variation in maternal care - between species - within species parent-offspring conflict mothering...
TRANSCRIPT
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Variation in maternal care- Between species- Within species
Parent-offspring conflict
MotheringCercopithecines
Baboon mother and infant
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Park Ride
Carried
Cling
Nest
Tree
Variation between speciesStyle (park/ride)
The main mammal pattern
(rodents, pigs, ungulates, etc)
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Park (nest, branch)
Park + Ride
Ride
Ancestral state unknown
Ross (2001) IJPTarsius
CeboidsCercopithecoidsHumanApes
Daubentonia
Indriidae
Lepilemur
Tree-shrews
Loris
Lorisoids
Galago/oides
Euoticus, Otolemur
Mouse lemurs
Phaner
Varecia
LemurHapalemur
Eulemur
1. Ancestral state = Parking
2. Riding evolved often
3. Riding is maintained
Riding is a repeated primate trait.
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Park (nest, branch)
Park + Ride
Ride
Ancestral state unknown Ross (2001) IJP Tarsius
Daubentonia
Indriidae
Lepilemur
Tree-shrews
Loris
Lorisoids
Galago/oides
Euoticus, Otolemur
Mouse lemurs
Phaner
Varecia
Hapalemur
Apes
Cercopithecoids Ceboids
Lemur
Eulemur
Riding: > 4 independent appearances
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Park (nest, branch)
Park + Ride
Ride
Ancestral state unknown
Ross (2001) IJPTarsius
CeboidsCercopithecoids
Apes
Daubentonia
Indriidae
Lepilemur
Tree-shrews
Loris
Galago/oides
Mouse lemurs
Varecia
Lemur
Hapalemur
Human
Lorisoids
Euoticus, Otolemur
Phaner
Park+Ride:
> 2 from Park*
> 1 from Ride+
*
*
+
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Costs (allometric) of riding, cf. parking:
(1) Smaller home ranges(2) Late weaning(3) Late age of first reproduction
Riding compared to parking:
(1) Same Inter-birth interval(2) Same Growth rate(3) Same Infant size(4) Same Brain size
Retrieving parked infant Arctocebus
So why has riding evolved?
Ho: Improved infant survival
Ross (2001) IJP
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Riders: Contact time similar in different species
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Mothers carry offspring more when needed
Altmann & Samuels 1992 BES
Kanyawara Field Station (Photo AH)
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Mothers are important for survival!
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… even when not dead
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Mothers are important for RS!
Large group
Small group
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Variation within species1. Ecological conditions:
Food availability
Gilgil
Amboseli
Drakensberg
High-quality
Low-quality
% time spent feeding by mother
………… Expected
Observed
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Lycett et al 1998 BES
P. c. ursinus Chacma
Harsh conditions:
Infants forage less independently (closer to mother)
Age
% Time
Good habitat
Intermediate habitat
Bad habitat
Twice more independent in a good habitat compared to a bad one !
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Lycett et al 1998 BES
P. c. ursinus Chacma
Drakensburg
Harsh conditions:
In harsh environment, mothers work harder.
(infants suckle more)
{ 5x diff.
Good habitatIntermediate habitatBad habitat
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Variation within speciesEcological conditions:
Predation Risk
No predation
Some prednMother’s hard work pays!
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Variation in maternal care3. Dominance rank / style
High-rank
mothers
carried
less.
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Mother’s Style
Laissez-faire Restrictive
Mother’s rank High Low
Infant allowed to leave < 2 months Later
M follows I Rare Common
M joins I Rare Common
M punishes I Less More
M-I contact Less More
Altmann (1980)
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Variation in maternal care
Maternal style
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Parent-Offspring conflict: OBSERVER’S Perspective
B = Benefit to Offspring(e.g. 10 cc of milk)
C = Cost to Mother(e.g. extra daysbefore next birth)
Age of Offspring
B/C
WEANING CONFLICTAGREED: M CARES
AGREED: M STOPS
B=C
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When to wean? MOTHER’S Perspective
B/2 = Benefit to Mother
? ??
B = Benefit to Offspring
C = Cost to Mother
Age of Offspring
B/C
Wean when “rB < C”i.e. when B/2 < C
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C/2 = Cost to Offspring
When to wean? OFFSPRING’S Perspective
B = Benefit to Offspring(e.g. 10 cc of milk)
C = Cost to Mother
Age of Offspring
B/C
Wean when “B < rC”i.e. when B < C/2
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Wean when “B < rC”i.e. when B < C/2
C/2
B
Age of Offspring
B/2C
Age of Offspring
Wean when “rB < C”i.e. when B/2 < C
“EARLY!!” “LATE!!”CONFLICT!!!
WHY PARENTS AND CHILDREN FIGHT
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Parent-offspring conflict
Infant age (weeks)
3030
25
50
Cayo Santiago
(feral, subtropic)
Madingley
(cages, UK)
% total nipple contacts
(attempted + successful) rejected by
mother
Mothers do reject young
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Infant age (weeks)
3012
Relative Responsibility Index: RRI
RRI = (% M-I body contacts initiated by infant) -
(% M-I body contacts terminated by infant)
RRI
0
25
50
25
50
I joins and/or M leaves
M joins and/or I leaves
Who’s responsible for declining maternal investment?
Mother’s investment falls
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Gelada:
More infant contact during
RESTING SOCIALIZING
than during
FEEDING MOVING
Contact MOTHER
Contact NIPPLE
Infants seek investment mostly when mother is not busy
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Infant age (months)
987654321
Re
jectio
ns p
er
min
of
activity
.040
.030
.020
.010
0.000
Context
Feeding
Socializing
Gelada: Rejections occur almost entirely when mother is feeding
No rejections were observed during
moving and resting
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Time on nipple
Ventral + Nipple
Suckling time decreases little!
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van Schaik & van Noordwijk (1999) Primates
Mothers need to wean early to increase their lifetime RS
Age-specific fecundity Macaques (3 species)
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Parent-offspring conflict:
Amount versus Timing
Barrett et al 1995 Anim Behav
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PlayFightingAllomotheringLearning
Juvenile relationships
Childhoodin
Macaques
Macaca fuscata
Japanese macaque
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Benefits of Play: cf. going to school
Benefit to young: Skills?
Lone play M = F
Play-fighting M > F NHP’s
Play-mother F > M NHPs’
Tool-use F > M Tai, Gombe
Social manipulation?
Mother distracting Infant (weaning)
Learning competition
Early coalitions/relationships?
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Benefits of Play: cf. going to schoolLone play M = F
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Benefits of Play: cf. going to schoolPlay-mother F > M NHPs’
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Benefits of Play: cf. going to schoolPlay-fighting M > F NHP’s
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Benefits of Play: cf. going to schoolTool-use F > M Tai, Gombe
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Rainfall
% play-time
Economics of play
(1) Benefit
More play when more food
Some primates never play!
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Economics of play(2) Cost
Vervets eaten when playing – Red-tailed infant playing at the very top of a fruit tree (Kibale) caught by eagle
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Vervet allomothers (captive)
Who?
Juvenile ff 65
(sibs; high-rank)
Adult ff 25
Juvenile mm 7
Adult mm 3
Benefit to mother
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Why be an alloparent?Females - nulliparous
*** Practice: improved mothering ** Nepotism: caring for kin * Market forces: care for HR’s offspring (get support in return)
Females - parous (e.g. langurs) * Nepotism * Non-kin: odd
suggesting friendship?
Males - juvenile * Nepotism * Practice mating * Market forces: care for HR’s offspring (get support in return)
Males - adult ** Paternal care*** Mating effort
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Wallen et al. (1995) Hormones and Behavior
14
7
0
DaughterSon
Yearling-controlled
proximity to mother
(% time)
Girls spend more time with mother than boys
Rhesus macaque
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Wallen et al. (1995) Hormones and Behavior
14
7
0
Daughter
Sons
Yearling-controlled
proximity to mother
(% time)
Neonatal T affects how much time boys spend with mother
T control T T
Rhesus macaque
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Lonsdorf et al 2004 Nature
% time at mound spent termite-fishing: girls > boys
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Termites per dip
Lonsdorf et al 2004 Nature
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Mother’s % dips at given tool length
Daughter’s % dips at given tool length
Lonsdorf et al 2004 Nature
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Mother’s % dips at given tool length
Son’s % dips at given tool length
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Juvenile relationships: anticipating adulthood
Dominance relations in FB-groups
Juv F Juv M
Juv F Maternal Rank Size
Juv M Size Size
e.g. Pereira, Amboseli baboons
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Juvenile relationships: anticipating adulthood
Infant Juvenile Adult
Pereira, Amboseli baboons
0 30 (months) 60
# Supplants received per hour from HR females Female
Male
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Juvenile relationships: anticipating adulthood
Juv F Juv M
Challenged by Adult FF YES NO
Supported by Adult FF YES NO
‘Head-flag’ to Adult FF YES NO
“ “ Adult MM NO YES
Non-kin supported HR-FF Juv MM
Pereira, Amboseli baboons
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Baboon rules for treating juveniles
Adult FF
Juvenile FF: support their rank
(winner support)
Juvenile MM: dominate as long as possible
Adult MM
Ignore sex/rank (dominate all)
Support possible offspring
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Old view New view
Juvenile Passive Active
Juv period Long, to learn Fast growth, risky
Major influences Proximate Ultimate
(experience) (social system)
Quality of life Relaxed Dangerous (>50% die)
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23 FF, 2 groups, 240 cases
90% held I’s = higher-rank
F > M
Kin > Unrelated
De Waal 1990 Primates
“Double-holding”:
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Play
Play partners
Infants Same size
HR preferred
Juv FF Infants (HR preferred)
Juv MM Juv MM