Infanticide and reproductive restraint in the archetypical polygynous primate.

Behavior and evolution seminar series. 

UC Davis. 
Center for behavior evolution and culture
Primate mating system. Reproductive strategies with polygamist males.
Objective: present data and testing validity of two objectives of the male, using
models of the way reproductive distributions works in groups. Multi male and
multi female species. 

The speaker walks out of the podium and use hand gestures.
Dominance hierarchy. Rank in the hiearchy - use it to see who has priority for
food or sex. 

Rank and Reproductive success. What determines the extent of an alpha male --
Why doesn't the alphamales sire all the kids?

Darwainian Male
Classical sexual selection. Why?  Fleshed out and validated (built upon) by
Bateman and Trivers. This classical view is outdated. The reworking by Trivers
has perpetuated the victorian and darwinian view. Key maxims- MAles RS
increases linearly with mating. 

Colloquiak "keep on banging out kids"

Triver noted that males unburdened by parental care. Males are essentially free. As
a consequence, they should pursue maximum potential reproductive output: invest
in competitive effort. We can ball them down into the essence of a Darwinian male.
Boil it down that the key of prediction from Trivers work that males should
maximize effort at the point of fertilization/conception.

Sexual selection through inter male competition

size matters. Weaponry. Competition for fertilization continues even after the
mating.
Why is there red in the graph?
what is the standard conclusion? Constraints- simply extraneous forces to pursuing
a total reproductive monopoly.
More formally- culturalize views- that the males want to sire all the kids, stopped by
extraneous contro
constraints vs. compromise- in the middle is limited control.
Limited control- opportuniy costs.
Graph of Alpha Monopoly vs. Reproductive Synchrony. – limited control:
opportunity costs
Receptive periods of female primates are in days. Males have to guard the females.
These receptive periods can overlap. As a consequence to ensure, he’s going to
confine his own attentions to only one. Therefore, subordinates can then have at the
other girl. The higher the overlap, the lower the male dominance in reproductive.
Clear negative association in the graph.

Maybe the male number may limit the dominance- rationale is that if there is a
increasing number of male, more burden in dominance. Forced to relinquish access
to females if there are more male. Prediction: number of male challenger associated
negatively with monopoly. Limited control: competitor pressure. (another scatter
plot graph). This effect is not seen that much.
third and final constraint is the energetic cost. Graph bar. act of guarding takes high
energetic costs. Prediction is there must be a point when males cannot guard 2
females anymore. guarding does impact activity. this must still be circumstantial
though. no actual direct evidence though.

What is the problem? quick recap

dominant males. traditionally, people tend to account for this variance using
classical sexual selection. argue that males don’t sire all the kids because of the 3
costraints. indeed, they are constrainted.
Argument: the above is too generalized. it doesn’t look at the system on the ground.
It ignores infant mortality (infanticide), constraints on female care (females can’t
really protect their kids), adult male mortality (you should worry about if you died,
you can’t protect your kids or remake kids).
Total reproductive monopoly may not be adaptive.
Classical sexual selection theory is wrong!

Study site and subjects Papio hamadryas ursinus. Two troops. 11 years. in the de
hoop nature reserve.

“I’m setting my argument, setting the scene for…”

Alphas sire between 60-80%
good classical sexual selection
dominant’s jaw is much larger than mid-ranking or low-ranking.
The only group that have these dent adaptations in lower jaw- they grind their jaw
to sharpen teeth.
You have to be good to be alpha male. Need weapon. Baboons

Infancticide- bar graph on percent infants born and depending on the nature
reserve. Four populations… anywhere from 15-20% death.

females really can’t do anything. Males friendships are crucial to surviving. males
should pay attention.

Residency of usurped alphas – step graph –

male babies who didn’t have a male in the first parts of their infancy are more likely
to die. There is this deficit in care due to this absence.

Hypothesis: Alpha males concede conceptions to encourage subordinate residency

and, in turn, increase chances of additional protection. we know that in female
solution is to predispose other males to protect their offsprings. get multiple males
inclined to protect their infants.
As an alpha male, you need to present incentives to the subordinate males to
participate.

Male residency
suboridinate residency is in short-term basis. What you find is that subordinate
males will respond to short-term chances. Relatively unpredictable.

Predictions
1) subordinate reproductive success within groups I a function of more than
factors constraining the guarding behavior of dominant males
2) subordinate residency is mediated by reproductive success
3) presence of subordinates is associated with a reduction in the risk of
infanticide.
In contrast to Classical S S.

Limited Control: Oestrous overlap

all of those conceptions- number of infant sired compared with alphas and
subordinates. Only 8 are overlapped. out of about 30 infants sired. dominants didn’t
try to limit access of subordinates with mating female.
Maybe it’s due to competition. Duration of alpha guarding limited control: male
number in dot plot. No real relationship.

Operational sex ratio- limited control: competitior pressure dot plot. Again, no
relationship.

Limited control: male number

expectation is that the more challengers, the more subordinates sires infants. But
there’s no association.

[questions were asked without hands raised]

You want to prevent subordinates from leaving by giving him more siring when
there’s less male subordinates.

Limited control: energetic constraints

Proportion of activity budget
argument reminded: guarding constraints disrupts the male activity budget and
energy and increases the amount they move. Net reduction of energetic intake.
Seasonality in cost-
proportion of activity budget
some difference across seasons- not excited about this difference. But there is some
sense in which males are subject to greater cost when guarding in winter.

Prediction 2: Residency
giving subordinates incentives to stay with the group.
subordinance curve. duration of residency stays longer when the subordinate has
kids.

Prediction3: reduced infanticide

during study: 26 periods of influx (immigration) ~ 1 influx/period of vulnerability
during influx: infants o alpha males
0.36 prob of being without primary protector
0.19 prob of being without surrogate protector
0.07 prob of having no protector

5 infanticides in 26 influxes
2 when primary + surrogate absent
2 when primary defending alpha status + no surrogate
“This is where the hand waving gets very vigorous”
1 when primary + surrogate present, but no mother

Summary:
(i) oestrous overlap accounts for 25% of subordinate conceptions. (try to
discredit classical view)
(ii) male number and competitor pressure do not compromise alpha male
guarding
(iii) alpha male guarding appears immune to variance in potential energetic
costs
(iv) subordinate residency is sensitive to reproductive success. They are
lingering because they already had some
(v) concession ensures infants have some form of protection
Take home message and implications:
male strategy aimed at maximizing number of reproductively viable offspring not
fertilizations
rather than just fertilizing you need to get those kids reproducing. Males are hoist by
their own petard. Inter-sexual conflict. Estrus advertisement.
lots of prevailing theory relies on this old-school view of sexual selection. Given
what we know that baboons are up to, we need to think harder about what these
models and theories are thinking.

Eugene Marais (1871-1936)

all these talks about infanticide- “It’s a wise baboon that knows its own father”. WE
almost imagine infants as passive vessels. Ultimately it’s the kid that’s the strongest
target in the infant. We should turn our attention to what makes these kids survive?