Variation in maternal care - Between species - Within species Parent-offspring conflict

1. Mothering Cercopithecines Variation in maternal care - Between species - Within species Parent-offspring conflict Baboon mother and infant

2. Variation between species Style (park/ride) Park Ride The main mammal pattern (rodents, pigs, ungulates, etc) Carried Nest Tree Cling

3. Tree-shrews Park (nest, branch) Park + Ride Ride Ancestral state unknown Riding is a repeated primate trait. Lorisoids Loris Euoticus, Otolemur Galago/oides 1. Ancestral state = Parking Mouse lemurs Phaner 2. Riding evolved often Varecia Eulemur 3. Riding is maintained Hapalemur Lemur Lepilemur Daubentonia Indriidae Apes Human Cercopithecoids Ceboids Tarsius Ross (2001) IJP

4. Tree-shrews Park (nest, branch) Park + Ride Ride Ancestral state unknown Riding: > 4 independent appearances Lorisoids Loris Euoticus, Otolemur Galago/oides Mouse lemurs Phaner Varecia Eulemur Hapalemur Lemur Lepilemur Daubentonia Indriidae Apes Cercopithecoids Ceboids Ross (2001) IJP Tarsius

5. Tree-shrews Park (nest, branch) Park + Ride Ride Ancestral state unknown Lorisoids Park+Ride: > 2 from Park* > 1 from Ride+ * Loris Euoticus, Otolemur Galago/oides Mouse lemurs Phaner * Varecia Hapalemur Lemur Lepilemur Daubentonia Indriidae Apes Human + Cercopithecoids Ceboids Tarsius Ross (2001) IJP

6. Riding compared to parking: (1) Same Inter-birth interval (2) Same Growth rate (3) Same Infant size (4) Same Brain size Costs (allometric) of riding, cf. parking: (1) Smaller home ranges (2) Late weaning (3) Late age of first reproduction So why has riding evolved? Ho: Improved infant survival Retrieving parked infant Arctocebus Ross (2001) IJP

7. Riders: Contact time similar in different species

8. Mothers carry offspring more when needed Kanyawara Field Station (Photo AH) Altmann & Samuels 1992 BES

9. Mothers are important for survival!

10. … even when not dead

11. Mothers are important for RS! Large group Small group

12. % time spent feeding by mother ………… Expected Observed Gilgil High-quality Variation within species 1. Ecological conditions: Food availability Amboseli Low-quality Drakensberg

13. Harsh conditions: Infants forage less independently (closer to mother) % Time Age Goodhabitat Intermediatehabitat Badhabitat P. c. ursinus Chacma Twice more independent in a good habitat compared to a bad one ! Lycett et al 1998 BES

14. { 5x diff. Harsh conditions: In harsh environment, mothers work harder. (infants suckle more) P. c. ursinus Chacma Drakensburg Badhabitat Intermediatehabitat Goodhabitat Lycett et al 1998 BES

15. Variation within species Ecological conditions: Predation Risk No predation Mother’s hard work pays! Some predn

16. Variation in maternal care 3. Dominance rank / style High-rank mothers carried less.

17. Mother’s Style Laissez-faireRestrictive 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)

18. Variation in maternal care Maternal style

19. B=C B/C B = Benefit to Offspring (e.g. 10 cc of milk) Age of Offspring C = Cost to Mother (e.g. extra days before next birth) AGREED: M STOPS AGREED: M CARES WEANING CONFLICT Parent-Offspring conflict: OBSERVER’S Perspective

20. When to wean? MOTHER’S Perspective ? ? ? B = Benefit to Offspring B/C C = Cost to Mother B/2 = Benefit to Mother Age of Offspring Wean when “rB < C” i.e. when B/2 < C

21. When to wean? OFFSPRING’S Perspective B/C B = Benefit to Offspring (e.g. 10 cc of milk) C = Cost to Mother Age of Offspring C/2 = Cost to Offspring Wean when “B < rC” i.e. when B < C/2

22. B/2 C B Age of Offspring Wean when “rB < C” i.e. when B/2 < C Wean when “B < rC” i.e. when B < C/2 C/2 Age of Offspring “LATE!!” “EARLY!!” WHY PARENTS AND CHILDREN FIGHT CONFLICT!!!

23. Parent-offspring conflict Mothers do reject young 50 Cayo Santiago (feral, subtropic) % total nipple contacts (attempted + successful) rejected by mother Madingley (cages, UK) 25 0 3 30 Infant age (weeks)

24. Who’s responsible for declining maternal investment? Relative Responsibility Index: RRI RRI = (% M-I body contacts initiated by infant) - (% M-I body contacts terminated by infant) 50 RRI I joins and/or M leaves 25 0 12 30 25 Infant age (weeks) M joins and/or I leaves 50 Mother’s investment falls

25. Contact MOTHER Infants seek investment mostly when mother is not busy Contact NIPPLE Gelada: More infant contact during RESTING SOCIALIZING than during FEEDING MOVING

26. Gelada: Rejections occur almost entirely when mother is feeding No rejections were observed during moving and resting

27. Suckling time decreases little! Time on nipple Ventral + Nipple

28. Age-specific fecundity Macaques (3 species) Mothers need to wean early to increase their lifetime RS van Schaik & van Noordwijk (1999) Primates

29. Parent-offspring conflict: Amount versus Timing Barrett et al 1995 Anim Behav

30. Childhood in Macaques Play Fighting Allomothering Learning Juvenile relationships Macaca fuscata Japanese macaque

31. 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?

32. Benefits of Play: cf. going to school Lone play M = F

33. Benefits of Play: cf. going to school Play-mother F > M NHPs’

34. Benefits of Play: cf. going to school Play-fighting M > F NHP’s

35. Benefits of Play: cf. going to school Tool-use F > M Tai, Gombe

36. Economics of play Benefit More play when more food % play-time Rainfall Some primates never play!

37. 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

38. Vervet allomothers (captive) Benefit to mother Who? Juvenile ff 65 (sibs; high-rank) Adult ff 25 Juvenile mm 7 Adult mm 3

39. 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

40. 14 Girls spend more time with mother than boys Yearling-controlled proximity to mother (% time) 7 Rhesus macaque 0 Daughter Son Wallen et al. (1995) Hormones and Behavior

41. T T 14 Neonatal T affects how much time boys spend with mother Yearling-controlled proximity to mother (% time) 7 Rhesus macaque 0 T control Daughter Sons Wallen et al. (1995) Hormones and Behavior

42. % time at mound spent termite-fishing: girls > boys Lonsdorf et al 2004 Nature

43. Termites per dip Lonsdorf et al 2004 Nature

44. Daughter’s % dips at given tool length Mother’s % dips at given tool length Lonsdorf et al 2004 Nature

45. Son’s % dips at given tool length Mother’s % dips at given tool length

46. 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

47. Juvenile relationships: anticipating adulthood # Supplants received per hour from HR females Female Male 0 30 (months) 60 Infant Juvenile Adult Pereira, Amboseli baboons

48. Juvenile relationships: anticipating adulthood • Juv FJuv M • Challenged by Adult FF YESNO • Supported by Adult FF YESNO • ‘Head-flag’ to Adult FF YESNO • “ “ Adult MM NOYES • Non-kin supported HR-FFJuv MM Pereira, Amboseli baboons

49. 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

50. 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)