Animal Behavior/Ethology and Behavioral Ecology

General Biology BI 04 Summer School Lecture Notes


Behavior is what an animal does

Patterns of behavior may be simple or complex

Brief History

First ethologists - pre-historic humans, studied behavior for practical purposes

prey behavior for hunting

examples - cave art

domestication - dogs, cats, ox, cattle, reindeer, horses, etc.

Middle Ages

Natural theologians document behavior as part of general biology of organisms

Early 1900s - formal discipline

Due to work of 3 ethologists

K. Lorenz studied waterfowl and other organisms

N. Tinbergen studied gulls and other organisms

K. Von Frisch studied communication in bees

Late 1900s and early 2000 - modern studies of animal behavior

descriptions of behavior

behavioral genetics

neurophysiology of behavior

sensory physiology

development of behavior


behavioral ecology

Causes of behavior

Proximate causation - immediate causes

Explains how behavior works - what stimulates behavior to occur

Study by measuring or describing the stimuli that elicit behavior

Internal - physiological events (hormones, nervous system)

External - environmental stimuli

Ultimate causation - historical explanations

Explains why a behavior evolved

Study by measuring influence on survival or reproduction

Example - bird migration

Proximate causes

External stimuli- changes in daylength

Internal stimuli - hormone levels

Ultimate causes - birds that migrate have a selective advantage over birds that don't/didn't, selected for over time, could be due to long term climate changes, glaciation, disease, taking advantage of food sources, etc.

Components of Behavior

2 Components

Nature/innate: instinct and genes determine behavior

Nurture/learned: experience and learning influence behavior

Two extremes are not mutually exclusive, but work together to influence behavior

Examples of Innate behavior

1) nest building in Lovebirds by Dilger

Fischer's Lovebird - uses long strips of nest materials, carries in beak, one at a time

Peach-faced Lovebird - tucks several short strips in feathers

Hybrids - intermediate lengths of nest materials, clumsy behavior trying to tuck strips into feathers, later will carry strips in bill but will still try and tuck into feathers

2) egg ejection by cuckoos (brood parasites)

3) freezing behavior of nestling birds when exposed to silhouettes (raptors versus waterfowl)

4) Parental feeding - brood parasites take advantage of parents

5) Freezing behavior of nestlings

6) incubation behavior of some birds (Oystercatchers)

7) Drosophila - 2 alleles of the dg2 gene

sitter allele (sedentary behavior)

rover allele (hyperactive, mobile)

Components of Innate Behavior

FAP - fixed action pattern, all or none response

Sign stimulus - causes release of FAP

Examples - colors of stickleback males during mating, oystercatchers and eggs during incubation (super-normal releaser)

Nature of sign stimulus - usually an obvious aspect of the morphology: red mark on beak of Herring Gulls, red belly of Sticklebacks, detection of ultrasounds from bats by prey species of moths

Learned Behavior

Simple learning - habituation, species of prey and the presence of predators

Lehrman's study of gull chick feeding behavior - how an instinct is learned

Learning and development - imprinting and Lorenz's classic experiments with Greylag Goose (critical period for learning) - geese forms social attachments shortly after birth , salmon and home stream, birds and breeding range, nesting materials, etc.

Sexual imprinting - Direct sexual behavior at member of one`s own species - cross-fostering studies, individuals raised by another species, recognizes foster species as its own when sexually mature, will attempt to mate with foster species

Imprinting in conservation biology - minimize/eliminate human presesence while raising California Condors

Song learning in birds

White-crowned sparrows reared under 3 conditions: 1) normal, 2) deafened, 3) in isolation, critical period

Continuos song learning - mimic thrushes, starlings and mynahs, some parrots and Cardueline finches

Classical/Pavlovian conditioning

animals make associations - Pavlov's dog associates bell with food, begins to salivate, can be extinguished and later followed by recovery (unconditioned stimulus - meat, unconditioned response - salivation, conditioned stimulus - bell, conditioned response - salivation)

Operant conditioning

reward/punishment for behavioral response, rats bar press for food

Observational learning - social imitation

Insight Learning

chickadees/tits and opening milk bottles

All examples of tool-using

Egyptian Vulture - uses rocks

Cocos Finch - uses splinters of wood

North American Gulls, Northwestern Crow - smash clams on sandy beaches

Play Behavior

young animals engage in play, precursors for adult behavior (e.g., fighting, sexual behavior, predation - cats, birds, Killer Whales), also stay in good physical condition


definition - usually defined as regular, seasonal movements from one area to another (wintering area to a breeding area), seasons can be fall/spring in temperate areas and rainy versus dry in tropical areas


Waterfowl flyways - well documented, due in part to hunting regulated and studied by US Fish and Wildlife Service, State fish and game agencies

Arctic Tern, North Pole to South Pole

some species of Gallinaceous and raptorial birds, altitudinal migration (valley to mountain peaks)

Salmon - return to native streams to breed after several years at sea

Deer and Caribou, African ungulates - mammals engaged overland migrations

Some sharks, whales (northern oceans for calving, southern areas for breeding) and other marine mammals - engaged in long distance oceanic migrations

Navigation and Orientation

Animals use cues in the environment to guide them during migration, must orient and navigate

Orientation - organism is capable of detecting compass direction (N, S, E, W) using cues from the environment

Navigation - organism is capable of detecting its position as well as orientation, (N, S, E, W of something - river. ocean, mountain range)

Example - study of migration in European Startlings

Juveniles - displaced individuals were able to orient only, could not navigate (never corrected for their displacement)

Adults - displaced individuals were able to orient and navigate (adults did correct for their displacement)

Birds can use several cues from their environment for navigation - visual (sun, stars, visual landmarks) and magnetic cues

Emlen's classic study of Indigo Buntings and their use of star constellations during spring and fall migrations

Timing of behavior

circaannual - behavior occurs on a seasonal/annual basis

Examples - hibernation in bears, frogs, toads, salamanders bury themselves in mud during the winter

circadian - behavior occurs on a daily basis


endogenous - hormonal

exogenous - external cues from the environment

Example - bird migration caused by exogenous cues like decreasing/increasing daylight, picked up by eye and visual cortex; endogenous - pineal gland and brain processes daylight information, brain contacts endocrine system, alters timing of secretion of prolactin and corticosterone

Predatory Behavior

Active pursuit - tarantula, scorpions, wasps, bats

Cooperative hunting

Wait and hide - web-building spiders, trap-door spider, many raptorial birds

Fishing by Angler Fish

Other - deception and camouflage

Anti-predatory Behavior

Batesian Mimicry

Mullerian mimicry

Camouflage, disruptive, cryptic coloration

Warning coloration

Maintenance Behavior



Behavioral Ecology - study of behavior in an ecological context, study ecological variables and relationship to behavior

Optimal foraging behavior - do species forage in an efficient manner that maximizes benefits and minimizes costs

1) Example - Zach's study of Northwestern Crows dropping shellfish to break the hard outer shell and make available the clam for food, experimenter dropped shells from various height to find the optimal height necessary to break shells, observed crows dropping shells and found their average height of 5.23 m was similar to the experimenter's optimal height

2) Example - Sunfish, provide predator with prey of different sizes and different densities, fish respond by foraging optimally (taking the most energetically rich prey under the appropriate conditions)

Social Behavior - any interactions between 2 or more individuals

Anti-predatory behavior - group defense in Musk Ox

Agonistic behavior

ritualized behaviors that substitute for physical contact and fighting - yawn of baboons, dogs and baring their teeth, cats and raising their fur, birds raising their feathers

fighting and physical contact, wolves, coyotes, seals, walruses, etc,

Dominance hierarchies

peck order of turkeys and chickens

wolves and primates - alpha males......


birds and birdsong - breeding, wintering, year-round territories, size of territory related to size of bird (small birds with small territories, large birds with large territories, some exceptions - colonial nesting species - large birds with small territories)

marking territories

birds - birdsong

mammals - urine

carpenter bees - constantly standing guard

Reproductive behavior - Courtship

highly ritualized, species specific behavior

Example - sticklebacks

Reproductive behavior - mating systems

monogamy - one male and one female


polygyny - one male and many females

polyandry - one female and many males

promiscuity - anything goes

tied to parental investment, length of family bond, ecological variables and their potential to be controlled

Communication systems

auditory - birds and humans, some insects (crickets)

olfactory - most mammals, moths

visual - bees and dancing, fireflies at night

Altruism - personal sacrifice for the good of the group

alarm call of mammals - Belding's Ground Squirrels

cooperative breeding - African bee-eaters, Scrub Jays

bees and other hymenopterous insects

Mole rats of Asia - similar to hymenopterous insect colonies with Queen and workers, workers protect and feed the Queen, sacrifice reproductive opportunities

Sociobiology - comparative study of animal culture

Study of animal cultures

Cognition - do animals think?



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Copyright 2001 Jay Pitocchelli. All rights reserved. The contents of this page are the intellectual property of Dr. Jay Pitocchelli for distribution to students enrolled in General Biology BI 04 at Saint Anselm College. These pages may not be copied, photocopied, reproduced, translated, or published in any electronic or machine-readable form in whole or in part without prior written approval of Jay Pitocchelli. Students enrolled in General Biology BI 04 at Saint Anselm College have permission to print this material for their lecture notes.