Subject: Ethology

Scientific Area:



64 Hours

Number of ECTS:




Overall objectives:

1 - OA1-Understanding of the interactions between behaviour and environment. Understand the complementarity of questions about behaviour following Niko Tinbergens (1963) 4 questions.
2 - OA2-Understand the role of natural selection in the evolution of behaviour and distinguish between proximate and ultimate causes. Understand the concepts of adaptation, fitness differences and individual selection.
3 - OA3-Gain a solid knowledge about the diversity of behaviour in the animal kingdom, including humans.
4 - OA4-Understand social evolution and the concepts of altruism, reciprocity, cooperation and eusociality.
5 - OA5-Learn how to formulate hypotheses about behaviour, how to measure behaviour, and how to analyse the data.
6 - Skills: Ability of students to express themselves in a suitable language for the specific area of the course; Capacity of analysis and synthesis; Critical thinking and organization of ideas.


1 - Ethology: Tinbergen's questions, history; behavioural genetics; selection & evolution; ontogeny, the senses, rhythms & biological clocks, orientation & navigation; learning, motivation, communication, cognition.
2 - Behavioural Ecology: optimality & the mathematical description of behaviour: optimal foraging, evolutionary stable strategies; life in groups, evolution of sexual reproduction & sexes, sexual selection, mating systems, parental investment, selfishness & altruism, kin selection, eusociality, social insects.
3 - Practicals: introduction to behavioural measurements & asking relevant questions about behaviour. Practicals use organisms that are abundant at Madeira, including humans. Statistics is seen as the basis of behavioural analysis, with special emphasis to non­parametric methods.


Alcock, J. , 2013 , Animal behavior: an evolutionary approach (10th ed.). , Sinauer Associates, Sunderland
Krebs, J.R. & Davies, N.B.  , 1993 , An Introduction to Behavioural Ecology (3rd ed.). , Blackwell Scientific Publications, Oxford.
Dugatkin, L.A. , 2014 , Principles of animal behavior (3rd ed.). , W. W. Norton & Company, Inc, New York
Martin, P. & Bateson, P. , 1994 , Measuring behaviour: an introductory guide (2nd ed.). , Cambridge University Press, Cambridge
Stamp Dawkins, M. , 2007 , Observing animal behaviour: design and analysis of quantitative data.  , Oxford University Press, Oxford.
Yoder, P.J. & Symons, F.J. , 2010 , Observational measurement of behavior. , Springer Pub. Co., New York
Bart, J., Fligner, M.A. & Notz, W. , 1998 , Sampling and statistical methods for behavioral ecologists , Cambridge University Press, Cambridge; New York.
Li, Y. & Baron, J. , 2012 , Behavioral research data analysis with R. Use R!  , Springer, New York, NY
Pardo, S.A. & Pardo, M.A. , 2018 , Statistical methods for field and laboratory studies in behavioral ecology , CRC
Krebs, J.R. & Davies, N.B. , 1997 , Behavioural Ecology: An Evolutionary Approach. (4th ed.) , Blackwell

Assesssment methods and criteria:

Classification Type: Quantitativa (0-20)

Evaluation Methodology:
The course includes theoretical lessons (lectures) (T), theoretical­-practicals (TP), laboratory practical (PL), and fieldwork (TC). Lectures: Lectures are predominantly expository, following the organization outlined before. Theoretical­-practical classes: Students are initiated in the search for relevant peer­reviewed literature, behavioural measurement, definition of hypothesis, data analysis and basic statistics. Laboratory practicals: Students perform their first behavioural research using captive insects and lizards. Fieldwork: these classes are divided into projects involving the whole class, and individual group projects. All projects use locally abundant and easily observable species like gulls, pigeons, cats, lizards, or various insects. Student assessment: 2 written tests (30% each) and 3­5 group reports (40% total).