responses to mesencephalic reticular stimulation in rats. Acta Physiologica Latinoamericana 25 (4): 332-338.

Kacelnik, A. & Segura, E. T. (1974). Respuestas cardiorespiratorias y electroencefalograficas a la estimulacion de la superficie de los hemisferios cerebrales en Tupinambis rufescens (Teiidae, squamata). Revista de la Sociedad Argentina de Biologia 50: 62-73.

Segura, E. T., Dejuan, A. O. R., Colombo, J. A. & Kacelnik, A (1971). Sexual clasp as a reticularly controlled nehavior in toad, Bufo arenarum Hensel. Physiology & Behavior 7 (2): 157-160.

Segura, E. T., de Juan, A. O. R., Kacelnik, A. & Albalustri, L. (1971). Comparacion de ciertas respuestas autonomicas 'estado dependientes' en Bufo arenarum Hensel and Rattus norvegicus. Revista de la Sociedad Argentina de Biologia 47: 193-206.