| Proximate questions | the how |
| Ultimate questions | the why |
| Behavioural ecology brief history | Was possibly the first science, paleolithic art showing humans observing animal behaviour. 1976 Blurton Jones observed hunter gathers who had been using behaviour ecology for centuries. 1880s C.O. Whitman coined term instinct to describe pidgins and was the first to develop analytical methods for behaviour 1871 Darwin wrote of evolution and natural selection. |
| Konrad Lorez | Nobel prize winner in 1973, 1903-1989 examined genetically programmed behaviours in young. |
| Karl Von Frisch | Nobel prize winner in 1973, 1886-1982 pioneered studies in bee communication and foraging. |
| Niko Tinbergen | Nobel prize winner in 1973, 1907-1988, formulated methods to study animal behaviour. |
| John Crook & David Lack | first to link social 6originisation of birds and primates to ecological factors. Pioneered the comparative approach, focused on ecological factors, and showed many characteristics may be inter-related and influenced by ecological factors |
| John Maynard Smith | introduced the concept of evolutionary stable strategy (ESS) as a part of game theory. Adapted form work of John Nash. ESS was an assemblage of behavioural and physical characters in a population resistant to replacement by any forms bearing a new trait. |
| Edward Wilson | father of socio-biology, revolutionised understanding of the behaviour and natural selection of colonial insects |
| Krebs and Davies | 1970-1980, a synthesis between evolutionary traditions of modern ethology and mechanistic studies of comparative psychology |
| Niko Tinbergen’s 4 explanatory levels | Proximate-looks at descriptions of organisms as a machine. Ontogeny- development of individual within it’s life span. Phylogeny- development over generations. Ultimate- examines the adaptive significance or usability of behaviour |
| A-F of behaviour | Animal-the organism. Behaviour- observable actions. Causation- proximate causes of behaviour. Development -ontogeny of behaviours. Evolution- phylogenetic content. Function- why? what is its use. |
| Fitness | the ultimate criteria on which all behaviours are scaled. When multiplied by the gene frequency will give the frequency in the next generation. Fitness of 0 meaning there are no offspring with this trait, 1 new generation has equal number as previous, fitness greater than one and next generation has higher number, less than one and next generation has less than previous |
| Inclusive fitness | personal fitness plus indirect fitness of others. Influenced relatedness |
| Hamilton’s rule | it can be of high fitness to die if your death allows your genes to be carried on my close kin. |
| Kin recognition | ability to recognise one’s kin is vital to inclusive fitness. |
| Rate | is the most important factor when choosing patch. The rate the energy can be consumed. if rate is too slow the resource may be wasted and energy will be lost collecting a resource that is not all used. |
| Profit | the net food value/ the time it takes to find food. The profit of the resource must outweigh the cost of obtaining it cost being energy and time. |
| Optimal prey selection | Predicts most profitable prey should never be ignored. Less profitable prey should be ignored depending on search time. Exclusion of less profitable prey should be all-or-nothing. Assumes that prey value is measurable. Handling time is fixed. Handling & searching cannot be done at the same time. Prey are recognized instantaneously. Prey are encountered sequentially & randomly. Energetic costs of handling are the same for different prey. Predators wish to maximize rate of energy. |
| Optimal prey selection study | Study by Krebs (1977) using Great Tits. Birds presented with small and large worms on a moving belt, varied ‘search time’ by varying distribution on belt.
Results ignored small when large abundant chose both when large less abundant .BUT, birds did not fit the zero-one rule partial preference for small worms. |
| Size restrictions | American Oystercatcher, eat mussels, optimal prey selection predicts that they would feed only on larger mussels. In actuality they have a preference on smaller mussels, this is due to the increase of handling time for larger mussels making them less profitable. |
| Nutrient constraints | optimal foraging is also constrained by nutrient value. It more important to herbivores as plants often lack needed nutrients, and a variety of plants must be consumed. |
| Optimal foraging constraints | nutrients, cognitive ability, predator avoidance, dominance higharchies and reproduction. |
| Age specific foraging behaviour | some species show difference in patch use between age groups, may be due to difference in nutritional needs. Strength to catch prey, cognitive ability to catch prey, ability to recognise prey, physiology, and skill development. |
| Patches | a group of or location of a resource e.g. patch of flowers or school of fish. Patch quality decreases with foraging behaviour due to depletion, increase of poison, diminishing prey quality, catching problems, antipredator behaviour, and load weight. |
| Marginal Value Theory MVT | Animals should, spend more time in poorer patches, spend less time in patch when average rate of energy gain is high, spend more time in patch when travel time increases. Assumes that, each patch type is recognized instantaneously, travel time between patches is known, gain curve is smooth, continuous travel time & searching within a patch have equal energy costs. |
| Marginal Value Theory test | Cowie (1977), using Great, mealworm hidden in cups around aviary. Predictions, when average feeding rate is high birds should spend less time at each patch, if travel time increases birds should spend more time at each. Results show this was consistent. |
| Factors effecting MVT | trade offs with other behaviours and constraints such as, time, psychological and behavioural. |
| Dynamic foraging models | Assume that travel times & patch quality vary in an unpredictable way. Therefore, animals might have a strategy for assessing patch quality & acquiring information by sampling. Caraco 1980 showed us that starvation changes the value of food judged by animals. |
| Central Place foraging | animals foraging around central location, often to return to young or cache of mates. Now optimal foraging must include energy to go to patch and back cost of travel is doubled. Maintenance costs as not foraging for just yourself. Cost to young whilst waiting. Increased predation risk to self and young. |
| Niche Separation inter-specific | avoidance of competition by using different resources. Closely related species altering food preference when sympatric. |
| Niche Separation Intra-specific | same species differing resource source. e.g colonial animals ensuring colonies have no overlap in patches so are not in competition with one another. |
| Niche Separation inter-sexual | different sexes in the same species using different resources. Developing difference in physiological ability to get resources e.g. size. Are sexually dimorphic animals such as size in sea lions or wing shape in albatross. |
| Crypsis | strategy for predator avoidance, avoid being seen by blending into environment. Can be inactive or actively changing with environment. Can also vary from season to season. Cost can be energetic costs to development and changing some also limit mauver ability. Benefit increases search and handling time for predators making them less profitable prey. If still seen can develop startling colours to scare prey. Could be confusing in sexual communication. Seasonal crypsis is energy costly and can be context specific if seasons change to slow you no longer blend in. |
| Aposematism | Making yourself unpalatable for predator. Must be advertised no use if predator does not know. Development of warning colouration. Evolved with convergent evolution across species to be clear. Costly to early developers as bright colours made them easy prey until predators learned not to eat them. Leads to development of Bastian mimicry of palatable prey pretending to be unpalatable |
| Physical predator avoidance | passive structures such as spikes or armour making handling time to long to eat or painful. Active structures such as horns and fangs to use as weapons. Or development of poisoning. |
| Chemical fouling | predator avoidance strategy that uses strong smells to make unprofitable prey, smell is unpleasant and nauseating but also lingers making predator easily detectable for long time making future hunt difficult. Giant pedrails also use fouling to spit at other birds that slicks feathers with oil making flight difficult |
| Chace avoidance | Must be vigilant. Moving to safety can be costly. Let predator know you have seen it and can run away so it will decide not to chase you at all. Becomes game of fake behaviour e.g. deer flicking tail to indicate has seen predator when it has not. |
| Temporal segregation | avoid predator altogether. Avoid times when predator is active, can have energetic costs for prey as best feeding times may be lost. : avoid predator altogether. Avoid times when predator is active, can have energetic costs for prey as best feeding times may be lost. |
| Spatial segregation | can be on micro- habitat scale e.g. nursing kangaroos avoiding open space sacrificing foraging efficiency. Or macro- habitat e.g. migration of Karabo or humpback whales swimming different parts of ocean to killer whales |
| Group defence | creating circles to defend animals in centre, group vigilance, dilution effect. |
| Incidental groups | living together because of shared resource, temporary dynamic. |
| Social group | reflect common dislike for being alone, common in mammals, fish and birds. Vary in size and can be single species or multiple. |
| Group advantages | foraging advantages- sharing information, group hunting, provisioning dominates gain more food than when alone and submissive about the same. Predation avoidance- group vigilance and defence subordinates need this more. Physiological l advantage- thermoregulation. Provisioning of young if genetics are shared to increase inclusive fitness. |
| Local enhancement | Individuals knowledge or resource location is gained by watching movement form others. As seen in study by Krebs 1972 with Great Tits. |
| Information centre hypothesis | shared knowledge to reduce search time amongst group members. Study by DeGroot 1980 showed Red Billed Quelea follow intention movements of knowledgeable members to resource location. |
| Effects of group size | benefits of reduced search time reach asymptote as search group size increases; members eventually overlap areas and interfere with one another. Increased size will increase energy needs that will eventually exceed energy gained by being in a group if too large. Avoiding predators, if group too large group vigilance not maintained and members will ‘cheat’. Optimal group size differs from species to species as well as time, patch quality and predation pressure. Temporary groups more often closer to optimal size than permeant as permeant change with births and increase steadily. |
| Social interactions | Benefits- shared resources, division of labour, increase indirect fitness, shared defences. Costs, sharing resources, parasitism, cheaters. |
| Mutualism | both benefit |
| Reciprocity | both benefit but if delayed to giver |
| Altruism | reviver benefits at cost to giver (selfless act) |
| Selfish behaviour | giver benefits at cost of receiver |
| Spite | both have negative cost. |
| Direct roles | behaviour of subgroup benefits another subgroup benefiting group as a whole. |
| Indirect role | selfish behaviour that is neutral or destructive to subgroups. More common in most vertebrates. e.g leadership and control. |
| Division of labour in naked mole rats | have Queen that is dominant and sole reproductive producer. Controls other females with aggression so too stressed to reproduce. Workers are male and female and smaller than Queen. Medium mole rats are diggers and largest care for young and may take Queen’s place one day. |
| Eusociality | a form of group living, happening when there is patchily distributed rescores, low reproductive success of individuals. Creates overlapping generations, increases relatedness through inbreeding. Cooperative care of young, division of labour, Haplo-diploid Hymenoptera. |
| Haplo-diploid Hymenoptera | evolved with eusociality to increase relatedness. Fitness from cooperative breeding must be greater than when alone for animals to continue to do it. Fathers are haploid and mother’s diploid. Females develop from fertilized eggs. Males develop from unfertilized eggs, have mothers, but no fathers, have daughters but no sons. Females have more investment in sisters than daughters with increased relatedness |
| How did Eusociality evolve | single queen colonies where female workers are more related to sisters than daughters. Female workers sex ratio of 3:1 is ideal but Queens 1:1 is ideal. Leading to conflict. Because males are haploid Queen controls primary sex ratio. Because workers raise young, they control secondary sex ratio. Ratio most biased in single queen colonies less in multiple queen colonies. |
| Reproductive casts | Queens vs workers. Created in larval stage by nutritional castration, difference in feeding, hormonal influences, social control (stressing them to not go into sexual maturity),environmental influences. |
| Behavioural casts | task specialisation in works. Temporal polytheism. |
| Morphological casts | morphologically and behaviourally specialised workers. |
| Eusociality in Naked Mole Rats | have Queen who controls workers. Harsh environment made colonies more beneficial. Relatedness of colony is 0.81. Queen aggressive to females to prevent them going into oestrous so are not overthrown. |
| Dispersal | tendency for once aggregated animals to become more widely dispersed. Is a one-way travel (un-like migration). Most commonly as travel away form breeding area. Scale varies from meters to kilometres. Massive costs in energy, time, risk of predation and unknow foraging grounds. Can be active by locomotion or passive by environment. Benefits, reduces intra-specific competition, colonises new habitats, reduces inbreeding. |
| Sex biased Dispersal | philopatry, tendency to breed in natural home ground. Dispersal not often equal between sexes. Dispersal of one sex away from other stops sibling on sibling inbreeding. Mammals males usually disperse, because female’s reproductive success reliant on nutrients and males’ number of females, females benefit more form familiarity. Birds females disperse more, mating biased of resource defence, males defend resource better to stay and defend good territory to get female. |
| Migration | defined route between locations. Associated with seasons or life stage. Requires enate sense of time. Reasons, food availability, mate availability, weather avoidance, predator avoidance, niche separation. |
| Orientation | compass sense, fixing on location and using cues to move in right direction. Mechanisms, sun compass use of circadian rhythm, limited by overcast. Star compass, magnetic compass, olfaction, ultra-low frequency sounds. |
| Navigation | compass sense plus map sense. Visual landmarks, geomagnetism, solar information (sun compass plus travel and time information) olfaction, combined information. |
| Territorial behaviour | benefits, protects resources including mates, access to the resource must increase fitness including inclusive fitness. Can be permanent, seasonal breeding season or weather, or dynamic. Costs, energy and risk of competition. High depletion rate of resource and unpredictability is unlikely to be economically defendable. Low depletion and high predictability rate more likely to be economically defendable |
| Super territories | Animals fitness would improve if the defend territories complete form all others making super territories. But this is unlikely. Would only increases fitness by reducing fitness of con-specifics, so selective advantage is low. If super territories became common advantage would decrease as cheaters would increase |
| Territory economics in Elephant seals | large males become beach masters that defend a territory. Gain 90% of all mating, have to be big enough to defend large area and all the females. Sneaky smaller males can infiltrate unseen to mate, would not likely win a fight so don’t even try. |
| Dear enemy effect | Fisher 1954 knowing neighbour helps decease cost of defending territory. Communication to neighbours indicating size and skill to avoid conflicted reduces unknow factors for both. Vigilance can be reduced if both know they will not attack. |
| Aggression | behaviours that are intended to inflict harm. Establishes territorial boundaries, maintenance of social higharchies, may be contagious. |
| Female aggression | defence against suckling, fence of young, defence against unwanted sexual encounters. |
| Agonism | behaviours as adjustment to situations of conflicts. Threats, submissions, physical combat. Intent not to bring harm. |
| Ritualised aggression | inter-specific aggression is usually ritualised fight with known pattens and uses weapons evolved to not harm each other. reduced blood shed for animals of same species. |
| Ritualised fighting in cervids | antlers designed to fend off blows. Antler points spread out for blows to side. When m/m aggression is occurring they direct blows away form side and into each -other antlers to minimise damage. |
| Agnostic behaviour threat displays | indicate readiness to fight and ability, may be symbolic, or redirect aggression or be intention movement such as first part of performance like baring teeth or turning into fighting position. |
| Agnostic behaviour Dominance displays | not directly related to fighting. Claims of dominance not necessarily true. May challenge or intimidate opponent. Threat and dominance displays may occur together. Usually to mane animal look bigger like broadside display showing size and lack of fear showing venerable side. Usually fight does not follow dominance display. |
| Starks and aggression | Appear territorial, parallel swimming, posture displays. Reduces needless conflicts, suggests well developed social higharchies |
| Space/territory claims | signal ownership of space, includes gland marking, wallowing, grazing, urinating, defecating, etc, occurs often in agonistic situations but is irrelevant to situation. |
| Displacement behaviours | occur in agonistic encounters, e.g. thrashing antlers in bushes etc ,may be an outlet. |
| Submissive | opposite to threat behaviour. |
| a reason why asexual reproduction may be less advantageous than sexual do to the accumulation of mutations over generations. | Mullers rachet hypothesis |
| Anisogamy | gametes of one sex larger than the other. bigger gametes increases zygote size and survival chances, smaller gametes can make more and have more chances at reproduction. Can lead to conflict amongst sexes as the sex producing bigger gametes have a larger investment in one singe reproduction larger gametes = bigger paternal investment |
| Sexual selection: | first named by Darwin. 2 forms male on male competition and female choice. |
| Monogamy | one mate for life or breeding season. Has high paternal investment. Usually happens when offspring would not survive with singe parent or females or resources are too widely distributed for male to monopolise. 92% of bird species. |
| Polygyny | many mates and low paternal investment. Leads to territorial and sexual dimorphism. Males mating with many females, happens by monopolising either the females FDP or the resources RDP. Decided by ecological factors, if females aggregate around resource then defend the resource if females don’t then protect groups of females. |
| Sneaker or Satellite | less investment in territory or aggression, often achieving results by deception. e.g. giant cuttlefish, males outnumber females 4: 1, smaller males mimic females to get close to females and not be detected by larger males, females mate with them for genetic variation, security and knowing if their sons are also small could also become sneakers. |
| Sperm competition | post copulatory competition, occurs when females mate with more than one male. Production of specialised sperm or more sperm. To defend against use mate guarding so they will be last one to mate with female. Use of nuptial gifts to entice longer mating higher chances of last mating. Females evolving post copulatory sperm selection to choose what sperm to use. Males evolving specialised mating structures such as scrubbers to remove other sperm. Or development of more pleasurable structures to increase mating duration. Sperm plugs not allowing other sperm to reach ova (common in reptiles and insects). |
| Female choice | choosing of best possible mate, limited by time, memory, location. Must choose quickly and wisely. |
| Ornamentation | exaggerated displays to show even when “disabled” by effort to grow display are still fit and heathy, may also be used in fighting. Sometimes runaway success and males evolve past the ornamentation being a true indicator of health. |
| Leks | gathering of males to display to females so females will choose mate. Males aggregate to spots where females are to show off, reduced risk of predation and greater resources than having to hunt down females. Females prefer to select mates form aggregations of males to compare. Most common in birds but also seen in whales, seals and some frogs and insects. Often in leks only one male gets all mates. |
| Polyandry | Males care for young and incubate eggs. Is rare. Females defend resource and attract mates and have more young. Often includes sex role reversal. |
| Parental care | is a costly behaviour, resources, time, predation risk, energy. Females usually carry burden of paternal care due to Anisogamy as have more investment in young. |
| Paternal... | Care -behaviour that increases fitness of offspring. Behaviour- contributes to survival rate of offspring can be maternal or paternal. Expenditure- use of paternal resource on offspring. Investment- how much the care of current offspring decreases parents’ future reproductive value. |
| Slow – fast continuum | Slow lane, slow development, delayed sexual maturity, small broods/litters, high parental care, high breeding success, higher adult survival, higher juvenile survival. Fast lane, fast development, early sexual maturity, large broods/litters, low parental care, high breeding success, low adult survival low juvenile survival. |
| Trivers and Willard (1973) hypothesis | females will invest more in female young more when in poor conditions and more in males when conditions are good. This is because males have a higher chance of having more young but need to be large and healthy to succeed. |
| Co-operative breeding | genetic parents found in 220 species of birds and 120 species of mammals. Approx. 3.2% of birds, 12% of all Australian species. Help for kin selection. Hamilton’s principle genetic relatedness times additional benefit must outweigh cost. May be used as practice, to gain future mates by proving ability to parent when unsuccessful in mating that season, may be due to limited resources to colonies take turns, may be sneakers who are secretly genetic parent. |
| Brood parasitism | tricking others into raising your young, e.g. Cuckoos and Cowbirds, Cuckoos lay eggs in Cowbirds nests and push out some of the Cowbirds eggs so they will devote energy to raising their young not their own. |
