Sequences Through The Mist

All animals are social, even if only to the extent of engaging in sexual reproduction and otherwise avoiding each other. Understanding the rules that govern social dynamics in general is a step to understanding our own lives (Axelrod 1984). Everything happens in context, so it is unsurprising that we advocate a contextual approach to studying society. Social relationships are the consequences of entwining sets of events and states, and our aim has been to explore how these may be disentangled through analyzing sequences of changes in time and space. Although some of these analytical approaches may seem daunting through their unfamiliarity, they concern a phenomenon— sequentiality—that not only is at the root of life in the patterning of nucleic acids, but also suffuses every aspect of our daily experience in strings of words and sentences and stories. One story can be told in different ways for different purposes, just as an individual butterfly is expressed differently as larva, pupa, and imago. A metamorphosis in storytelling shows how the perceptions of Robert Henryson, a fifteenth-century Scottish poet, might, for the purposes of measuring the dynamics of mammalian society, be stripped to their essentials. In his Moral Fables, Henryson penned these words:

This country mouse lay flattened on the ground, fearing every minute that she would be killed; for her heart was pounding with strokes of fear, and feverishly she trembled hand and foot. And when her sister found her in such straits, for the sake of pity first she grieved a bit, then comforted her with words as sweet as honey.

It is common for stories to be peppered with anthropomorphism, and one class of these, akin to mock anthropomorphism (Kennedy 1986), enriches prose with insight. Certainly, understanding is not the sole prerogative of those who wield the colorless pen of late-twentieth-century science. On the other hand, another class of anthropomorphism is blind to life's processes and, in its fancifulness, more often than not corrupts literature by diminishing understanding. Stripping out these distractions is a step toward understanding, and interpretation of the essence that remains hinges on the meaning of sequences, whether they be words or behaviors. In the contextual study of animal behavior the narration that precedes or succeeds a given action has equal weight to the action itself. As a step toward such contextual analysis, we might wonder whether Henryson had this in mind:

One rural female wood mouse (Apodemus sylvaticus) adopted a dorsoven-trally flattened posture, exhibiting, in an unspecified but seemingly fearful context, tachycardia and elevated body temperature, and associated tremulous movements of the fore- and hind-paws. These signs appeared to be stress-induced. A conspecific female approached and initiated prolonged nonreciprocal grooming, perhaps with the consequence of diminishing the signs of putative stress. Subsequent molecular analyses confirmed the hypothesis that these individuals were siblings.

The only merit of the aridity of this version is that it exposes the states and events that can usefully be defined in an ethogram and exposes the sequences of those that will become grist for analysis:

(I) ETHOGRAM: crouch: dorsoventral flattened posture with head close to the (i.e., modifier) ground; shake: tremulous movement of body (no modifier), or shaking fore-paw(s) (modifier 1) or shaking hind-paw (modifier 2); approach: one individual moves toward another; groom opposite: one individual grooms the other on the head, or neck, or back, or flank (modifier 1).

(II) SEQUENCE: BEHAVIOR: A-crouch ^ A-shake ^ B-approach-A ^ B-nose-to-nose-A ^ B-groom-A.

MODIFIER 1: ground paw

MODIFIER 2: foot

DURATION: (tj - t0) (t2 - tj) (t3 - t2) (t4 - t3) (tn - t4) SUBJECTS: 2 female wood mice (Apodemus sylvaticus)

Clearly, there are many ways of describing an observation. Henryson's original and our coded version may be quests for different sorts of understanding. The point is that having designed an ethogram and having chosen the correct method of observation, we can distill an observed set of events or states into a pattern that facilitates greater precision of interpretation and decreased risk of fanciful anthropomorphism.


We are grateful to M. Dawkins, R. Dunbar, D. Johnson, and two anonymous reviewers for comments on an earlier draft. We also thank Luigi Boitani and Todd Fuller for their invitation to participate in the conference that spawned this book and for their forbearance as editors.

Literature Cited

Albonetti, M. E., J. Lazarus, D. W. Dickins, A. Whybrow, R. Schonheiter, J. Prenter, R. Elwood, J. D. Newman, F. O. Odberg, L. Kaiser, M. H. Pham-Delegue, V. Verguelen, J. C. Fentress, and M. L. Bocia. 1992. Software multiple review. The Observer: Software for behavioural research, version 2.0. Ethology, Ecology and Evolution 4: 401—414. Altmann, J. 1974. Observational study of behaviour: Sampling methods. Behaviour 49: 227-267.

Appleby, M. C. 1983. The probability of linearity in hierarchies. Animal Behaviour 32: 600-608.

Arnoldmeeks, C. and J. J. McGlone. 1986. Validating techniques to sample behavior of confined, young-pigs. Applied Animal Behaviour Science 16: 149-155. Axelrod, R. 1984. The evolution of co-operation. New York: Basic Books. Axelrod, R. and W. D. Hamilton. 1981. The evolution of cooperation. Science 211: 1390-1396.

Barbault, R. 1988. Body size, ecological constraints, and the evolution of life-history strategies. In M. K. Hecht, B. Wallace, and G. T. Prance, eds., Evolutionary biology, 261-285. New York: Plenum.

Bekoff, M. 1977. Quantitative studies of three areas of classical ethology: Social dominance, behavioural taxonomy, and behavioural variability. In B. Hazlett, ed., Quantitative methods in the study of animal behavior, 1-46. New York: Academic Press. Bekoff, M. and D. Jamieson. 1997. Ethics and the study of carnivores: Doing science while respecting animals. In J. L. Gittleman, ed., Carnivore behavior, ecology and evolution, vol. 2, 15-45. Ithaca, N.Y.: Cornell University Press. Berdoy, M., P. Smith, and D. W. Macdonald. 1994. Stability of social status in wild rats:

Age and the role of settled dominance. Behaviour 132: 193-212. Berger, J. and C. Cunningham. 1995. Predation, sensitivity and sex: Why female black rhinoceroses outlive males. Behavioural Ecology 6: 57-64. Bernstein, I. S. 1991. An empirical comparison of focal and ad-libitum scoring with commentary on instantaneous scans, all occurrence and one-zero techniques. Animal Behaviour 42: 721-728.

Borries, C. 1992. Grooming site preferences in female langurs (Presbytis entellus). International Journal of Primatology 13: 19—32.

Bovet, J. 1972a. On the social behaviour in a stable group of long-tailed field mice (Apodemus sylvaticus). I. An interpretation of defensive postures. Behaviour 41: 43—54.

Bovet, J. 1972b. On the social behaviour in a stable group of long-tailed field mice (Apodemus sylvaticus). II. Its relations with distribution of daily activity. Behaviour 41: 55—67.

Bressers, M., E. Meelis, P. Haccou, and M. Krok. 1991. When did it really start or stop: The impact of censored observations on the analysis of durations. Behavioural Processes 23: 1-20.

Brown, J. L. 1975. The evolution of behaviour. New York: Norton.

Burley, N. T. 1988. Wild zebra finches have band colour preferences. Animal Behaviour 36: 1235-1237.

Cairns, S. J. and S. J. Schwager. 1987. A comparison of association indices. Animal Behaviour 35: 1454-1469.

Chase, I. D. 1982. Dynamics of hierarchy formation: The sequential development of dominance relationships. Behaviour 80: 218-240.

Cheney, D. L. 1992. Intragroup cohesion and intergroup hostility: the relation between grooming distributions and intergroup competition among female primates. Behav-

iouralEcology 3: 334-345.

Clout, M. N. and M. G. Efford. 1984. Sex differences in the dispersal and settlement of brushtail possums (Trichurus vulpecula). Journal of Animal Ecology 53: 737-749.

Clutton-Brock, T. H., F. E. Guinness, and S. D. Albon. 1982. Red deer: Behaviour and ecology of two sexes. Chicago: University of Chicago Press.

Clutton-Brock, T. H. and G. A. Parker. 1995. Punishment in animal societies. Nature 373: 209-216.

Colgan, P. 1978. Quantitative ethology. New York: Wiley.

Creel, S. 1997. Cooperative hunting and group size: Assumptions and currencies. Animal Behaviour 54: 1319-1324.

Creel, S. and D. W. Macdonald. 1996. Sociality, group size, and reproductive suppression among carnivores. Advances in the Study of Behaviors 24: 203-257.

Creel, S., S. L. Monfort, D. E. Wildt, and P. M. Waser. 1991. Spontaneous lactation is an adaptive result of pseudopregnancy. Nature 351: 660-662.

Cuthill, I. 1991. Field experiments in animal behaviour: Methods and ethics. Animal Behaviour 42: 1007-1014.

Dawkins, R. 1976. The selfish gene. Oxford, U.K.: Oxford University Press.

Dawkins, R. 1989 (2d ed.). The selfish gene. Oxford, U.K.: Oxford University Press.

Doncaster, C. P. 1990. Non-parametric estimates of interaction from radio-tracking data. Journal of Theoretical Biology 143: 431-443.

Drews, C. 1993. The concept and definition of dominance in animal behaviour. Behaviour 125: 283-313.

Dugatkin, L. A. 1997. Cooperation among animals: An evolutionary perspective. Oxford, U.K.: Oxford University Press.

Dunbar, R. I. M. 1976. Some aspects of research design and their implications in the observational study of behaviour. Behaviour 58: 78—98.

Dunbar, R. 1988. Primate social systems. London: Croom Helm.

Evans, P. G. H., D. W. Macdonald, and C. L. Cheeseman. 1989. Social structure of the Eurasian badger (Meles meles): Genetic evidence. Journal of Zoology 218: 587—595.

Fagen, R. M. 1978. Information measures: Statistical confidence limits and inference. Journal of Theoretical Biology 73: 61—79.

Fagen, R. M. and R. N. Goldman. 1977. Behavioural catalogue analysis methods. Animal Behaviour 25: 261-274.

Faver, J., M. Mendl, and P. Bateson. 1986. A method for rating individual distinctiveness of domestic cats. Animal Behaviour 34: 1016-1025.

Findlay, C. S., R. I. C. Hansell, and C. J. Lumsden. 1989. Behavioural evolution and bio-cultural games: Oblique and horizontal cultural transmission. Journal of Theoretical

Biology 137: 245-269.

Geffen, E., M. E. Gompper, J. L. Gittleman, H. K. Luh, D. W. Macdonald, and R. K. Wayne. 1996. Size, life-history traits, and social organization in the Canidae: A reevaluation. American Naturalist 147: 140-160.

Geyer, L. A. and C. A. Kornet. 1982. Auto- and allo-grooming in pine voles (Microtus pine-torum) and meadow voles (Microtus pennsylvanicus). Physiology and Behavior 28: 409412.

Ginsberg, J. R. and T. P. Young. 1992. Measuring association between individuals or groups in behavioural studies. Animal Behaviour 44: 377-379.

Golani, I. 1976. Homeostatic motor processes in mammalian interactions: a choreography of display. In P. P. G. Bateson and P. H. Klopfer, eds., Perspectives in ethology, vol. 2, 69-134. New York: Plenum.

Golani, I. 1992. A mobility gradient in the organisation of vertebrate movement: The perception of movement through symbolic movement. Behavioural and Brain Sciences 15: 249-266.

Goosen, C. 1987. Social grooming in primates. In G. Mitchell and J. Erwin, eds., Comparative primate biology, Vol. 2: Behaviour, cognition, and motivation, 107-131. New York: Alan R. Liss.

Haccou, P. and E. Meelis. 1992. Statistical analysis of behavioural data. Oxford, U.K.: Oxford University Press.

Haccou, P. and E. Meelis. 1995 (2d ed.). Statistical analysis of behavioural data. Oxford, U.K.: Oxford University Press.

Hailman, J. P. and J. J. I. Dzelzkalns. 1974. Mallard tail-wagging: Punctuation for animal communication? American Naturalist 108: 236-238.

Hailman, E. D. and J. P. Hailman. 1993. UNCERT software (beta version) user's guide. ([email protected]).

Hansson, L. 1992. Fitness and life history correlates of weight variations in small mammals. Oikos 64: 479-484.

Hansson, M. B., L. J. Bledsoe, B. C. Kirkevold, C. J., Casson, and J. W. Nightingale. 1993.

Behavioural budgets of captive sea otter mother-pup pairs during pup development.

Zoo Biology 12: 459-477.

Hare, J. F. 1992. Colony member discrimination by juvenile Columbian ground squirrels (Spermophilus columbianus). Ethology 92: 301-315.

Harris, S. and P. C. L. White. 1992. Is reduced affiliative rather than increased agonistic behaviour associated with dispersal in red foxes? Animal Behaviour 44: 1085-1089.

Hazlett, B. A. 1977. Quantitative methods in the study of animal behaviour. New York: Academic Press.

Hemelrijk, C. K. 1990. Models and tests of reciprocity, unidirectionality and other social interaction patterns at group level. Animal Behaviour 39: 1013-1029.

Hemelrijk, C. K. and A. Ek. 1991. Reciprocity and interchange of grooming and support in captive chimpanzees. Animal Behaviour 41: 923-935.

Herrera, E. A. and D. W. Macdonald. 1989. Resource utilisation and territoriality in group-living capybaras (Hydrochoerus hydrochaeris). Journal of Animal Ecology 58: 667-679.

Herrera, E. A. and D. W. Macdonald. 1993. Aggression, dominance, and mating success among capybara males (Hydrochoerus hydrochaeris). Behavioural Ecology 4: 114-119.

Hinde, R. A. H. 1981. Social relationships. In D. McFarland, ed., The Oxford companion to animal behaviour, 521-527. Oxford, U.K.: Oxford University Press.

Hinde, R. A., ed. 1983. Primate social relationships. An integrated approach. Oxford, U.K.: Blackwell Scientific.

Hofer, H. and M. East. 1993. Loud calling in a female-dominated mammalian society. II Behavioural contexts and functions of whooping of spotted hyaenas, Crocuta crocuta. Animal Behaviour 42: 651-669.

Hoogland, J. L. 1995. The black tailed prairie dog: Social life of a burrowing mammal. Chicago: University of Chicago Press.

Iacobucci, D. 1990. Derivation of subgroups from dyadic interactions. Psychological Bulletin 107: 114-132.

Johnsingh, A. J. T. 1983. Large mammalian prey-predators in Bandipur. Journal of the Bombay Natural History Society 80: 1-53.

Jouventin, P. and A. Cornet. 1980. The sociobiology of pinnipeds. Advances in the Study of Behavior 11: 121-141.

Kawanaka, K. 1996. Observation time and sampling intervals for measuring behavior and interactions of chimpanzees in the wild. Primates 37: 185-196.

Kennedy, J. S. 1986. Some current issues in orientation to odour sources. In T. L. Payne, M. C. Birch, and C. E. J. Kennedy, eds., Mechanisms in insect olfaction, 11-25. Oxford, U.K.: Clarendon.

Kenward, R. 1987. Wildlife radio-tagging. London: Academic Press.

Kerby, G. and D. W. Macdonald. 1988. Cat society and the consequences of colony size. In D. Turner and P. Bateson, eds., The domestic cat: The biology of its behaviour, 67-81. Cambridge, U.K.: Cambridge University Press.

Kraft, B., C. Colin, and P. Peignot. 1994. Diving-for-food: A new model to assess social roles in a group of laboratory rats. Ethology 96: 11-23.

Krebs, J. R., and N. B. Davies. 1991 (3d ed.). Behavioural ecology. Oxford, U.K.: Blackwell Scientific.

Krishnaiah, P. R. and T. J. Rao. 1988. The handbook of statistics. Amsterdam: Elsevier.

Kruuk, H. 1972. The spotted hyena. Chicago: Chicago University Press.

Kruuk, H. 1978. Foraging and spatial organisation of the badger (Meles meles). Behavioural Ecology andSociobiology 4: 75—89.

Langton, S. D., D. Collet, and M. R. Sibly. 1995. Splitting behaviour into bouts; a maximum likelihood approach. Behaviour 132: 9—10.

Ledger, D. W. and I. A. Didrichsons. 1994. An assessment of data pooling and some alternatives. Animal Behaviour 48: 823—832.

Lehner, P. N. 1996 (2d ed.). Handbook ofethologicalmethods. Cambridge, U.K.: Cambridge University Press.

Lindgren, B. W. 1968. Statistical theory. London: Macmillan.

Lindquist, T. and I. S. Bernstein. 1987. A feminist perspective on dominance relationships in primates. American Journal of Primatology 12: 355.

Lindstedt, S. L., B. J. Miller, and S. W. Buskirk. 1986. Home range, time, and body size in mammals. Ecology 67: 413-418.

Lorenz, K. 1981. The foundations of ethology. New York: Springer-Verlag.

Lott, D. F. 1991. Intraspecific variation in the social systems of wild vertebrates. Cambridge, U.K.: Cambridge University Press.

Macdonald, D. W. 1983. The ecology of carnivore social behaviour. Nature 301: 379-384.

Macdonald, D. W. 1981. Dwindling resources and the social behaviour of capybaras (Hydrochoerus hydrochaeris). Journal of Zoology 194: 371-391.

Macdonald, D. W. 1995. Wildlife rabies: The implications for Britain. Unresolved questions for the control of wildlife rabies: Social perturbation and interspecific interactions. In Rabies in a changing world, 33-48. Cheltenham: British Small Animal Veterinary Association.

Macdonald, D. W. and C. J. Amlaner, Jr. 1980. A practical guide to radio tracking. In C. J. Amlaner and D. W. Macdonald, eds., A handbook on biotelemetry and radiotracking, 143-159. Oxford, U.K.: Pergamon.

Macdonald, D. W., P. J. Apps, G. Carr, and G. Kerby. 1987. Social behaviour, nursing coalitions and infanticide in a colony of farm cats. Advances in Ethology 28: 1-66.

Macdonald, D. W., F. G. Ball, and N. G. Hough. 1980. The evaluation of home range size and configuration using radio-tracking data. In D. W. Macdonald and C. J. Amlaner, eds., A handbook on biotelemetry and radio tracking, 143-159, 405-426. Oxford, U.K.: Pergamon.

Macdonald, D. W. and G. M. Carr. 1989. Food security and the rewards of tolerance. In V. Standen and R. A. Foley, eds., Comparative socioecology: The behavioural ecology of humans and other mammals, 75-99. Oxford, U.K.: Blackwell Scientific.

Macdonald, D. W. and O. Courtenay. 1996. Enduring social relationships in a population of crab-eating zorros, Cerdocyon thous, in Amazonian Brazil. Journal of Zoology 239: 329-355.

Macdonald, D. W., N. Yamaguchi, and G. Kerby. In press (2d ed.). Group-living in the domestic cat: Its sociobiology and epidemiology. In D. Turner and P. Bateson, eds., The domestic cat: The biology of its behaviour. Cambridge, U.K.: Cambridge University Press.

Machlis, L. 1977. An analysis of the temporal patterning of pecking in chicks. Behaviour 63: 1-70.

Machlis, L., P. W. D. Dood, and J. C. Fentress. 1985. The pooling fallacy: Problems arising when individuals contribute more than one observation to the data set. Zeitschrift für Tierpsychologie 68: 201-214.

Martin, P. and P. Bateson, eds. 1993 (2d ed.). Measuring behaviour: An introductory guide. Cambridge, U.K.: Cambridge University Press.

Matloff, N. S. 1991. Statistical hypothesis testing: Problems and alternatives. Environmental Entomology 20: 1246-1250.

May, R. M. 1975. Patterns of species abundance and diversity. In M. L. Cody and J. Diamond., eds., Ecology and evolution of communities, 81-120. Cambridge, Mass.: Harvard University Press.

McFarland, D. J. 1981. The Oxford companion to animal behaviour. Oxford, U.K.: Oxford University Press.

McKnight, B. L. 1995. Behavioural ecology of hand reared elephants (Loxodonta africana) in Tsavo East National Park, Kenya. African Journal of Ecology 33: 242-256.

Milinski, M. and G. A. Parker. 1991 (3d ed.). Competition for Resources. In J. R. Krebs and N. B. Davies, eds., Behavioural ecology: An evolutionary approach, 137-168. Oxford, U.K.: Blackwell Scientific.

Mitchell, B. W., N. S. Thompson, and H. L. Miles, eds. 1997. Anthropomorphism, anecdotes and animals. New York: State University of New York Press.

Moehrenschlager, A., D. W. Macdonald, and C. Moehrenschlager. In press. The effects of trapping and handling swift foxes. In L. Carbyn and M. Sovoda, eds., Swift fox biology. Edmonton, Alberta: Canadian Circumpolar Institute.

Morell, V. 1993. Primatology: Seeing nature through the lens of gender. Science 260: 428-429.

Moser, R., M. Cords, and H. Kummer. 1991. Social influences on grooming site preferences among captive long-tailed macaques. International Journal of Primatology 12: 217-230.

Noe, R., C. P. van Schaik, and J. A. R. A. M. van Hoof. 1991. The market effect: An explanation for pay-off asymmetries among collaborating animals. Ethology 87: 97118.

Noldus Information Technology. 1994. The Observer, support package for video tape analysis. Reference manual, Version 3.0. Wageningen, the Netherlands: Noldus Information Technology.

Noldus Information Technology. 1995. The Observer, base package for Windows. Reference manual, Version 3.0. Wageningen, the Netherlands: Noldus Information Technology.

Odagiri, K. and Y. Matsuzawa 1994. Application of an instantaneous sampling method to lamb's play behaviour. Japanese Journal of Animal Psychology 43: 9-16.

Packer, C. and T. Caro. 1997. Foraging costs in social carnivores. Animal Behaviour 54: 1317-1324.

Pearl, M. C. and S. R. Schulman. 1983. Techniques for the analysis of social structure in animal societies. Advances in the Study of Behavior 13: 107-146.

Plesner Jensen, S. 1993. Temporal changes in food preferences of wood mice (Apodemus sylvaticus L.). Oecologia 94: 76-82.

Pollock, G. B. 1994. Social competition or correlated strategy? Evolutionary Ecology 8: 221-229.

Poysa, H. 1991. Measuring time budgets with instantaneous sampling: A cautionary note. Animal Behaviour 42: 317-318.

Quera, V. 1990. A generalised technique to estimate frequency and duration in time sampling. Behavioral Assessment 12: 409-424.

Roberts, B. A. 1992. Sequential data in biological experiments. An introduction for research workers. London: Chapman & Hall.

Rogers, L. M., C. L. Chesseman, and S. Langton. 1997. Body weight as an indication of density-dependent population regulation in badgers (Meles meles) at Woodchester Park, Gloucestershire. Journal of Zoology 242: 597-604.

Roper, T. J. 1992. Badger (Meles meles) sett architecture, internal environment and function. Mammal Review 22: 43-53.

Rowell, T. E. and C. A. Rowell. 1993. The social organization of feral Ovis aries ram groups in the pre-rut period. Ethology 95: 213-232.

Rubenstein, D. I. 1993. The ecology of female social behaviour in horses, zebras and asses. Physiology and Ecology Japan 29: 13-28.

Schino, G., F. Aureli, and A. Troisi. 1988. Equivalence between measures of allogrooming: an empirical-comparison in 3 species of macaques. Folia Primatologica 51: 214-219.

Schleidt, W. M., G. Yakalis, M. Donnelly, and J. McGarry. 1984. A proposal for a standard ethogram, exemplified by an ethogram of the bluebreasted quail (Coturnix chinensis). Zeitschrift für Tierpsychologie 64: 193-220.

Seyfarth, R. M. 1983. Grooming and social competition in primates. In R. A. Hinde, ed., Primate social relationships, 182-190. Oxford, U.K.: Blackwell Scientific.

Sharman, M. and R. I. M. Dunbar. 1982. Observer bias in selection of study group in baboon field studies. Primates 23: 567-573.

Shrader-Frachette, K. S. and E. D. McCoy. 1992. Statistics, costs and rationality in ecological inference. Trends in Ecology and Evolution 7: 96-99.

Sibley, R. M., H. M. R. Nott, and D. J. Fletcher. 1990. Splitting behaviour into bouts. Animal Behaviour 39: 63-69.

Sillero-Zubiri, C., P. J. Johnson, and D. W. Macdonald. 1998. A hypothesis for breeding synchrony in Ethiopian wolves (Canis simensis) Journal of Mammalogy 79: 853-858.

Smuts, B. 1985. Sex and friendship in baboons. New York: Aldine.

Stevenson-Hinde, J. 1983. individual characteristics and the social situation. In R. A. Hinde, ed., Primate social relationships, 28-35. Oxford, U.K.: Blackwell Scientific.

Stewart, P. D. 1997. The social behaviour of the European badger (Meles meles). Doctoral thesis. Oxford, U.K.: Oxford University Press.

Stewart, P. D., S. A. Ellwood, and D. W. Macdonald. 1997. Video surveillance of wildlife: An introduction from experience with the European badger (Meles meles). Mammal Review 27: 185-204.

Stewart, P. D. and D. W. Macdonald. 1997. Age, sex, and condition as predictors of moult and the efficacy of a novel fur clip technique for individual marking of the European badger (Meles meles). Journal of Zoology 241: 543-550.

Stonehouse, B. 1978. Animal marking: Recognition marking of animals in research. Baltimore, Md.: University Park Press.

Stopka, P. and D. W. Macdonald. 1998. Signal interchange during mating in the wood mouse (Apodemus sylvaticus): The concept of active and passive signalling. Behaviour

Stopka, P. and D. W. Macdonald. 1999. The market effect in the wood mouse, Apodemus sylveticus: Selling information on reproductive status. Ethology 105: 913-1008.

Suen, H. K. and D. Ary. 1984. Variables influencing one-zero and instantaneous time sampling outcomes. Primates 25: 89-94.

Suen, H. K. and D. Ary. 1986a. Poisson cumulative probabilities of systematic errors in single-subject and multiple subject time sampling. Behavioural Assessment 8: 155169.

Suen, H. K. and D. Ary. 1986b. A post hoc correction procedure for systematic errors in time sampling duration estimates. Journal ofPsychopathology and Behavioral Assessment 8: 31-38.

Swinton, J., F. Tuyttens, D. W. Macdonald, D. J. Nokes, C. L. Cheeseman, and R. Clifton-Hadley. 1997. A comparison of fertility control and lethal control of bovine tuberculosis in badgers: The impact of perturbation induced transmission. Philosophical Transactions of the Royal Society of London B352: 619-631.

Taber, A. B. and D. W. Macdonald. 1992. Communal breeding in the mara Dolichotis patagonium. Journal of Zoology 227: 439-452.

Tew, T. E. and D. W. Macdonald. 1994. Dynamics of space use and male vigour amongst wood mice, Apodemus sylvaticus, in the cereal ecosystem. Behavioral Ecology and Socio-

biology 34: 337-345.

Thierry, B., C. Gauthier, and P. Peignot. 1990. Social grooming in Tonkean macaques

(Macaca tonkeana). International Journal of Primatology 11: 357-375.

Thompson, S. K. 1992. Sampling. New York: Wiley.

Tinbergen, N. 1951. The study of instinct. Oxford, U.K.: Oxford University Press.

Tinbergen, N. 1963. On aims and methods in ethology. Zeitschrift für Tierpsychologie 20: 410-433.

Todd, I. A. 1992. Wildtrak: Non-parametric home range analysis for the Macintosh. Oxford, U.K.: Isis Innovations.

Tuyttens, F. A. M. and D. W. Macdonald. In press. Social perturbation caused by population control: Implications for the management and conservation of wildlife. In M.

Gosling, W. Sutherland, and M. Avery, eds., Behaviour and conservation. Cambridge, U.K.: Cambridge University Press.

de Vries, H., W. J. Netto, and L. H. Hanegraaf. 1993. MATMAN: A program for the analysis of sociometric matrices and behavioural transition matrices. Behaviour 125: 157-175.

de Vries, H. 1993. The rowwise correlation between two proximity matrices and the partial rowwise correlation. Psychometrica 58: 53-69.

de Vries, H. 1995. An improved test of linearity in dominance hierarchies containing unknown or tied relationship. Animal Behaviour 50: 1375-1389.

de Waal, F. B. M. 1992. Coalitions as part of reciprocal relations in the Arnhem chimpanzee colony. In A. H. Harcourt and F. B. M. de Waal, eds., Coalitions and alliances in humans and other animals, 233-258. Oxford, U.K.: Oxford University Press.

White, G. C. and R. A. Garrott. 1990. Analysis of wildlife radio-tracking data. London: Academic Press.

White, P. C. L. and S. Harris. 1994. Encounters between red foxes (Vulpes vulpes): Implications for territory maintenance, social cohesion and dispersal. Journal of Animal Ecology

Wilson, E. O. 1975. Sociobiology. Cambridge, Mass.: Belknap.

Wong, J., P. D. Stewart, and D. W. Macdonald. 1999. Vocal repertoire in the European badger (Meles meles): Structure, context, and function. Journal of Mammalogy 80: 570-588.

Woodroffe, R. 1993. Alloparental behaviour in the European badger. Animal Behaviour 46: 413-415.

Woodroffe, R. and D. W. Macdonald. 1993. Badger sociality: Models of spatial grouping. Symposium of the Zoological Society of London 65: 145-169.

Woodroffe, R. and D. W. Macdonald. 1995a. Costs of breeding status in the European badger (Meles meles). Journal of Zoology 235: 237-245.

Woodroffe, R. and D. W. Macdonald. 1995b. Female-female competition in European badgers (Meles meles): Effects on breeding success. Journal of Animal Ecology 64: 12-20.

Wrangham, R. W. 1974. Artificial feeding of chimpanzees and baboons in their natural habitat. Animal Behaviour 22: 83-93.

Wratten, S. D., ed. 1994. Video techniques in animal ecology and behaviour. London: Chapman & Hall.

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