ISSN 1479-5647

The Federation of Zoological Gardens

of Great Britain and Ireland

Zoo Research Guidelines

Project Planning and Behavioural Observations
© The Federation of Zoological Gardens of Great Britain and Ireland 2003

All rights reserved. No part of this publication my be reproduced or transmitted in any form or by any means, electronic or
mechanical, including photocopy, recording or any information storage and retrieval system, without permission in writing
from the publisher.

Wehnelt, S., Hosie, C., Plowman, A., and A Feistner (2003) Zoo Research Guidelines: Project Planning and Behavioural
Observations The Federation of Zoological Gardens of Great Britain and Ireland, London

First published 2003

Published and printed by:

The Federation of Zoological Gardens of Great Britain and Ireland,
Zoological Gardens,
Regent’s Park,
London NW1 4RY,
United Kingdom

ISSN 1479-5647
 Zoo Research Guidelines:
Project Planning and Behavioural Observations
Stephanie Wehnelt1, Charlotte Hosie2, Amy Plowman3, Anna Feistner4
1
North of England Zoological Society, Chester Zoo, Upton, Chester CH2 1LH, UK;
2
Chester College, Department of Biological Sciences, Parkgate Road, Chester CH1 4BJ, UK;
3
Paignton Zoo Environmental Park, Totnes Road, Paignton, Devon TQ4 7EU, UK;
4
Durrell Wildlife Conservation Trust, Les Augres Manor, Trinity, Jersey JE3 5BP, British Isles

Acknowledgements for valuable comments:

Richard Brown, Geoff Hosey, Filippo Aureli, Colleen Schaffner

Scientific investigations are the basis for understanding the animals in a zoo, assessing
the way they are cared for and how they interact with their zoo environment. Through
careful research, we can improve animal management, husbandry, welfare, breeding,
diets etc. Research conducted in zoos can also contribute to the conservation of
animals in situ. In zoos it is possible to study animals closely, which can be an excellent
training for observations in the field. Zoos also provide a valuable opportunity to train
young scientists and other interested parties in scientific research techniques.

These guidelines are designed to clarify the series of steps that are usually involved in
developing and conducting a research project in a zoo environment. They discuss those
challenges that are characteristic for zoo research projects and that may differ from
those encountered in laboratory or field studies. The guidelines are intended to assist
those relatively new to zoo research in carrying out sound, well-designed projects that
have scientific validity.

The first volume of this series of Zoo Research Guidelines is organised into sections
identifying the steps in the research process: 1. Formulating the research question, 2.
Developing the research design, 3. Data analysis and interpretation,
4. Compiling the project report. Please note that these steps may be cyclical and
interdependent, e.g. experimental design and analysis should be considered together. It
is advisable to read all sections before starting and consider carefully any cross-
references between sections. The final section 5. presents a wide range of useful
references and identifies key texts for each stage of the project.
1. FORMULATING THE RESEARCH QUESTION & INITIAL PRACTICALITIES
• Identify the question(s) and the species upon which you wish to focus. Consult your
supervisor to make sure that these are appropriate for a zoo project using perhaps
preliminary reference to other studies. A thorough literature search on both the
species and the question is then the next step. Electronic databases such as ‘Web of
Science’ are the easiest way to search for scientific papers on a particular topic.
The Internet and reports on previous zoo projects, which can be obtained by
contacting Zoo Libraries or Zoo Research Coordinators, can be useful. However, the
source and accuracy of Internet material should be carefully considered. Also
previous project reports may be unreliable and inaccurate due to the restricted
input from zoo personnel or poor supervision. Read unpublished projects critically
and decide if the information is valid. Avoid following on from a scientifically
unsound project unless you can make changes and improve on it. Do not base your
project too closely on a previous study as this may be classed as plagiarism. If in
doubt consult your project supervisor.
Example: If you were interested in studying the effect of visitor numbers on the
behaviour of a group of macaques, your literature search should cover 'macaque
behaviour' both 'in the wild' and 'in zoos' plus 'visitor effects on zoo
animals/primates'. You could specify your research and measure noise levels or note
down age structure of the visitor crowd.

• Discuss the project with your supervisor before contacting the zoo or animal
collection. A list of suggested projects may be available from your supervisor, the
Zoo Library or the Zoo Research Coordinator. Check carefully what your course
requires with regards to the execution and write-up of the project and whether
these requirements can be fully met by the intended zoo project. For example, are
descriptive statistics adequate or is statistical hypothesis testing a requirement?

• If you are planning to investigate physiological parameters to back-up your

behavioural observations, make sure you are familiar with the legislation and zoo-
specific regulations for invasive sampling procedures. The Zoo Federation Research
Group has produced Sampling Guidelines that provide information on what samples
can be taken from animals without incurring a Home Office 'regulated procedure'
license (see section 5.). Most zoos do not hold a Home Office Licence. You can find
further information on ethical aspects of animal research procedures in the
reference section 5. If in doubt, consult the Research Coordinator at the zoo.

• Find out the registration procedure for conducting a project at the zoo you have
chosen. Start by contacting your supervisor who might have zoo registration forms
or information leaflets. If there is no information available, contact the Zoo
 Research Coordinator. It is helpful, and often a requirement, to have a short, typed
project proposal to submit to your supervisor and the zoo.

• Identify your study animals and determine any group structure. Ask the zoo for
identification sheets or a copy of the taxon report for the species of your interest.
The taxon report gives individual information on age, sex, parents, etc. Check
through any previous project reports that are available on these animals (ask the
Research Coordinator or the Zoo Librarian). Previous reports may not only hold
information on the individuals but often also contain enclosure maps. Determine
whether identification of individuals is possible within your time constraints. It
might be sufficient for your particular research question to subdivide the population
by sex or age group.

• Define the specific question(s) you wish to address before you start any data
collection. Do your literature searches direct you towards specific hypotheses? You
should be able to justify your project aims from the literature.
2. DEVELOPING THE RESEARCH DESIGN
2.1 Practicalities and Responsibilities
• Consider what types of projects are appropriate for a zoo environment. For example
invasive studies that require a Home Office Licence may be possible in a laboratory
but will not be accepted in a zoo. For ethical reasons it is unlikely that you will be
able to collect data that would require separating individual animals from a group
(e.g. for individual feeding trials) or catching or anaesthetising the study animals.

• Discuss with your supervisor and if necessary with the Zoo Research Coordinator
how feasible your initial design will be to implement. The project needs to work
around zoo regimes (e.g. animal feeding times, keepers rota) and avoid disrupting the
animals or the keepers. Check the husbandry regime for your study animals and bear
in mind that you may not be able to observe them at certain times (e.g. outside of
zoo opening times or when they cannot be seen from public viewing areas). By
handing in a preliminary proposal well before the start of your project, the zoo will
have the opportunity to inform you about any possible problems.

• Always keep to the arrangements that you made with zoo staff (e.g. for putting
enrichment devices into enclosures) and ensure that you turn up on the days and at
the times agreed. Be sure to inform the keepers and/or the Research Coordinator if
you make changes to your research plans.

2.2 Experimental Design

Please note that any statistical tests you may do depend closely on the design of your
experiment and the way in which you collect your data. You should read sections 2. and
3. in conjunction with each other. If you are in any doubt you should consult detailed
literature (see section 5.) and/or your supervisor.

• Identify independent and dependent variables in your experiment. The independent

variable is the factor that you, as the researcher, will manipulate. The dependant
variable is what you will measure. The independent variable can be manipulated
either directly or indirectly.
Example: Independent variables might include the addition of enrichment devices
(directly controlled by you) or visitor numbers (indirect measure since you cannot
control it). The dependent variable might be position of the animal(s) in the
enclosure or behaviour frequencies or durations.

• If your project requires observation of detailed behaviour you will need to

construct an ethogram. This is a detailed list of objectively described, mutually
exclusive behaviours that you will focus on to answer your research question(s).
 These behaviours provide you with measurable units for which you can record
frequencies and durations or latencies. Literature research (e.g. see section 5.),
previous studies on the same animals and preliminary observations can help to
identify and define these behaviours.

• Think carefully about the context of your data collection – what else might be going
on at the same time and possibly affecting it? Eliminate as many confounding
variables as possible so that the only factor that affects your dependent variable
is/are your independent variable/s. Those that cannot be eliminated should be
controlled for as far as possible and acknowledged and discussed in your project
report (see section 3.).
Example: Animals may reduce their activity when visitor numbers increase but does
this also coincide with higher temperatures due to better weather?

2.3 Choosing Sampling Methods

• Exactly what data you collect depends on your hypothesis. The variables you
measure should be chosen to best describe what is of interest to answer the
research question. Avoid taking unnecessary data by remaining focused on your
hypothesis.
Example: Instead of collecting general data on, for example, everything that each
individual of a group does, you might focus specifically on where individuals are
found in the enclosure and behaviour shown there. If you are looking at the effect
of enrichment devices, you might focus on behavioural elements relevant to the
device (e.g. foraging and feeding) or focus more broadly on changes in time budgets,
but it would not be necessary to collect data on details of e.g. social behaviours.

• Decide when to collect the data. You need to consider time of day, visitor numbers,
temperature, weather conditions, animal breeding season etc (also see section on
confounding variables in 2.2.). Make sure you plan your data collection so that you
have enough time to answer your questions within such constraints and allow extra
time for unexpected problems. Please note when interpreting the results of a short-
term project that the outcome cannot be seen as being representative for the
animals' behaviour all year round! – see section 3.

• Consider how/when/how often etc. to record the data and ensure you are using the
most appropriate sampling method to answer your question. This is important in
determining whether your data are suitable for statistical analysis, so do check if
you intend such analysis. (See section 5.0 for helpful references on sampling and its
relationship to statistical tests.)
 Example: You might study one focal animal at a time, select individuals out of sub-
groups or scan the behaviour of the whole group at the same time point. Sampling
methods include; ad lib, focal and scan behaviour sampling. Recording can be
continuous, instantaneous, frequency, etc.

• The number of observations you take will depend on how many animals you have
available, how many different experimental conditions you are planning to compare,
the probability tests you have decided to use (if any) and how strong the effect of
your independent variable is. Decide how long you will leave between sampling periods
(among other factors, this will depend on the duration of the behaviours observed)
and how you will avoid bias when choosing a subject to sample. It is important that
these issues are carefully considered to avoid generating errors in interpreting
results and/or any statistical analyses. If certain individuals are more active/visible,
their behaviour maybe recorded repeatedly (and so be over-represented in the
results) instead of obtaining data from all individuals evenly.
Example: To avoid bias, pick subjects at random or in a set order rather than
focusing on the animal displaying the most interesting behaviour at that time. This
might result in lots of, e.g. 'inactive' or 'out of sight' behaviour being recorded but,
even if this seems less interesting to you, the observer, it is still valid data.

• Once you have planned your data collection it is very valuable to carry out
preliminary observations (a pilot study) to iron out any unforeseen problems with
your research design or sampling strategies. It also enables you to judge
realistically how long the data collection for the whole project will take and to plan a
timetable accordingly (with extra time added for further unforeseen problems!).
3. DATA ANALYSIS AND INTERPRETATION
• If you are intending to carry out probability testing, you should have decided on the
tests that you will use before starting to collect your data (as discussed in section
2.). This will ensure that you collect data in the right form and are not left
wondering what test you can use on your results!

• To consider which statistical tests to use, try consulting general statistics books
and texts, particularly those that are especially directed at zoo or field researchers
(see section 5.). You will need to consider (and test) whether the data you have
collected are normally distributed as this will determine whether parametric or non-
parametric tests can be used. Generally parametric tests are more powerful but
depend closely upon having normally distributed data (among other constraints).
Where data are not normally distributed and transformations are not recommended
or helpful, similar non-parametric tests are available. Bear in mind that it may not
always be appropriate or necessary to use hypothesis testing with statistics.
Descriptive statistics and graphical representations of data may be more
appropriate than trying to force data into an inappropriate test that may provide
invalid or unrepresentative results and lead to erroneous conclusions.

• As with the previous stage in the process, preliminary analyses after some data have
been collected are extremely useful to check whether any further refinements to
the data collection are needed. Therefore allow time for preliminary observations
and analyses when planning your observation timetable.

• Some statistical problems can be typical for zoo projects (e.g. pseudoreplication and
pooling errors) usually because of small sample sizes, inappropriate repeated
measures etc (see also section 2.3). Statistics books and previous research that
address these issues (see section 5.) can help to prevent the occurrence of these
errors. There are, for example, different correction methods available but a major
way of avoiding these problems is through carefully planned experimental design.

• Take care when interpreting your results to reconsider the planning and design part
of the process and think back over any confounding or extraneous variables. Did any
others arise during the course of your data collection (e.g. new animal arrived in
collection, new feeding method, strong weather change)? How might these,
individually and together, have affected your results? You need to discuss these
thoroughly; in some studies they have a larger effect than the independent variable.
Consider the value of your results for the observed animals (in study zoo) and also,
if appropriate, how applicable these results are for the subject species in other
zoos/wild.
4. THE PROJECT REPORT
• The style of the project report will obviously depend on the requirements of your
course and academic institution but it is essential that you produce at least one copy
for the Zoo. Check whether you should pass it to the zoo personally and to whom it
should be addressed (usually to the Research Coordinator) or whether your
academic institution collects it to pass on. Without a written report the knowledge
gained from the research will not be available for zoo staff, which means your work
does not benefit the animals. Be sure to agree on the ownership of any data
collected with the Zoo Research Coordinator before you start your project. Some
zoos set as a requirement that they are provided with an electronic copy of your
data at the end of your project. Be sure to comply with any such requests to ensure
valuable links and good relations between zoos and academic institutions are
maintained.

• The most useful reports include information that would allow another person to
follow the same study on different animals or in other situations. The report should
state the background of the study and why it was carried out. It must include
details of the methods used, particularly when exactly the data was collected, how
many hours of observations were collected and details of behavioural sampling
methods and experimental design. State exactly what the results were (including
any statistical details) and include a discussion of how the results can be
interpreted, using knowledge of the theoretical background. It is often useful for
the zoo if you add recommendations for husbandry procedures that result from your
research.
5. USEFUL REFERENCES
5.1 Zoo Research – Background and Contexts
Key references:
• Crockett, CM (1996) Data collection in the zoo setting emphasising behavior. In:
Mammals in Captivity; Principles and Techniques, Kleiman, DG, Allen, ME, Thompson,
KV and Lumpkin, S (eds). University of Chicago Press, Chicago, pp.545-565.
• Federation Research Group (2002) Research Sampling Guidelines for Zoos. The
Federation of Zoological Gardens of Great Britain and Ireland, London.
Other useful texts and key papers dealing with general zoo research issues:
ASAB (2001) Guidelines for the treatment of animals in behavioural research and
teaching. Animal Behaviour 61: 271-275.
Benirschke, K (1996) The need for multidisciplinary research units in zoos. In: Mammals
in Captivity; Principles and Techniques, Kleiman, DG, Allen, ME, Thompson, KV and
Lumpkin, S (eds). University of Chicago Press, Chicago, pp.537-544.
Bostock, S St C (1993) Zoos and Animal Rights. Routledge, London. (see chapter on
‘Science in Zoos’)
Burghardt, GM (1996) Introduction: Research and welfare in animal exhibit facilities.
In The Well-being of Animals in Zoo and Aquarium Sponsored Research, Burghardt, GM,
Bielitski, JT, Boyce, JR and Schaeffer, DO (eds), Scientists Center for Animal
Welfare, Greenbelt, pp. 1-3.
Feistner, ATC and Price, EC (2000) Working together for conservation: A win-win
strategy for zoos and universities. In Proceedings of the 2nd Annual Symposium on Zoo
Research, Plowman, AB (ed.), Federation of Zoological Gardens of Great Britain and
Ireland, London, pp. 23-30.
Fitzroy Hardy, D (1996) Current research activities in zoos. In: Mammals in Captivity;
Principles and Techniques, Kleiman, DG, Allen, ME, Thompson, KV and Lumpkin, S (eds).
University of Chicago Press, Chicago, pp.531-536.
Fouts, R (1995) Science in zoos: arrogance of knowledge versus humility of ignorance.
In Ethics on the Ark: Zoos, Animal Welfare and Wildlife Conservation, Norton, BG,
Hutchins, M, Stevens, EE and Maple, TL (eds) (1995) Smithsonian Institution Press,
London, pp.277-285.
Hutchins, M (1988) On the design of zoo research programs. International Zoo
Yearbook 27: 9-18.
Hutchins, M, Dresser, B and Wemmer, C (1995) Ethical considerations in zoo and
aquarium research. In Ethics on the Ark: Zoos, Animal Welfare and Wildlife
Conservation, Norton, BG, Hutchins, M, Stevens, EE and Maple, TL (eds) (1995)
Smithsonian Institution Press, London, ppp.253-276.
Hutchins, M, Paul, E and Bowdoin, JM (1996) Contributions of zoo and aquarium
research to wildlife conservation and science. In The Well-being of Animals in Zoo and
Aquarium Sponsored Research, Burghardt, GM, Bielitski, JT, Boyce, JR and Schaeffer,
DO (eds), Scientists Center for Animal Welfare, Greenbelt, pp. 23-39.
IUDZG/CBSG(IUCN/SSC) (1993) The World Zoo Conservation Strategy; The Role of
the Zoos and Aquaria of the World in Global Conservation. Chicago Zoological Society,
Chicago.
Kleiman, DG (1992) Behavior research in zoos: past, present, and future. Zoo Biology 11:
301-312.
Kleiman, DG (1995) Criteria for the evaluation of zoo research projects. Zoo Biology 4:
93-98.
Kleiman, DG (1996) Special research strategies for zoos and aquariums and design of
research programs. In The Well-being of Animals in Zoo and Aquarium Sponsored
Research, Burghardt, GM, Bielitski, JT, Boyce, JR and Schaeffer, DO (eds), Scientists
Center for Animal Welfare, Greenbelt, pp. 15-22.
Kleiman, DG, Allen, ME, Thompson, KV and Lumpkin, S (eds) (1996) Wild Mammals in
Captivity; Principles and Techniques. University of Chicago Press, Chicago.
Lindburg, DG (1993) Curators and applied science. Zoo Biology 12: 317-319.
Mellen, JD (1994) Survey and interzoo studies used to address husbandry problems in
some zoo vertebrates. Zoo Biology 13: 459-470.
Norton, BG, Hutchins, M, Stevens, EE and Maple, TL (eds) (1995) Ethics on the Ark:
Zoos, Animal Welfare and Wildlife Conservation. Smithsonian Institution Press, London.
O’Connell, M (2000) Threats to waterbirds and wetlands: Implications for conservation,
inventory and research. Wildlfowl 51: 1-15.
Robinson, MH (1998) Enriching the lives of zoo animals: Where research can be
fundamental. Animal Welfare 7: 151-175.
Ryder, OA (1995) Zoological parks and the conservation of biological diversity: Linking
ex situ and in situ conservation efforts. Journal of Environment and Development 4(2):
105-120.
Ryder, OA and Feistner, ATC (1995) Research in zoos: A growth area in conservation.
Biodiversity and Conservation 4(6): 671-677.
Stoinski, TS, Lukas, KE and Maple, TL (1998) A survey of research in North American
zoos and aquariums. Zoo Biology 17: 167-180.
Thompson, SD (1993) Zoo research and conservation: Beyond sperm and eggs toward
the science of animal management. Zoo Biology 12: 155-159.
Wemmer, C, Rodden, M and Pickett, C (1997) Publication trends in zoo biology: A brief
analysis of the first 15 years. Zoo Biology 16: 3-8.

5.2 Developing the Research Design

Key references: These provide a good starting point for developing your research
design; the first two cover what their titles suggest, the others give valuable
information on how to plan behavioural observations, devise ethograms, determine
sampling strategies etc (see also the separate specialist reference sections 5.3 and
5.4).
• Adams, DC and Anthony, CD (1996) Using randomisation techniques to analyse
behavioural data. Animal Behaviour 51: 733-738.
• Bart, J, Fligner, MA and Notz, WI (1998) Sampling and Statistical Methods for
Behavioural Ecologists. Cambridge University Press. Cambridge.
• Dytham, C (1999) Choosing and Using Statistics – A Biologists Guide. Blackwell
Science Ltd. Oxford.
• Martin, P and Bateson, P (1993) Measuring Behaviour. An Introductory Guide,
Second Edition. Cambridge University Press, Cambridge.
• Crockett, CM (1996) Data collection in the zoo setting emphasising behavior. In:
Mammals in Captivity; Principles and Techniques, Kleiman, DG, Allen, ME, Thompson,
KV and Lumpkin, S (eds). University of Chicago Press, Chicago, pp.545-565.
• Saudargas, RA and Drummer, LC (1996) Single subject (small N) research designs
and zoo research. Zoo Biology 15: 173-181.
• Still, AW (1982) On the number of subjects used in animal behaviour experiments.
Animal Behaviour 30: 873-880.
Other useful general texts and key papers dealing with experimental
design/statistical issues:
Crawley, MJ (1993) GLIM for Ecologists. Blackwell Scientific, Oxford.
Grafen, A (2002) Modern Statistics for the Life Sciences. Oxford University Press,
Oxford.
Hurlbert, SH (1984) Pseudoreplication and the design of ecological field experiments.
Ecological monographs 54(2): 87-211.
Little, RJA (1987) Statistical Analysis with Missing Data. Wiley, New York.
Machlis, L, Dodd, PWD and Fentress, JC (1985) The pooling fallacy: Problems arising
when individuals contribute more than one observation to the data set. Zeitschrift für
Tierpsychologie 68: 201-214.
McConway, K (1992) The number of subjects in animal behaviour experiments: is Still
still right? In Ethics in research on animal behaviour: readings from Animal Behaviour,
Dawkins MS and Gosling M (eds), Academic Press for the Association for the Study of
Animal Behaviour and the Animal Behaviour Society, London.
Medawar, PB (1981) Advice to a Young Scientist. Pan Books, London.
Miller, S (1984) Experimental Design and Statistics. Second Edition. Methuen, London.
Sutherland, WJ (2000) The Conservation Handbook: Research, Management and Policy.
Blackwell Science Ltd, Oxford.
Sutherland, WJ (ed.) (2000) Ecological Census Techniques, a Handbook. Cambridge
University Press, Cambridge.

5.3 Principles of Behavioural Observation

This section focuses specifically on the principles underlying behavioural observation
and how it can be used to answer particular questions. (See also section 5.4.)
Key references:
• Altmann, J (1974) Observational study of behavior: sampling methods. Behaviour
49: 227-267.
• Dunbar, RIM (1976) Some aspects of research design and their implications in the
observational study of behaviour. Behaviour 58: 78-98.
• Federation Research Group (2002) Research Sampling Guidelines for Zoos. The
Federation of Zoological Gardens of Great Britain and Ireland, London.
• Martin, P and Bateson, P (1993) Measuring Behaviour; An Introductory Guide Second
Edition. Cambridge University Press, Cambridge.
• Saudargas, RA and Drummer, LC (1996) Single subject (small N) research designs
and zoo research. Zoo Biology 15: 173-181.
Other useful texts dealing with principles of behavioural observations:
ASAB (2001) Guidelines for the treatment of animals in behavioural research and
teaching. Animal Behaviour 61: 271-275.
Bostock, S St C (1993) Zoos and Animal Rights. Routledge, London. (see chapter on
‘Science in Zoos’)
Harcourt, AH (1978) Activity periods and patterns of social interaction: a neglected
problem. Behaviour 66: 121-135.
Hutchins, M (1988) On the design of zoo research programs. International Zoo
Yearbook 27: 9-18.
Hutchins, M, Dresser, B and Wemmer, C (1995) Ethical considerations in zoo and
aquarium research. In Ethics on the Ark: Zoos, Animal Welfare and Wildlife
Conservation, Norton, BG, Hutchins, M, Stevens, EE and Maple, TL (eds) (1995)
Smithsonian Institution Press, London, pp.253-276.
Kleiman, DG (1992) Behavior research in zoos: past, present, and future. Zoo Biology 11:
301-312.
Lehner, PN (1987) Design and execution of animal behavior research: An overview.
Journal of Animal Science 65: 1213-1219.
Lindburg, DG and Fitch-Snyder, H (1994) Use of behavior to evaluate reproductive
problems in captive mammals. Zoo Biology 13: 433-445.
Thiemann, S and Kraemer, HC (1984) Sources of behavioral variance: implications for
sample size decisions. American Journal of Primatology 7: 367-375.
Tinbergen, N (1963) On aims and methods of ethology. Zeitschrift für Tierpsychologie
20: 410-433.

5.4 Observing behaviour

This section addresses techniques and problems of behavioural observation as a large
number of zoo research projects rely on reliable and accurate behavioural observations.
(See also section 5.3.)
Key references:
• Altmann, J (1974) Observational study of behavior: sampling methods. Behaviour
49: 227-267.
• Bernstein, IS (1991) An empirical comparison of focal and ad libitum scoring with
commentary on instantaneous scans, all-occurrences and one-zero techniques. Animal
Behaviour 42: 721-728.
• Dunbar, RIM (1976) Some aspects of research design and their implications in the
observational study of behaviour. Behaviour 58: 78-98.
• Martin, P and Bateson, P (1993) Measuring Behaviour; An Introductory Guide Second
Edition. Cambridge University Press, Cambridge.
• Saudargas, RA and Drummer, LC (1996) Single subject (small N) research designs
and zoo research. Zoo Biology 15: 173-181.
Other useful texts dealing with techniques and problems of behavioural
observations:
Caro, TM, Roper, R, Young, M and Dank, GR (1979) Inter-observer reliability. Behaviour
69: 303-315.
Chow, IA and Rosenblum, LA (1977) A statistical investigation of the time-sampling
methods in studying primate behavior. Primates 18: 555-563.
Harcourt, AH (1978) Activity periods and patterns of social interaction: a neglected
problem. Behaviour 66: 121-135.
Hinde, RA (1973) On the design of check-sheets. Primates 14: 393-406.
Kraemer, HC (1979) One-zero sampling in the study of primate behaviour. Primates 20:
237-244.
Lindburg, DG and Fitch-Snyder, H (1994) Use of behavior to evaluate reproductive
problems in captive mammals. Zoo Biology 13: 433-445.
Rhine, RJ and Ender, PB (1983) Comparability of methods used in the sampling of
primate behavior. American Journal of Primatology 5: 1-15.
Rhine, RJ and Flanigon, M (1978) An empirical comparison of one-zero, focal animal and
instantaneous methods of sampling spontaneous primate behavior. Primates 19: 353-
361.
Rhine, RJ and Linville, AK (1980) Properties of one-zero scores in observational studies
of primate social behavior: the effect of assumptions on empirical analyses. Primates
21: 111-122.
Thiemann, S and Kraemer, HC (1984) Sources of behavioral variance: implications for
sample size decisions. American Journal of Primatology 7: 367-375.
Tinbergen, N (1963) On aims and methods of ethology. Zeitschrift für Tierpsychologie
20: 410-433.
Tyler, S (1979) Time-sampling: A matter of convention. Animal Behaviour 27: 801-810.