Behavioral Ecology Notes
BEHAVIORAL ECOLOGY - ALTRUISM
Definition of Altruism
- An act is altruistic if it reduces the actor's fitness (cost, ) while increasing the recipient's fitness (benefit, ).
Hamilton's Rule & Inclusive Fitness
- Inclusive fitness = direct fitness (your own reproduction) + indirect fitness (reproduction of relatives, weighted by relatedness ).
- Hamilton's Rule: r \times B > C,
- where:
- : coefficient of relatedness
- : benefit to the recipient
- : cost to the actor
- where:
Kin Selection
- Genes promoting altruism spread when directed toward relatives.
- Example: Ground squirrels give alarm calls to warn kin of predators.
Eusocial Insects: Extreme Altruism
- Sterile worker castes care for queen's offspring.
- Haplodiploidy (ants, bees, wasps) sisters share .
- Raising sisters can yield higher indirect fitness than having own offspring.
Reciprocal Altruism
- Helping non-kin with expectation of returned favor.
- Conditions (Trivers 1971): repeated interactions, recognition & memory, B > C, punishment of cheaters.
- Example: Vampire bats share blood meals with roost-mates.
Alternative Frameworks
- Multilevel selection: interplay of individual- and group-level benefits.
- Groups of cooperators may outcompete selfish groups under certain conditions.
Medical-Biology Connections
- Apoptosis: programmed cell death eliminates damaged cells (actor cost) for organism's health (benefit).
- Microbial Public Goods: bacteria release enzymes that benefit colony; cheater dynamics influence infection & resistance.
Key Takeaways
- Altruism can evolve via kin selection (r \times B > C) or reciprocity.
- Relatedness and repeated interactions are crucial.
- Principles apply across taxa-from squirrels to cells.
BEHAVIORAL ECOLOGY - LIVING IN GROUPS
What Is Group Living?
- Definition: Repeated, close-proximity association among conspecifics with coordinated or emergent behaviors.
- Ubiquity: Occurs in all major animal taxa—from planktonic swarms to primate troops to eusocial insects.
Benefits of Group Living
- Predator Defense
- Many-Eyes Effect: Shared vigilance lowers individual scanning effort.
- Dilution Effect: Per-individual predation risk decreases as group size increases.
- Cooperative Foraging & Information Sharing
- Local Enhancement: Observing foragers leads naïve individuals to resources.
- Division of Labor: Specialists (e.g., scouts vs. harvesters) increase efficiency in social insects.
- Thermoregulation
- Huddling conserves heat; e.g., emperor penguins rotate positions in dense clusters.
- Reproductive & Kin-Selection Advantages
- More mating opportunities in aggregations.
- Helping close relatives enhances inclusive fitness.
- Predator Defense
Costs of Group Living
- Resource Competition: More individuals compete for limited food, space, or mates.
- Disease Transmission: Close contact facilitates parasites and pathogens.
- Increased Detectability: Large groups can be more conspicuous to predators.
- Social Conflict: Hierarchies and aggression can stress lower-ranked members.
Types of Group Structures
- Aggregations
- Characteristics: Temporary, loose-often in response to external stimuli
- Example: Fish schooling when threatened
- Colonies
- Characteristics: Site-attached, long-term
- Example: Seabird rookeries; bat caves
- Eusocial Societies
- Characteristics: Sterile worker castes; reproductive division
- Example: Ant, bee, termite colonies
- Fission-Fusion
- Characteristics: Fluid membership; split and rejoin
- Example: Chimpanzee communities; some deer
- Aggregations
Key Theoretical Models
- Selfish Herd Theory (Hamilton, 1971): Individuals move toward group center to minimize personal predation risk.
- Ideal Free Distribution: Animals distribute among resource patches to equalize individual gain.
- Game Theory: Payoff matrices predict when cooperation or defection (group joining or leaving) maximizes fitness.
Case Studies
- Meerkats (Suricata suricatta)
- Sentinel system: Individuals take turns watching for predators while others forage.
- Cooperative care: Helpers feed and guard pups, boosting group survival.
- Honey-Bee Colonies (Apis mellifera)
- Waggle dance communicates direction and distance of flower patches.
- Caste system: Workers, drones, and queens specialize in tasks.
- Mixed-Species Flocks (Neotropical Birds)
- Species with different foraging niches flock together.
- Collective vigilance yields lower individual predation risk.
- Meerkats (Suricata suricatta)
Balancing Costs & Benefits
- Adjustable Group Size: Individuals join or leave groups based on predation risk and resource availability.
- Behavioral Synchrony: Coordinated movement and signals maintain cohesion (e.g., flock turns).
- Social Immunity: Collective grooming or antimicrobial secretions reduce disease spread in eusocial insects.
Implications Beyond Ecology
- Human Sociality & Epidemiology: Insights into crowd behavior, disease outbreaks, and public-health strategies.
- Conservation & Management: Understanding grouping can inform species reintroduction, habitat design, and wildlife corridors.
Key Take-Home Messages
- Group living evolves when its net benefits (anti-predator, foraging, thermoregulation, kin selection) exceed net costs (competition, disease, detectability, conflict).
- A spectrum of group structures exists-each shaped by ecological pressures and life-history trade-offs.
- Theoretical models (Selfish Herd, Ideal Free, Game Theory) help predict when and how animals should form, maintain, or leave groups.
BEHAVIORAL ECOLOGY - RESOURCE ASSESSMENT
Why Resource Assessment Matters
- Animals have limited time, energy, and safety. To survive and reproduce they must decide:
- Resource Value (RV) - How valuable is this item right now?
- Resource-Holding Potential (RHP) - How likely am I to win or defend it?
- Accurate answers reduce wasted effort and injury.
- Animals have limited time, energy, and safety. To survive and reproduce they must decide:
Key Definitions
- Resource Value (RV)
- Plain-English Meaning: Immediate fitness payoff of winning the item
- Typical Measurement or Example: Fat in a seed; number of eggs a nest can support
- Resource-Holding Potential (RHP)
- Plain-English Meaning: Fighting ability relative to rivals
- Typical Measurement or Example: Body size in beetles; antler span in deer
- Signal
- Plain-English Meaning: Trait or action that conveys information about RV or RHP
- Typical Measurement or Example: Roars, plumage colour, chemical scent
- Evolutionarily Stable Strategy (ESS)
- Plain-English Meaning: Strategy (or mix) that cannot be invaded by an alternative once common in the population
- Typical Measurement or Example: Hawk-Dove mix when injury cost > resource value
- Resource Value (RV)
Contest Assessment Strategies
- Self-Assessment - Individual quits when its own energy or risk threshold is reached.
- Example: Salmon stop fighting when aerobic reserves are depleted.
- Mutual (Sequential) Assessment - Rivals exchange increasingly risky signals, quitting if clearly outmatched.
- Example: Red-deer stags often settle contests by roaring alone.
- Cumulative Assessment - Loser quits when total costs inflicted by the opponent exceed its limit.
- Example: Field crickets accumulate scrape damage until one retreats.
- Reality: Species may blend strategies depending on context.
- Self-Assessment - Individual quits when its own energy or risk threshold is reached.
Classic Theoretical Models
- Hawk-Dove Game (Maynard Smith)
- Core Idea: Individuals choose to fight (Hawk) or display (Dove).
- Key Prediction: When injury cost > resource value , a mixed ESS emerges.
- Real-World Evidence: Speckled-wood butterflies: more Hawks over high-sunlight spots.
- War of Attrition (Parker & Thompson)
- Core Idea: Contestants avoid dangerous combat and "out-wait" each other with costly displays.
- Key Prediction: Contest length is highly variable; longer if resource value is high.
- Real-World Evidence: Fiddler-crab waving bouts vary widely in duration.
- Sequential Assessment Model (Enquist & Leimar)
- Core Idea: Rivals sample RHP step-by-step, escalating risk only when necessary.
- Key Prediction: Contest duration increases sharply as RHP difference shrinks.
- Real-World Evidence: Cichlid fish mouth-wrestling lasts longest in size-matched pairs.
- Cumulative Assessment
- Core Idea: Quit when cumulative costs received reach a personal threshold.
- Key Prediction: Losers show higher immediate injury or energy debt than winners.
- Real-World Evidence: Loser crickets have greater scrape damage after pushing bouts.
- Hawk-Dove Game (Maynard Smith)
Signals: Keeping Information Honest
- Costly to fake: Deep croaks require large bodies; small toads cannot mimic.
- Social policing: Paper wasps attack rivals with "cheater" facial patterns.
- Handicap principle: Honest signals persist because they impose real costs on high-quality senders.
Beyond Fighting
- Foraging
- Assessment Rule: Marginal Value Theorem - leave a patch when current intake < average elsewhere
- Example: Bumblebees track nectar refill rates
- Territory
- Assessment Rule: Ideal Free Distribution - individuals distribute so none improves payoff by moving
- Example: Ducks among feeding ponds
- Mate Choice
- Assessment Rule: Sample and assess potential partners; often apply "best-so-far" rule
- Example: Female birds ranking male song complexity
- Foraging
Take-Home Points
- Define your terms - always translate acronyms back to real-world meanings.
- Assessment saves resources - wrong decisions waste energy or cause injury.
- Models guide hypotheses - each predicts specific, testable patterns.
- Same logic, diverse settings-feeding, mating, territoriality all rely on cost-benefit assessment.