Lecture 11 - Mating Systems and Parental Care
Learning Objectives
- Describe the full spectrum of mating-system classifications.
- Understand ecological drivers that link resource distribution to mating strategies.
- Explain the diversity, costs, benefits, and evolution of parental care.
Key Concepts & Definitions
- Anisogamy
- Females produce few, large, energetically expensive eggs.
- Males produce many, tiny, energetically cheap sperm.
- Sets up male–male competition and potential for males to mate with several females ⇒ polygyny.
- Sexual selection acts more strongly on the sex with the higher variance in reproductive success (usually males).
- Polygyny threshold
- Minimum territory or male-quality difference at which a female gains higher fitness joining an already-mated male than pairing with an unmated male of lower quality.
- Philopatry – return to natal site for breeding; prominent in jacanas.
- Iteroparity vs. Semelparity
- Iteroparity: repeated breeding at intervals.
- Semelparity: breed once explosively then die (e.g., coho salmon).
- r– and K-selection
- r ≈ intrinsic reproductive rate; K ≈ carrying capacity.
Spatial Distribution of Resources & Mating Systems
- Fig 18.1 demonstrates that where food is patchy and predators present, large groups form; multiple mating for both sexes likely ⇒ polygyny.
- Dense forests hinder signalling → monogamy becomes adaptive because finding multiple mates is costly.
Classification by Social Association
• Monogamy – 1 ♂ : 1 ♀
• Polygyny – 1 ♂ : ≥2 ♀♀
• Polyandry – 1 ♀ : ≥2 ♂♂
• Promiscuity – no prolonged bonds; multiple mating by ≥1 sex.
• Polygamy (generic) – umbrella term for all non-monogamous systems.
Ecological (Operational) Classification
- Monogamy – neither sex can monopolise >1 mate.
- Polygyny
- Resource-defence
- Female-defence
- Male-dominance (lek)
- Polyandry
- Resource-defence polyandry
- Female-access polyandry (♀♀ limit access to ♂♂ without defending resources)
Monogamy
- Occurs when renewable resources are scarce & widely spaced.
- Benefits
- Mate assistance in rearing offspring.
- Reduced search time; long-term pair bonds.
- Sea-gulls: re-pairing with former mate ⇒ higher success (less aggression; synchronised courtship).
- Stats: ≈90\% of bird species are socially monogamous.
- Small social units reduce predation risk.
Polygyny
Resource-Defence Polygyny
- ♂ defends critical sites (feeding / nesting).
- Female choice depends on territory quality.
- Example: walnut flies; variable territory quality explains polygyny threshold.
Female-Defence Polygyny
- Females are already gregarious (e.g., seals limited to haul-out sites).
- ♂♂ guard clusters of females; extreme male–male competition and high variance in male fitness.
Male-Dominance Polygyny (Lekking)
- ♂ provides no resources or care.
- ♂♂ aggregate in leks = small defended display arenas.
- Females visit, choose, copulate, depart to rear young alone.
- No direct benefits; indirect (good genes) or copying.
- Hammer-head bats: 6\% of ♂♂ obtain 79\% of copulations.
- Ungulates:
- Topi: large ♂♂ solo territories most successful; smaller ♂♂ cluster in leks.
- Uganda kob: opposite pattern—lek males outperform solitary ones.
Scramble Competition Polygyny
- No territories or dominance.
- Explosive breeding assemblages: frogs in temporary ponds; horseshoe crabs intercept females on shore.
Polyandry
- Rarer because egg investment > sperm.
- Conditions favouring: unpredictable food, high nest predation ⇒ females lay multiple clutches while ♂♂ incubate.
- Birds with male-only incubation (e.g., spotted sandpipers, jacanas).
Case Study: American Jacana
- ♂ territories small; ♀ super-territories encompass several ♂ nests.
- ♀ 50 % larger; provide little direct care; dominance over ♂♂.
- Functional reversal of polygyny: ♀ specialise in egg production.
Ecology & Mating: Blackbird Case Study
- Red-winged blackbird (polygynous)
- Stable, long-lasting food (seeds/insects).
- ♂ returns early, defends territory for months.
- ♂ provides no care; ♀ builds nest & feeds young.
- Tricolored blackbird (nomadic, quasi-monogamous pairs within huge colonies)
- Food in temporary, concentrated patches.
- Colony relocates once food depleted; synchronous, rapid breeding (≤1 wk).
- Both sexes invest heavily but over short period.
- Conclusion: resource temporal stability shapes social system.
Alternative Reproductive Tactics (ARTs)
Coho Salmon
- Two irreversible life-history tactics chosen at ~6 mo:
- Hooknose: large, red, weaponised; tactic = fight.
- Jack: small, early-maturing; tactic = sneak.
- Reversible behavioural tactics while on spawning ground: fight or sneak (matrix in Fig 18.8).
Marine Isopod (Paracerceis sculpta)
- Three male morphs with equal lifetime success:
- Alpha: large, defend sponge harems.
- Beta: medium, female mimics; gain coaxial matings.
- Gamma: tiny, mobile sneakers; infiltrate harems.
Parental Care Spectrum
- Increases with organismal complexity.
- Extremes:
- Broadcast spawning fish/invertebrates ⇒ no care.
- Primates: up to 25\% of offspring lifespan invested.
- Birds: diverse strategies
- Megapodes bury eggs in compost heaps for heat.
- Penguins fast & incubate without feeding.
Ecological Determinants of Parental Effort
- Reproductive effort = energy + risk now that reduces future reproduction.
- Stable environments (K-selected)
- Large body, slow development, low fecundity, high care.
- Home ranges; competition intense; strategy — quality over quantity.
- Fluctuating environments (r-selected)
- Rapid dev., high fecundity, low care.
- Populations shaped by abiotic mortality.
- Coho salmon: huge pre-spawning costs ⇒ semelparity; death after breeding.
- Scarce/difficult food promotes prolonged dependency for learning.
Which Sex Should Provide Care?
Confidence of Parentage (Trivers 1972)
- Internal fertilisation ⟹ male uncertainty; may desert.
- Female certain ⇒ higher likelihood to stay.
Association with Embryos (Williams 1975)
- Care evolves in sex physiologically linked to embryos.
- Internal fert. & gestation: favours female care.
- External fert. in male territory: favours male care.
- Teleost data (Table 18.1):
- Internal fertilisation families – mostly female care (14 vs 2 ♂).
- External fertilisation families – male care dominates (245 families).
Taxon-Specific Rules
- Birds: bi-parental care common; males incubate, feed, even secrete crop milk (pigeons).
- Mammals: gestation/lactation restrict direct male care; hence polygyny common, especially in precocial species. Altricial young require more biparental input.
- Insects show extremes from none to male egg-brooding (water bugs, Fig 18.13).
Differential Investment Curve (Fig 18.10)
- Early: male territory defence costly > female.
- Mid: egg production swings cost to female.
- Later: male incubation > female.
- Implication: dynamic cost‐benefit influences mate desertion decisions.
Example: Elephant Seals
- Highly polygynous colonies.
- Sexual dimorphism: ♂ ~3× size; enlarged proboscis & chest shield.
- Dominance hierarchy; <\tfrac13 males copulate.
- Zero paternal care; pups can be trampled inadvertently during copulations.
- Graph (Fig 18.12) shows positive relationship between dominance rank & copulatory success.
- Prediction: males at lower left (low rank) likely young/subordinate; upper right are older, larger, high-status.
Biparental & Male-Biased Examples in Mammals
- Silver-backed jackals: both sexes hunt & defend territory cooperatively.
- Tamarins/marmosets: males carry neonates immediately after birth ⇒ maternal feeding efficiency.
Parent–Offspring Recognition
- Adaptive to avoid misdirected care.
- Modal cues vary:
- Birds: vocal signatures; cliff swallows have complex individual calls vs barn swallows (Fig 18.15) due to colony size differences.
- Dolphins: individual whistle signatures used for mother–offspring & group recognition.
Parent–Offspring Conflict (Trivers 1974)
- Ratio of cost to mother / benefit to offspring increases with offspring age (Fig 18.16).
- <1 ⇒ mutual benefit.
- 1–2 ⇒ conflict zone; offspring still benefits, mother fitness declining.
- >2 ⇒ both lose; offspring weans.
Ethical & Evolutionary Implications
- Understanding mating systems informs conservation (lek species vulnerable to over-harvest of top males).
- ARTs illustrate that multiple strategies can be evolutionarily stable (isopod morphs equalised fitness).
- Human parallels: resource distribution & parental investment theories contribute to anthropology and social policy debates.