The main focus of Sterck et al (1997) proposal was to evaluate the original ecological model,
designed by Wrangham (1980) for its competence and suggest areas in which this model needed
improving, in order to increase its effectiveness for explaining the different reasons why females
social relationships differ. Sterck et al (1997) define different categories of female social structures
as Dispersal Egalitarian which has no female philopatry, and Resident Egalitarian, Resident Nepotistic
and Resident Nepotistic Tolerant all of which have female philopatry.

The result was the Socio-Ecological model, which includes the suggestion that predation is the
ultimate cause for social relationships in females, a theory first put forward by Van Schaik (1989).
Isbell (1991) expanded on Wrangham͛s (1980) theory implying that resource distribution and
abundance was the main reason for females forming social relationships, not entirely food resources
alone. This suggestion is supported by Digby (2000), suggesting that females compete for a variety of
resources including helpers or nest sites and this can also impact on their social structure. Sterck et
al (1997) still acknowledge feeding competition as a valid second causal effect; regarding this they
suggested 4 modes of feeding competition, Within Group Scramble (WGS) and Within Group Contest
(WGC); Between Group Scramble (BGS) and Between Group Contest (BGC). Dispersal egalitarian
females have low WGC and BGC; Resident egalitarian females has low WGC but high BGC; Resident
nepotistic females has high WGC and low BGC and Resident nepotistic tolerant females have high
instances of both WGC and BGC. These modes are relevant as the dissemination of food sources
occurs both between and within groups. WGC and BGC (BGC is still a vague category and not much is
known but there are hypotheses) regard estimations that these categories occur when food
resources are abundant thus competition between females is likely when food abundance is of
substantial size and is clumped in an immediate area in relation to the size and location of the group.
Within WGC, formal submission of females is predicted. Sterck et al͛s (1997) model suggests that
submission such as bared teeth display is observed across different species all with the WGC
category but (Koenig 2001) argues that no current datum supports the theory that formal
submission of females is exclusively linked to WGC. WGS and BGS are linked to predictions of more
limited resources. Stevenson and Castellanos (2000) observed that no primate populations fitted
into the egalitarian or tolerant structure proposed by Sterck et al (1997). Koenig (2001) argues that
the correlation between female social relationships and feeding competition are only supported in
certain areas and that within a BGC population no evidence was found for the egalitarian or tolerant
social structures; Koenig (2000) suggests that BGC is possibly not linked to distribution of food but to
the abundance of it. . This author suggested that this is probably because very little studies carried
out use measures of reproductive success or overall energy gain to gather information on feeding
competition and therefore they are lack validity. Koenig (2001) also states that when aggression
rates were tested across numerous primate populations the rates always fell short of estimations for
the competition categories.

However Koenig (2002) suggests that females are affected more by food competition and group
living females will probably experience a mixture of scramble and contest both within and between
groups over their lifetime, and agrees that social group formation is linked to predation risks.
Stanford (2002) concurs that predation risk massively influences social relationships in primates.
However, Boinski et al (2002) disagrees that predation risk affects female social relationships. Many
authors have noted that predation risk has not been effectively measured independently. Hill and
Dunbar (1998) claim that using predation rates to estimate predation risk is not valid as primates
have various anti predator defence strategies in operation to reduce predation and using predation
rate data does not take this into account. In order to effectively analyse the overall impact of
predation risk on female social relationships, Numm and Van Schaik (2001) asserts that predator
grouping could be measured to predict the risk of predation on groups of primate and use this
information in conjunction with competition data on the observed species to categorically confirm if
this is an intrinsic factor.

Sterck et al (1997) advocated that infanticide and demography should be considered in the socio-
ecological model as these have major impacts on the formation of female social relationships. Sterck
et al (1997) states that infanticide effects female reproductive success which in turn affects their
social structures. Infanticide relates to mating relationships and Watts et al (2000) states that male
sexual strategies could have an impact on their social relationships, as although they are carry out
infanticide they also may intervene in disputes between females. Koenig (2001) agrees that male
sexual strategies have an effect on food competition between females. Boinski (1999) also
acknowledges that female social relationships are developed in concurrence with male as male
behaviour affects within group feeding competition. Crockett and Janson (2000) indicate that
infanticide is a likely causal factor in small group size, however Wright (1999) states that small group
size is due to severe ecological conditions. Most opinions on infanticide do not note infanticide as
having much importance in the explanation of social system formation according to Dagg (1999) as
infanticide rates are rather low to qualify as a major aspect. However, van Schaik (2001) conveys
that the rates could possibly be low due to the countermeasures already in operation. Van Schaik
(2000) implies that in order to effectively measure infanticide risk a ratio of lactation and gestation
rate would be more accurate and this would also provide further information on longevity rates
Janson (2003).

There are further alternative explanations for female social relationships; Koenig (2001) puts
forward that there may perhaps be have a phylogenic explanation to social system disparities, this is
supported by Kappeler and van Schaik (2001) who agree that life histories are a key component in
understanding primate social systems. Nunn et al (2000) suggests that disease occurrences in
primate populations could effect behaviour and therefore affect social relationships. A further
suggestion is that societies are structured differently due to the self structuring phenomena as put
forward by Hemelrijk (1999), stating that when given behavioural rules are in operation, social
structures would emerge. Matsumura and Kobayashi (1998) suggest that dominance relationships
are not built on ecological terms alone but are also requisites of evolutionary stable strategies.
Koenig (2000) draws attention to the fact that the Sterck et al (1997) model does not cover all
populations as habitat destruction has caused unstable situations and therefore makes it difficultto
make predictions. Finally Barrett and Henzi (2001) state that the availability of partners could have
an impact on female social relationships but this is not acknowledged by the socio-ecological model,
they also suggest that body size and seasonality could have an effect on forming coalitions within
female relationships.

Although the socio-ecological model still is relevant today, it emits certain areas of consideration
that could affect female social systems.




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