The evolutionary and ecological basis of behaviours, how animals adapt their behaviour to the environment, and how behaviour affects survival and reproduction
Decisions
1. Why do animals do what they do?
2. What are the selective advantages of these behaviours?
3. What triggers a change or choice of behaviour?
Behavioural Ecology
Many branches and aspects that affect the ecology of the animals, here tying to focus on those that impinge on other species and ecosystems and less on e.g., reproductive strategies
Ethology
The study of animal behaviour from an evolutionary prospective
Proximate mechanisms
Neuronal, hormonal, anatomical mechanisms
Ultimate causes
Selection pressures that shaped the evolution of the behaviour
Tinbergen's four questions
Adaptation / Function - how does the behaviour increase the animal's fitness? (Ultimate)
Evolution - how did the behaviour evolve and how has selection changed it over time? (Ultimate)
Causation - what are the triggers or stimuli that cause the behaviour to be performed? (Proximate)
Ontogeny - how has the behaviour changed over the lifetime of the animal? (Proximate)
Inheritance of behaviours
Behaviours can be learned or inherited
Much of animal behaviour is unlearned (innate) and stereotypic (always exactly the same)
Stereotypic behaviour is often species-specific
Hawk/Goose effect
Baby birds show a reaction to a hawk-like silhouette (cower) but no reaction to a goose-like silhouette
Lorenz's study of courtship behavior in ducks
Hybrid offspring expressed some elements of each parent's display, but in novel combinations – they were inherited
Females were not interested in males performing hybrid displays: evidence that sexual selection had shaped these genetically determined behaviors to be reproductive isolating mechanisms
Genes do not encode behaviour; but gene products such as enzymes can affect behaviour by starting series of gene-environment interactions that underlie development of proximate mechanisms
Inheritance of songbird songs
Male songbirds have species-specific songs used in territorial displays and courtship
White-crowned sparrows must hear the song when they are nestlings, even though they do not sing it for a whole year
As males approach maturity, they begin singing and match song with the stored memory
If the bird is deafened before starting to sing, it will not develop the species-specific song
Learning
Animals are capable of learning and developing a new response to a stimulus
Behaviours can be quite plastic
Animals are capable of learning from each other
Behavioural influences
There may be a multitude of potential triggers (situations that produce innate responses) but especially for complex or learned behaviours there can be numerous influences or conflicting pressures
Costs to behaviours
Animals have limited resources and so have to make decisions about how they spend these
This can be considered in terms of a cost-benefit analysis – will they get more back than they have expended? What are the risks involved?
Three basic costs to actions
Energetic: the actions of the animal cost energy
Risk: increased chance of death, injury or infection etc. as a result of the behaviour
Opportunity: potential benefits that are forfeited or lost from taking the action
Dominant male deer during breeding season
Encounters an unhealthy-looking female accompanied by a young male - what are the potential costs if he tried to mate with her?
Optimal Foraging
Foraging (finding and processing food) in particular is well studied in terms of costs and benefits
How long will it take to find a given food item, how much effort is involved, what will be the return?
The state of the animal is important in these calculations
Optimal foraging
Predicts that animals will make choices that maximise their energy intake
The more rapidly they can take in the fuel they need, the less opportunities are lost and the fewer risks are taken
Reindeer foraging
Prefer to feed on plant species where a single bite takes in the most leaf mass, maximising energy intake and reducing predation risk
In areas of low predator density, they can persists on smaller plants by foraging time increases
Factors that vary in food
How easy it is to find
How easy it is to process
Energetic value
Nutritional value
Palatability
Crab foraging on mussels
Prefer mid-sized mussels as a prey item
Small mussels are easy to open, but contain very little inside
Large mussels contain a lot of meat inside but take a lot of time to process
Mid-sized mussels have the optimal trade-off of effort to process and energy return for that effort
Foraging by time
Some animals may balance effort primarily by time rather than energy
Some bees chose to forage according to the time taken to reach a location and parasitoid wasps by the amount of time taken to find a potential host
Galapagos marine iguana
Juveniles primarily feed in the intertidal zone, adults underwater
Are more efficient at swimming than walking, but get cold in the sea – takes time to warm up again
Larger animals chill slower in water
Intertidal times are limited
Ends with a complex balance of time and energy to forage effectively at different sizes and at different times
Landscape of Fear
Foraging is not just about food, but it is also about risk
Animals can carry a mental map of areas to avoid or limit their time there
Deer and elephants
Deer avoid valleys in winter in the western US due to deep snow and vulnerability to wolves
Elephants that raid crops approach and leave farms in a similar way to waterholes to minimise risk of exposure
Ideal Free Distribution
Describes how animals should optimise where they forage to maximise their intake
Ideal means they have a perfect knowledge of the environment and the quality of patches, and Free means they are free to move between patches
Analyses of real world examples show some support for the Ideal Free Distribution model, but the real world is rather more complex
Resource defence
When is a resource worth defending?
Dung beetles take three basic approaches to using this resource
Ideal free distribution (IDF) model
Describes how animals should optimise where they forage to maximise their intake
Assumes animals have perfect knowledge of the environment and quality of patches
Assumes animals are free to move between patches
Analyses of real world examples show some support for the IDF model
Examples supporting IDF model
Bumblebees distribute themselves 'evenly' among patches of flowers when scaled by flower number and nectar availability
In African wild dogs, pack size correlated with prey density when travelling time was factored in
The IDF model provides a baseline or null model for examining animal distributions
Resource defence
When a resource is worth defending
Dung beetle strategies for using dung
Roll some of it away to bury
Make a ball within the pat
Tunnel into the soil beneath to make an egg chamber
Dung beetle strategies
Tunnellers are horned animals that defend a resource, and rollers are hornless and do not
Dung is a limited but valuable resource
Rollers can steal each other's balls, but by taking it away they are protecting it
Female tunnellers may compete with each other for dung to make the biggest brood ball, while males guard the dug tunnels to protect the ball and the egg
Living in groups
Provides a cost-benefits trade off
Competition is usually most intense with other members of the same species
Risks include increased competition, disease spread, attracting predators, and risk of EPCs for low ranking animals
Advantages of living in groups
Vigilance - multiple animals scouting for predators, can increase feeding times
Dilution - when attacked, there are numerous possible targets so any individual is less at risk
Group defence - groups may be capable of fending off threats that individuals are not
Group foraging
If resources are plentiful, groups may form passively
May be increased efficiency (food escapes to another)
May be able to tackle new (especially bigger) prey
May be able to find food more effectively if it is patchy
Can also share knowledge (how to handle food, learn of new patches etc.)
Bovids
Antelope and their kin
All are herbivores (grazers and browsers) and vary from c. 1 kg to 1 t
All species are predated by various carnivorans
Must trade off between feeding and vigilance
Also issues of body size (need more food, but can digest tough plants) and competition for reproduction