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systems and parental investment Mating systems

Pattern of in a population

green anole

Antithesis =

Polygyny Polygyny

Scramble: no attempts to defend females, resources horseshoe crabs

Northern barred frog Female defense: must be clustered elk Montezuma’s oropendola

Dulichiella spp.

Polygyny Resource distribution

Resource defense: males defend food, nest sites Distribution of females affects Red-winged blackbird Lamprologus cichlid males’ ability to guard them

Males cannot monopolize wide-ranging females

dunnock

1 Polygyny threshold Polygyny threshold

Male with no other females () Male with other female(s)

polygyny threshold

??? Quality of male’s territory

Polygyny threshold Male dominance polygyny

When females and sage Polygyny threshold = point at which it’s resources too dispersed, better to be polygynous on a good territory males compete

Leks = communal display arenas

hammerhead bat Uganda kob

Leks Leks

High variance in male mating success – 10-20% males achieve >50% copulations – one male got 75% copulations

Classical lek: males display in sight of each other

Exploded lek: males rely wire-tailed manakin on vocal communication, e.g. kakapo

2 Leks Leks

• Hotshots • Hotshots – Females attracted to lek by dominant male – Females attracted to lek by dominant male • Hotspots – Leks located in high-use areas

Leks Leks

• Hotshots Position of most successful – Females attracted to lek by dominant male male territory shifts (hot shot?) • Hotspots black grouse – Leks located in high-use areas • Female preferences – Females prefer large groups of males

Removal of dominant male may cause lek to shift, disband

Leks Paradox of the lek

Lek placement varies with respect to female density (hot spot??) How is genetic variation maintained?

Hamilton Rowe and Zuk and Houle

3 Paradox of the lek

• Usually characterized by If females mate once, why do leks form? -role reversals • Factors favoring polyandry include – Male-biased sex ratio Wilson’s – Precocial offspring jacana phalarope – Superabundant food

Polyandry Monogamy

Given sex differences in gamete production, why monogamy?

spotted sandpiper

x

Clutch manipulations Selects for production reveal incubation of two clutches constrained at four eggs

Monogamy Monogamy

Occurs in where females are widely distributed, rarely encountered

Duration may be variable

Rare in mammals (<5%) mantis shrimp common in birds (>90%)

4 Mate assistance Mate assistance

Important when young are Important where habitat is energetically demanding harsh or offspring survival is low

cotton-top tamarins emperor penguins

Female + male Female only

Female enforcement Extra-pair copulations

Conflict of interests between the regarding In some species, >50% offspring are extra-pair young care of offspring, opportunities for additional matings 9 8 Female untethered 7 Female tethered 6 5 4 3 2 1 Number of broods of Number 0

0.0 - 0.10 0.91 - 1.0 burying beetle 0.11 - 0.20 0.21 - 0.30 0.31 - 0.40 0.41 - 0.50 0.51 - 0.60 0.61 - 0.70 0.71 - 0.80 0.81 - 0.90 superb fairy wren < 6 min > 6 min Proportion of extra-pair young (EPY) Duration of male pheromone emission 50-90% EPY (avg. 78%)

Why do females solicit EPCs? Why do females solicit EPCs?

Material benefits? Material benefits?

scorpionfly • EPCs are less “institutionalized” version of polyandry

• Many of the same benefits apply

Adelie penguins get rocks from EPC partners

5 Why do females solicit EPCs? Why do females solicit EPCs?

Male defense Genetic benefits …against nest predators …against infanticidal males – Guard against poor sperm quality of mate

red-winged blackbird Hanuman langur 32% on females with EPY 54% predation on females with no EPY Females may mate often pseudoscorpion to confuse paternity

Why do females solicit EPCs? Why do females solicit EPCs?

Genetic benefits Genetic benefits – Guard against poor sperm quality of mate – Guard against poor sperm quality of mate – Increase genetic diversity of offspring – Increase genetic diversity of offspring – Mate with genetically superior male

bluethroat

blue tit

EPCs in humans Parental investment

Population % nonpaternity southern (!Kung) 2 rural Michigan, USA ~10 Munich and Copenhagen >10 Italy 13.2 15.3 urban Michigan, USA 20.1 Liverpool, UK 20-30

6 Parental investment Parental investment

Behavior that increases Nutrition in egg fitness of offspring at cost to Pre-hatching (nest site, protecting eggs) Post-hatching investment is rare

Post-hatching (provisioning, lowland gorilla protecting young)

Patterns of parental investment Patterns of parental investment

mallee fowl • Female-only care predominates • Male, biparental care is rare =

How do we account for the general female bias golden- tamarin in parental care?

Richard Dawkins Logic suffers from “sunk cost” or What about the abrupt “Concorde” fallacy termination of investment in polyandrous species?

Patterns of parental investment Importance of paternity

Explained by order of gamete release? 9 8 7 6 Extra-pair Mode of fertilization 5 4 paternity 3 2 1 may select for Provider sex Internal External broods of Number 0 decreased guppy 0.0 - 0.10 0.91 - 1.0 tree 0.11 - 0.20 0.21 - 0.30 0.31 - 0.40 0.41 - 0.50 0.51 - 0.60 0.61 - 0.70 0.71 - 0.80 0.81 - 0.90 male care Male Few Many piranha swallow Proportion of extra-pair young (EPY)

Female Many Few If costs to male and female to rear young are the same, but only 3 of 4 chicks are his… How to explain the predominance of male He should reduce his care in frogs where males deposit sperm first? dark-eyed level of investment junco

7 Recognition of unrelated young? Recognition of unrelated young?

No apparent discrimination against EPY Alpha, beta male dunnocks care for young in proportion to time spent with female male female

Male tree swallows show Male yellow warblers do no feeding bias to own Females may use mating not adjust feeding rate chicks, feed those closest solicitations to manipulate male in response to cuckoldry or most in need parental investment

Experimental = male Recognition of unrelated young? Control = male bluegill on nest with bluegill on nest “sneaker” in a jar Why the difference between care of eggs and care of fry?

Tested defensive responses to nest predator

Sex differences in costs of investment Sex differences in costs of investment

St. Peter’s fish giant water bug Male care persists where costs to females exceed costs to males?

Male care depends on costs/benefits to male ~11 days ~7 days

Large eggs require high level of care

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