Male Services and Mating Success: Sexual Selection As Instigator of Male Cooperation

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Male services and mating success: Sexual selection as instigator of male

cooperation
Thesis · April 2017

DOI: 10.13140/RG.2.2.22170.49605
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Master Thesis
Male services and mating success
Sexual selection as instigator of male

cooperation
Simona Mandra
April 2017
Supervision by
Dr. Erik P. Willems
University of Zurich
Department of Anthropology
Male services and mating success Simona Mandra
Sexual selection as instigator of male cooperation
Abstract
Explaining cooperative behaviours in animals remains a key challenge in evolutionary biology. In many
species, males perform cooperative acts that are unreciprocated in kind by female beneficiaries. This opens
up the possibility that, whenever females can exert some degree of mate choice, sexual selection may be the
driving force behind such one-sided male cooperative behaviours (i.e. male services). The aim of this study
was to investigate this hypothesis and establish whether a relationship exists between the extent of male
helping behaviour and mating success. Behavioural data were collected on three wild groups of vervet mon-
keys (Chlorocebus aethiops pygerythrus) from two study populations in South Africa. Two different dyadic
association indices were correlated with dyadic mating proportions for a total of 8 males and 16 females.
The results do not support the hypothesis that males who behave more cooperative towards females profit
from increased mating success. Both males (τ = 0.558, 95%, CIBCa = 0.045 - 0.794, ndyads = 18) and females
(τ = 0.524, 95%, CIBCa = 0.137 - 0.719, ndyads = 23) may prefer higher ranking mating partners under certain
circumstances. High ranking females are found more often in spatial association with males which is con-
sistent with the assumption that male proximity is valuable to females. These findings suggest that male
services do not always translate directly into increased mating access and that the beneficial effects of indi-
vidual dominance are moderated by additional factors.
Introduction
MALE SERVICES
In primates, certain cooperative behaviours are frequently performed primarily by males. These one-sided
acts of assistance (henceforth male services) are provided to females or juveniles in the absence of recipro-
cation with the same kind of cooperative behaviour. Male services can be found in a wide range of taxa and
along a spectrum of behaviours, including predator defence (Stanford, 2002; van Schaik and van Noordwijk,
1989), agonistic support (Palombit et al., 1997), conflict mediation (Flack et al., 2005), food sharing (Goffe
and Fischer, 2016; van Noordwijk and van Schaik, 2009) and grooming (Norscia et al., 2009).
According to Darwin (1859), a trait can only be favoured by natural selection if it leads to an increase in
fitness for its bearer. If an individual provides assistance to others, the resulting fitness benefits to the service
provider and their underlying mechanisms are not immediately apparent. The recipients of these acts of
assistance appear to be able to exploit the donor’s help and thus it is a puzzle how cooperative acts are
evolutionarily stable (Nowak, 2006; West et al., 2007).
Two main evolutionary pathways have been proposed to explain cooperative behaviour: Indirect fitness
benefits can be secured when an actor enhances the fitness of another individual sharing copies of its own
genes, most plausibly through kin selection in which an actor promotes a relative’s reproduction (Hamilton,
1964; Maynard Smith, 1964; West-Eberhard, 1975). The second pathway is constituted by direct fitness
benefits, accrued by an actor through reciprocation of a cooperative act (Trivers, 1971). Over the past couple
of decades, Trivers’ original concept of reciprocity has been extended to encompass a myriad of non-mutu-
ally exclusive proximate mechanisms that tackle the question of how cooperation may lead to direct fitness
benefits (Bshary and Bergmüller, 2008). One possible direct fitness benefit a male may receive for providing
services is an enhanced reproduction through increased mating access, which can come about through sev-
eral mechanisms:
(i) An unequivocal proximate mechanism explaining male service provision is based on by-product bene-
fits resulting from a male’s self-serving behaviour (Brown, 1983; West-Eberhard, 1975). If a male per-
forms a behaviour to exclusively maximise his own benefits, e.g. by defending a home range to mo-
nopolize mating access to females, a beneficial by-product for females might be resource defence. This
principle has been introduced by Rubenstein (1986) and proposed to be relevant e.g. in colobine mon-
keys (Fashing, 2001).
(ii) Another way in which cooperative intent might enhance mating access is based on direct reciprocity
(Trivers, 1971). In group-living primates with individualised and long-term bonds, such reciprocity has
2
been found to be rather relaxed and based on emotional bookkeeping, and is often referred to as friend-
ship (Nguyen et al., 2009; Schino and Aureli, 2009; Silk, 2002b). This concept is contrasting with first
advances in explaining cooperation between non-relatives as it does not presume a strict tit-for-tat like
symmetry (Axelrod and Hamilton, 1981) which has often been criticised for being too cognitively de-
manding in large groups (Hammerstein, 2003; Stevens and Hauser, 2004).
Mating access could constitute a female act of reciprocation in exchange for male services such as
conflict support, grooming or social tolerance. In yellow baboons, long-term affiliative bonds between
males and females have indeed been shown to have a sexual selection component that includes mating
access and a high paternity probability for cooperative males (Nguyen et al., 2009), whereas in long-
tailed macaques (Gumert, 2007) and rhesus macaques (Massen et al., 2012), a similar association be-
tween male-female grooming and mating access has been found.
(iii) When mating partners don’t equally invest in offspring, the sex that provides more infant care is ex-
pected to be choosier with regard to mating partners and the less invested sex is expected to compete
for mating access (Trivers, 1972). In species where females invest more in offspring, they are assumed
to choose mating partners based on previous experience and available information (Jennions and Petrie,
1997). Cooperative intent could serve as a phenotypic signal, analogous to visual ornaments that pro-
motes mating access (Raihani and Smith, 2015; Roberts, 1998; Zahavi, 1975). If male cooperation is
deployed as a signal of quality, a female that responds with granting a higher mating access is perform-
ing a self-serving act rather than an act of cooperation as it is in the female’s direct interest to mate with
the most suitable partner, e.g. to acquire valuable genetic material for her offspring. In this case, the
exchange of male services for mating access is an example of pseudo-reciprocity (Bergmüller et al.,
2007).
Male services as signals of quality may also indirectly lead to direct fitness benefits: Performing a
cooperative act may enhances a male’s reputation and thus his perceived value as a mating partner with
females who witnessed the service provision without directly benefitting from it. Thus, signalling may
be part of indirect pseudo-reciprocity in which bystanders of a cooperative act respond by benefitting
the actor with a self-serving behaviour (Alexander, 1987; Bshary and Bergmüller, 2008; Lotem et al.,
2003; Nowak and Sigmund, 1998). Experimental studies with humans (Griskevicius et al., 2007; Iredale
et al., 2008; Rusch et al., 2015; Tognetti et al., 2012) and research on hunter-gatherer (Apicella, 2014)
as well as agro-pastoralist societies (Glowacki and Wrangham, 2015), suggest cooperative intent to be
a trait of interest for male and female mate choice based on an enhanced reputation.
AIM OF THIS STUDY
A recent study by Arseneau et al. (2015) found a positive association between male support of females in
between-group conflicts and subsequent mating success in vervet monkeys (Chlorocebus aethiops
pygerythrus), suggesting that male services in a high risk context may affect female mate choice in this
species. The aim of the present study is to build on this previous finding by investigating whether male
vervet monkeys who behave cooperatively towards females in low-risk contexts gain enhanced mating ac-
cess as well. In order to answer this question, this study focusses on the effects of low-cost socio-positive
behaviours in the within-group context, more specifically male social tolerance, both throughout the entire
diurnal activity period and during the feeding context only.
One way in which a female might profit from being tolerated in a male’s proximity is enhanced safety from
predators. There is evidence that male vervet monkeys are more vigilant than females (Baldellou and Henzi,
1992), a finding that was corroborated in the current study (Supplementary figure 1). Moreover, when fe-
males are in proximity, captive male green monkeys (Chlorocebus aethiops sabaeus), a species closely
related to vervet monkeys, have been found to give more alarm calls compared to situations in which only
other males are present (Cheney and Seyfarth, 1985). Therefore, reduced predation risk is a likely benefit
for females who spend time in a male’s proximity. Male vigilance may also be directed at extra-group males,
and might thus additionally provide protection against potentially infanticidal intruders (Isbell et al., 2002).
3
Such advantages might be particularly valuable to females in the feeding context, as extraction of food items
requires attention which is likely to be an obstacle to predator detection (Wickler, 1985) or because valuable
food sites could be located at places with an increased predation risk (Cowlishaw, 1997; Josephs et al.,
2016). Apart from a more efficient food intake (Haunhorst et al., 2017), male tolerance in feeding situations
may lead to females profiting from quality food sources (Dubuc et al., 2012). Hence, male proximity might
allow females to be less vigilant compared to situations in which they are alone or with other females,
thereby increasing foraging efficiency and also enabling the exploitation of higher risk areas or high value
resources.
In the present study, the effects of male services on mating success are analysed in two wild populations of
vervet monkeys in South Africa. To estimate the amount of male tolerance and occurrence of co-feeding
with females, association indices are constructed that quantify male proximity to females throughout the
entire activity period, as well as in the feeding context in particular. To measure copulation success, dyadic
mating indices are calculated and then correlated with the association indices in order to evaluate the effect
of male services on mating access.
Studies analysing social bonds in nonhuman primates often rely on data collected in group scan sampling
(Borgeaud et al., 2016; Young et al., 2017). While this method allows for the collection of large quantities
of data across many individuals, important drawbacks are the susceptibility to bias towards more conspicu-
ous behaviours and individuals as well as non-independence between data points collected on different in-
dividuals in the same group scan (Altmann, 1974; Martin and Bateson, 1993). In this study, it was attempted
to reduce such drawbacks as much as possible. To achieve this, systematically collected focal animal data
in which each group member was represented equally as well as evenly across the photoperiod, were used
to calculate association indices.
Another commonly used method in the study of primate associations are composite sociality indices that
sum up the number of observations of different affiliative behaviours such as grooming and proximity be-
tween two individuals and put them into relation with the individuals’ observation time (Borgeaud et al.,
2016; Cords and Nikitopoulos, 2015; Haunhorst et al., 2016; Ostner et al., 2013; Silk et al., 2003). These
indices implicitly assume that all the included behaviours are equally strong indicators of bondedness be-
tween individuals. No compelling evidence for such equivalidity of different affiliative behaviours has yet
been described for any primate species and some studies even find opposite effects in the analysis of differ-
ent affiliative behaviours (Young et al., 2017). Therefore, the association indices used in this study are based
on distinct categories of affiliative behaviours.
HYPOTHESES AND EXPECTATIONS
In line with the above mentioned findings on the high-risk male service of participation in between-group
conflicts (Arseneau et al., 2015), it is hypothesized that a male’s mating access will increase with his provi-
sioning of low-risk services in the present study. Thus, a positive correlation of male mating access with
tolerance and co-feeding is expected.
In order to take into account confounding effects, the influence of male and female rank on mating success
and proximity are included in the analysis. High ranking females have been found to be preferred mating
partners in green monkeys and other nonhuman primates (Keddy, 1986; Samuels, 1980; Seyfarth, 1978;
Wolfe, 1979) and various studies report socially dominant male primates to be more successful at mating
(Andelman, 1987; Freeman et al., 2016; Keddy, 1986; Massen et al., 2012). High ranking individuals are
thus expected to be preferred mating partners or more successful competitors which is hypothesized to
translate into a positive correlation between individual rank and mating access.
Under the assumptions that male services lead to an enhanced mating success, and that high ranking females
are favoured mating partners, males are expected to provide more services to preferred females leading to a
positive correlation between female rank and service reception.
As a response to the expected disadvantageous effect of low rank on male mating success, service provision
is hypothesized to be deployed by low-ranking males as a means to compensate for this shortcoming and as
4
an expression of an alternative mating strategy (Soltis et al., 1997; Wroblewski et al., 2009). Therefore it is
hypothesized that the amount of tolerance a male provides to females in general and in feeding contexts is
negatively correlated to his rank.
An alternative hypothesis regarding male proximity to females is that males seek female’s spatial association
in order to gain mating access. Should this be the case, a positive correlation between a male’s spatial asso-
ciation to a female and the allocation of copulations to that female is expected. In order to test this alternative
hypothesis, the correlation between male matings and association with females is analysed.
Material and Methods

FIELD SITES
Data were collected on 3 groups of free ranging vervet monkeys at two field sites, both located in the eastern
part of South Africa: Lajuma research centre (29°26'05"E, 23°02'23"S) which is part of a conservancy situ-
ated in the Soutpansberg Mountain Range, and Mawana research site (31°12'35''E, 28°00'33''S), which is a
distinct area on a privately owned game reserve in KwaZulu-Natal.
BEHAVIOURAL DATA COLLECTION
At both field sites, Palm® handheld computers with the Pendragon Forms® software were used to collect
behavioural data.
At Lajuma, data were collected on one study group which on average consisted of 17.8 individuals and
contained 1-5 adult males and 5-8 adult females. Adult individuals were followed throughout their entire
diurnal activity period, and focal data were systematically collected during ten minute sampling periods
using instantaneous point sampling (Altmann, 1974; Martin and Bateson, 1993) at one-minute intervals.
The day was divided into 4 time zones and each adult individual was sampled three times per time zone per
month. To maximise independence between focal sessions, the same individual was never sampled in con-
secutive time zones on the same day. When a focal session yielded less than seven successful data point
recordings, all data were discarded and a new session was recorded at a later time. Apart from general
information about the ecological setting and activity of the focal individual, the identity of and distance to
the focal individual’s nearest neighbour was recorded when located within a distance of up to 5m. A total
number of 14’159 sample points were collected over a period of 17 months from January 2004 to April
2006 (Willems, 2007).
The focal protocol used at Mawana is a modified version of the protocol used at Lajuma and therefore
identical in most regards. In order to collect detailed social data, the protocol was altered to maximize the
information about a focal individual’s neighbours within a distance of up to 10m. Each neighbouring indi-
vidual in up to 5m, up to 2m or within an arm’s length was recorded. When there was no neighbour within
a distance of 5m or less, the nearest neighbour was also recorded if located at a distance of up to 10m. To
allow for this more detailed collection of information, the sample interval was increased to 2 minutes. 10
data points were collected during focal sessions of 20 minutes. The photoperiod was divided into three
equally long parts and it was ensured that the number of collected data points was unbiased across individ-
uals and these three diurnal time zones. Between two focal sessions for the same individual, a minimum of
two hours was maintained and only sessions where the focal individual was in sight in at least six sample
points were considered to be valid. Data were collected between April and September 2016. A number of
7’548 sample points were recorded for 15 individuals in two groups (NH: 6 females and 2 males, AK: 4
females and 3 males). Throughout the observation period, the groups consisted of 29-33 (NH) and 20-22
(AK) individuals in total.
Sexual behaviours in both groups were recorded ad libitum every time they were observed during group
follows (Altmann, 1974; Martin and Bateson, 1993). A total of 55 successful matings was recorded at La-
juma and 104 at Mawana. As it is difficult to observe and record all matings in the field, the data set is likely
to be incomplete which gives rise to a possible bias. It should be noted, though, that during a data collection
5
procedure in which the observer’s main activity is focal animal sampling evenly distributed across individ-
uals, the observation of sexual behaviours can be expected to be less biased towards conspicuous individuals
as well.
DATA PREPARATION
Dyadic data
In order to estimate dyadic social tolerance, co-feeding and mating success, association and mating indices
were calculated based on the collected focal animal and ad libitum data. For each dyad, the calculation was
based on the sample points in which two individuals co-resided in the group, i.e. on the periods in which
there were focal data available for both individuals. All calculated indices were stored in intersexual dyadic
matrices.
As a proxy for tolerance, general association indices (AI overall) between two individuals were calculated
employing a method adjusted from Whitehead (2008): The sum of all focal sample points for which two
opposite-sex adult individuals were located within 5m of each other was divided by the total number of
focal sample points collected for these two individuals:
( )+ ( ) (1)
=
+
Af(Bn) is the amount of sample points in which A was the focal individual while individual B was its neigh-
bour and, accordingly, Bf(An) is the sum of sample points in which B was the focal individual and A was
located within 5m proximity. Af and Bf are the total number of focal sample points collected for individuals
A and B. In the calculation of an association index in feeding contexts (AIfeeding), the same logic as in the
calculation of general association indices applied to determine a proxy for co-feeding. The number of sample
points in which one member of a dyad was feeding in up to 5m vicinity of the other individual was divided
by the total number of sample points in which each of the two focal individuals were feeding.
To evaluate copulation success, for each adult individual’s total number of matings the proportion in which
each opposite-sex individual was the partner was determined. Thus, mating indices were calculated regard-
ing the proportion of matings each male (MImale) and female (MIfemale) obtained of each opposite-sex partner:
= (2)
MatAB is the total number of observed matings between the individuals A and B while MatA is the total
number of data points in which individual A was involved in a successful copulation.
Rank
Each individual’s intra-sexual dominance rank was determined based on approach-retreat interactions, col-
lected ad libitum. The highest ranking male in each group was regarded as dominant while all other male
group members were considered non-dominant. In the Lajuma population, the females that were in the upper
half of the dominance hierarchy were considered high-rankers while individuals in the lower half of the
hierarchy were considered low-rankers. As Mawana is a long-term study site, more detailed demographic
data are available and in this study population females could therefore be categorized into high, middle and
low dominance classes depending on their matriline.
STATISTICAL ANALYSES
The fact that each individual was part of several dyads when calculating the dyadic measures of AI and MI
lead to an inherent non-independence among data-points. This was taken into account by using random
resampling with replacement: Nonparametric correlation tests were used to determine the strength of the
relationship between dyadic association matrices on the one hand, and the dyadic mating proportion matrix
6
on the other. Significance of Kendall‘s correlation coefficients was estimated by employing the bias-cor-
rected and accelerated bootstrap (Efron, 1987), based on 9000 resampling events with replacement to cal-
culate 95% confidence intervals around the parameter estimate.
To investigate the role of individual traits, mating indices were correlated with individual rank to reveal
potential preferences of males and females to mate with high ranking partners in the manner described
above. Furthermore, the statistical relationship between the association indices and individual rank was de-
termined to analyse dominance-related patterns in individual space use.
In the calculation of the nonparametric tests, only dyads for which there was a minimum of 300 data points
available were included in the analysis. One female from the Lajuma population (AF 8) exhibited a lower
level of habituation than the rest of the group and – suspectedly as a result of that – the amount of ad libitum
data collected on her was low. Dyadic data including AF 8 were thus excluded from the analysis to prevent
a biased result. Consequently, the analysis for Lajuma was conducted for a total of 18 dyads that were
composed of 3 males and 6 females. For the Mawana population, the data of both groups were internally
standardized to z-scores and then lumped together to allow inclusion in the same analyses. 5 males and 10
females were part of the analysis and there were sufficient data for 23 out of 24 possible dyads.
All statistical analyses were run in R version 3.3.1 using the “boot” package version 1.3-18 (Canty, 2002)
to perform bootstraps.
Results
DETERMINANTS OF MALE MATING SUCCESS
The proportion of a female’s matings that a male obtained at Lajuma was neither related to his dominance
rank nor to the proportion of time he spent within 5m of her, regardless of whether proximity was assessed
exclusively in the feeding context or overall (Table 1). At Mawana, the proportion of time a male was
located within 5m of a female did not show a significant effect on mating access in overall situations as well
as feeding contexts. Furthermore, male rank was significantly positively correlated to the proportion of
female matings a male obtained in the Mawana population.
Table 1 Kendall correlation coefficients and bootstrap confidence intervals of the correlation
between male proportion of female matings (MImale) and association indices as well as the
correlation between MImale and male rank.
Study population Parameter Effect size (τ) 95% CIBCa
Lajuma AIoverall 0.187 -0.289 0.592
AIfeeding 0.072 -0.382 0.482
Male rank 0.126 -0.230 0.496
Mawana AIoverall -0.016 -0.339 0.329
AIfeeding -0.072 -0.365 0.262
Male rank 0.524 0.137 0.719
DETERMINANTS OF FEMALE MATING SUCCESS

A female’s rank was positively associated with the proportion of male matings she obtained at Lajuma
(Table 2). This effect was not found in the Mawana population where female rank did not correspond to the
proportion of a male’s matings that a female received. In both populations males did not appear to assign a
higher share of their copulations to females with whom they were more often spatially associated in overall
contexts as well as in feeding situations.
7
Table 2 Kendall correlation coefficients and bootstrap confidence intervals of the statistical
relationship between a female’s obtained proportion of male matings (MIfemale ) and proxim-
ity as well as the correlation of MIfemale with female rank.
Study population Parameter Effect size (τ) 95% CIBCa

Lajuma AIoverall 0.235 -0.215 0.611
AIfeeding 0.229 -0.275 0.650
Female rank 0.558 0.045 0.794
Mawana AIoverall 0.087 -0.235 0.438
AIfeeding 0.119 -0.208 0.464
Female rank 0.206 -0.143 0.504
DETERMINANTS OF PROXIMITY
At both field sites female rank and male spatial proximity overall and specifically in the feeding context
appeared to be associated: The correlational analyses yielded effect sizes of τ = .422 and τ = .487 at Lajuma
as well as τ = .543 and τ = .334 at Mawana. However, only the association of AIoverall at Mawana reached
statistical significance at the 95% level (Table 3). Male rank and female spatial proximity, in contrast, did
not exhibit any consistent pattern of association.
Table 3 Kendall correlation coefficients and bootstrap confidence intervals of the statistical relationship between
general spatial proximity and male as well as female rank.
Study population Parameter 1 Parameter 2 Effect size (τ) 95% CI BCa

Lajuma AIoverall Female rank * 0.422 -0.066 0.672
AIoverall Male rank -0.095 -0.465 0.318
AIfeeding Female rank * 0.487 -0.042 0.735
AIfeeding Male rank -0.076 -0.465 0.325
Mawana AIoverall Female rank 0.543 0.149 0.738
AIoverall Male rank 0.132 -0.285 0.483
AIfeeding Female rank 0.334 -0.085 0.620
AIfeeding Male rank 0.044 -0.360 0.413
* p < 0.1
Discussion
This study set out to investigate the effects of low-cost male services on mating success. It found no support
for the hypothesis that the provision of male tolerance in overall situations and feeding contexts was asso-
ciated with a higher mating success in both analysed study populations.
Dominant males did not gain a higher share of female matings at Lajuma but were more successfully ac-
quiring copulations at Mawana. Female rank on the other hand was positively correlated to mating success
in the Lajuma population while it was not statistically related to the allocation of male copulations at
Mawana. These mixed results indicate that there might be interesting site-specific factors moderating the
effect of an individual’s dominance on the proportion of acquired matings in the two study populations.
Further correlational analyses of the calculated association indices revealed that males did not mate more
with females they were found in proximity with more often in general or while feeding. Moreover, dominant
females were to some extent more often spatially associated with males while male rank did not show any
effect on the propensity to be located in a female’s vicinity.
These results may have various implications.
8
LOW-COST MALE SERVICES AND MATING SUCCESS

The employed measures of tolerance and co-feeding were not significantly correlated to male mating suc-
cess. This finding is not consistent with the hypothesis that male provision of low-cost services is driven by
the prospect of gaining an enhanced mating access. Possible explanations are that females don’t value male
presence or that male tolerance provision is leading to other benefits than mating access. Another explana-
tion would be that service provision was equally distributed across female group members which would
lead to enhanced difficulties to detect a resulting higher mating access. Visual inspection of standardized
association indices regarding general proximity in 5m reveals a considerable variation in male proximity to
females: Most males spent different amounts of time with different females and the amount of time spent in
the vicinity of females was variable between males as well (Supplementary figure 2). Hence, an equal dis-
tribution of male services across females does not appear to apply as an explanation for the absence of a
correlation with mating success while the value of male presence to females remains unclear.
THE EFFECT OF DOMINANCE ON MATING SUCCESS
The findings regarding the effect of dominance on mating success are only partly in line with the hypothesis
that high ranking individuals are generally preferred mating partners. Rank does not seem to have an equally
positive effect on mating access in every situation which might be due to external or demographic factors
(Berard, 1999; Cowlishaw and Dunbar, 1991). One salient difference between the two study populations
analysed here is predation rate. While predation events were common at Lajuma, they occurred extremely
rarely at Mawana. The difference in predation may have influenced the study subjects’ behaviour or even
the incentive for males to act cooperatively. Moreover, predation risk could augment the costs of mating:
The trade-off between mating and vigilance could lead to increased risk of mating at high-predation sites.
Furthermore, an increased predation pressure could generally result in lower energy availability, e.g. as a
consequence of elevated glucocorticoid levels (Emery Thompson and Georgiev, 2014). While males are
often expected to maximize their access to females (Andersson, 1994; Bateman, 1948; Trivers, 1972), en-
ergetic costliness of male mating effort could be considerable (Bercovitch and Nürnberg, 1996; Emery
Thompson and Georgiev, 2014) and as a consequence impose restrictions on the total amount of male mat-
ings in a given time. Furthermore, there is evidence that dominant females gain fitness benefits, manifesting
themselves e.g. in a higher infant survival in vervet monkeys (Whitten, 1983) and other primates (Pusey et
al., 1997; Silk, 2007; van Noordwijk and van Schaik, 1999). It is thinkable that the advantages of being
dominant are more pronounced in high predation areas. Therefore, there could be an incentive for males to
mate predominantly with preferred females when the costs of mating are elevated (Alberts et al., 2006) or
when dominant females can reap a high amount of rank-related fitness benefits.
An alternative explanation for the finding that high ranking females at Lajuma gained a higher share of male
matings is female competition. While originally thought to be relevant only in species with “reversed sex
roles” (Emlen et al., 1998; Jones et al., 2000), this dimension of sexual selection has recently gained atten-
tion also in other systems including primates (Doran-Sheehy et al., 2009; Kvarnemo and Simmons, 2013;
Palombit et al., 2001; Rosvall, 2011). One of the two Mawana study groups analysed here (AK) is known
for its extraordinary tolerance among females (Borgeaud and Bshary, 2015) that corresponds to a high re-
latedness between females in this group (Schnider, 2016). This might suggest less pronounced female com-
petition for male matings at Mawana and could be responsible for the absence of a female rank effect on
male copulation allocation. On top of that, flat female hierarchies might also entail less distinct male pref-
erences for high ranking females.
SPATIAL ASSOCIATION AND MALE MATING ALLOCATION
If males seek proximity to preferred female mating partners, one would expect a correlation between spatial
association and male copulation allocation. As proximity to a female was not correlated with the proportion
of matings allocated to that female, there is no indication for proximity reflecting male mating preferences.
9
SPATIAL ASSOCIATION AND SOCIAL DOMINANCE

The finding that in both populations female rank was to some extent positively associated with male prox-
imity could be the result of high ranking females gaining priority of access to the valuable resource of male
tolerance. On the other hand, males might seek the vicinity of high ranking females in order to form alliances
with these potentially preferred association partners (Young et al., 2017). As spatial association was not
correlated with mating access, this result – even though in line with the initial expectations – is not likely to
be the consequence of an increased male mating effort with high ranking females.
EMPLOYED ESTIMATES AND DATA QUALITY
In order to estimate a dyad’s co-feeding rate, the data points in which the focal individual was feeding while
a neighbour was located in a distance of 5m or less were used. As the deployed focal protocols do not
include any information about the neighbour’s activity, it is expected that in some cases only one of the two
members of a dyad was feeding. However, it was commonly observed in the field that all individuals in
sight were feeding when conducting focal animal sampling. A co-feeding estimate based on scan data col-
lected at Lajuma supports this impression: An average of 56.6% of the sampled individuals were feeding in
the scan samples that included at least one feeding individual. For that reason, the number of focal sample
points in which the neighbour was non-feeding while the focal individual was feeding is expected to be low
and without a meaningful influence on AIfeeding.
When cooperative acts are exchanged reciprocally, it is unclear within which time frame a recipient will
repay a donor. Here, the relation between long-term association of a dyad and their mating behaviour has
been analysed. This is in line with previous research on primates that found an association of long-term but
not short-term grooming patterns with mating access (Massen et al., 2012). However, there is a possibility
that socio-positive behaviour only has an influence on mating success when it occurs shortly before mating
(Gumert, 2007) or that mating could be followed rather than preceded by cooperative or socio-positive
behaviour.
Furthermore, male services such as predator defence, allogrooming and support in within-group conflicts
were not abundant enough in the datasets used here to allow for a meaningful analysis. Employing data
recording methods that are more systematic than scan or ad libitum sampling but more specifically aimed
at recording rare behaviours than focal animal sampling, e.g. focal behaviour sampling (Martin and Bateson,
1993), could provide an approach to investigate the effects of relatively rare cooperative behaviours on
mating success in vervet monkeys. Two thesis works that used different methods to analyse the effect of
grooming on copulation frequency in vervet monkeys did not find a relation between these two variables
(Freeman, 2012; Hellard, 2007).
The relatively small data set did not allow for a detailed statistical analysis with more than one co-variate.
Effects of confounding variables such as seasonality, type of food patch, physical condition, maternity/pa-
ternity status and female fertile phase had to be neglected due to lack of observability or statistical power.
Another important confounding variable is demography: Male mating success is likely to depend on the
presence or absence of other males and potentially on the number of females relative to the number of males
(Alberts, 2012; Cowlishaw and Dunbar, 1991). In the Mawana population, group composition did not un-
dergo major changes during the observation period. Demography effects can thus be expected to have re-
mained stable over this time. For the Lajuma population, each individual’s share of matings was multiplied
with the weighted average of total number of same-sex group members in an attempt to correct for “diffi-
culty” to obtain a high share of matings (Supplementary table 1). As this correction factor did not markedly
change the order of received proportions of female copulations, a meaningful influence on the nonparamet-
ric test results appears unlikely and hence the initially calculated values were used in the statistical analysis.
DIFFERENCES BETWEEN THE TWO FIELD SITES
The partly dissimilar findings in the two study populations could be caused by variations in demographic
and habitat-related factors. In addition to the differences mentioned above, variable climate and food abun-
dance might lead to different preconditions in the two study populations. At Mawana, the data were collected
10
during a period of extraordinary drought which may constitute such a difference. Furthermore, occasional
provisioning of the Mawana population in connection with experimental research (Borgeaud and Bshary,
2015; van de Waal et al., 2013) might have influenced the results.
Apart from using slightly different focal protocols, data for the two study populations were not collected
across equally long time spans. While the data from Lajuma cover more than a year, the sample points at
Mawana were collected only over a period of six months. The difference between the two field sites could
e.g. be a result of the fact that the data for the Mawana population were collected mainly during the mating
season while the data for the Lajuma population cover the whole year. Furthermore the adult group members
remained the same at Mawana during the time of data collection. The observation period at Lajuma on the
other hand coincided with male immigration events that were connected to power take-over and eviction of
the previous dominant male. Hierarchical instability among males could thus have been a stronger influenc-
ing factor in the Lajuma population.
PROXIMITY AS A PROXY FOR MALE SERVICES
Demonstrating the cooperative nature of a behaviour is not trivial. Apart from the difficulties in evaluating
the costs and benefits for the involved individuals, demonstrating selection pressure on the beneficial effect
on the recipient is a challenge (Clutton-Brock, 2009). To establish male tolerance as a service, one would
need to provide further evidence that females are gaining a fitness benefit from it and it would be necessary
to show an evolutionary incentive for a male to exert this cooperative act.
One important issue to note is that proximity might not be equally relevant in each kind of situation. Here,
feeding situations have been analysed separately to take into account a potentially more relevant condition
(also see Supplementary table 2 and Supplementary figure 3). Apart from an individual’s activity, factors
such as group spread, habitat and food type could also influence the beneficial effect of a male’s proximity
for a female. While proximity may be an expression of tolerance and cooperative intent in many situations,
it is also a side-effect or even the result of agonistic events in other situations. Furthermore, there is a pos-
sibility that intersexual proximity is an expression of female rather than male tolerance. The ability to fully
recognize and record the different facets of proximity would allow for a more accurate investigation regard-
ing the effects of male tolerance on female mate choice.
Kin selection often explains cooperative behaviour (Clutton-Brock, 2002; Silk, 2002a) and paternal care
comprises an important alternative explanation for increased male-female proximity. Nonhuman primates
are thought to have cues to recognize their offspring, but these don’t appear to be equally represented across
taxa: Several studies report that males are more likely to be associated with their own offspring compared
to unrelated infants (Buchan et al., 2003; Langos et al., 2013; Moscovice et al., 2010; Ostner et al., 2013)
but in other species, paternity does not show any promoting effect on male-infant relationships (Ménard et
al., 2001; Rosenbaum et al., 2015). In the case of vervet monkeys, genetic paternity has been shown to
influence a male’s reaction to infant distress calls (Hauser, 1986) and estimated paternity probability partly
explains participation in between-group conflicts (Arseneau et al., 2015). However, experimental studies
with green monkeys find that males behave more affiliative towards infants when the infant’s mother is in
visual contact which suggests that paternity is not the only factor influencing male-infant relations (Keddy-
Hector et al., 1989). Further studies are needed to disentangle the different, non-mutually exclusive factors
that may lead to male bonding with mothers of dependent offspring.
Where male-infant association is not explained by paternal care, it might in fact be a case of a male service
leading to enhanced mating success (Smuts and Gubernick, 1992; van Schaik and Paul, 1996). Evidence for
this “care-then-mate” hypothesis has been found in barbary macaques (Ménard et al., 2001) and crested
macaques (Kerhoas et al., 2016). An attempt to analyse this in the present data set is provided in the sup-
plementary material (Supplementary table 2).
OUTLOOK
While Arseneau et al. (2015) found an association of male participation in territory defence with subsequent
mating success in vervet monkeys, the present study did not find a similar effect for social tolerance and co-
11
feeding which are presumably considerably less costly services. Providing different kinds of male services
might lead to different kinds of rewards. Pre-copulatory mate choice is only one component of sexual se-
lection (Darwin, 1859; Huxley, 1938) and only one of several reproductive stages at which sexual selection
can influence the evolution of a behaviour (Kvarnemo and Simmons, 2013). As a consequence, sexual se-
lection may offer incentives for males to provide services to females also in the absence of a higher mating
access.
Apart from mating access, an important parameter to consider is paternity success which is explained by
mating frequency in some studies (Alberts et al., 2006; Bercovitch and Nürnberg, 1996; de Ruiter and van
Hooff, 1993), but not found to be a reliable predictor in other cases (de Ruiter and van Hooff, 1993; Ménard
et al., 2001). While mating is a crucial precondition to siring offspring, it is not the only relevant factor. In
many organisms, cryptic female choice is a determinant of paternity (Birkhead, 1998; Eberhard, 1996); this
dimension of sexual selection has not been described extensively in primates (Drea, 2005). The female
orgasm is hypothesized to have a reproductive function in encouraging fertilization in primates (Allen and
Lemmon, 1981; Milton, 1985) and could be a responsible factor for the mismatch sometimes found between
copulation frequency and paternity success. Furthermore, phenomena such as selective sperm removal
(Reeder, 2003; Strier and Ziegler, 1997) or selective maternal investment in unborn infants (Pereira, 1983)
have been shown to play a role in some cases and might turn out to be relevant in future studies. In cases
where female choice continues to have an influence after copulation, male service provision could be a
factor favouring the fertilization of females.
Male services may not only translate directly into an increased mating success, but also indirectly, e.g.
through group membership and social integration, tenure length and dominance rank (Figure 1). To gain
regular mating access to the females of a group, an established membership in that group is a likely precon-
dition (Muller and Emery Thompson, 2012). Alliances with females could be crucial to establish group
membership and service provision might be a means to form such alliances. As a consequence, social toler-
ance could be indirectly beneficial to gain mating access by promoting a faster acceptance or prolonged
tenure in a group.
Male services and reproductive success
indirect direct
influence influence
Group tenure Mating
Male services
Dominance Fertilization
Figure 1 The provision of male services could promote male repro-

ductive success directly via mating success or paternity probability
as well as indirectly by enhancing group tenure or the acquisition
of a high rank.
Paternity in vervet monkeys is, at least in some cases, achieved mainly by dominant males (Minkner et al.,
2017; Weingrill et al., 2011). If these findings are substantiated in the future, dominance will have to be
regarded as the primary influencing factor on reproductive success in this species. Achieving a high rank
could hence be a most effective way to increase fitness and again, male service provision could promote
this. Females have been suggested to have leverage over male careers in vervet monkeys (Young et al.,
2017) and green monkeys (Raleigh and McGuire, 1989). Therefore, bonds with females could be an instru-
ment to not just prolong group tenure but also to positively influence a male’s power trajectory.
12
Same as the present study, most research analysing the effect of male cooperative behaviour on sexual
selection has focused exclusively on mate choice in humans (Arnocky et al., 2016; Iredale et al., 2008;
Raihani and Smith, 2015), non-human primates (Clarke et al., 2010; Gumert, 2007; Massen et al., 2012)
and other animals (Christy, 1987; Clutton-Brock et al., 1988; Curio et al., 1983; da Cunha et al., 2017;
Gosling and Petrie, 1990; Zahavi, 1995). A highly interesting additional issue to investigate are the effects
of male services on fertilization probability and indirect factors influencing lifetime reproductive success
such as group tenure and dominance acquisition. The present study highlights the importance of pursuing a
differentiated approach that takes into account the various effects of behaviours that seemingly fall into the
same category.
Cooperation and sexual selection are two concepts with numerous components. Future research will further
elucidate the ways in which these elements are connected.
Acknowledgements
First, I would like to thank my supervisor Erik Willems for the amazing collaboration with countless helpful
inputs and virtually endless support. I am extremely grateful to Carel van Schaik for giving me the great
opportunity to do my master thesis at the Anthropological Institute in Zurich and for helpful discussions and
feedback. I thank Erica van de Waal, Redouan Bshary and Klaus Zuberbühler for enabling me to do field
work at the Mawana research site. Furthermore, I would like to thank Arend van Blerk, Axelle Bono, Miguel
Gareta and all my fellow students and volunteers in the field. Charlotte Bergerat, Severine Caré, Eve Holden
and Maxime Clerc deserve a special thank you for making my time at Mawana a great experience. In addi-
tion, I thank the A.H. Schultz foundation and the University of Zurich for supporting this project financially.
A big thank you goes to all the members of the Anthropological Institute in Zurich for making my work
there very pleasant. I especially thank Franca Eichenberger, Sabine Frei, Benjamin Laubi, Laura Damerius
and Julia Mörchen for their friendship. Finally, I cordially thank my friends and family for all their moral
support and for never losing their patience with me.
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18
Supplementary Material
Sexual selection as instigator of male cooperation Supplementary material
Sex differences in vigilance

A study by Baldellou and Henzi (1992) reports a higher vigilance in male vervet monkeys compared to
females. While males were vigilant in 9.6 – 27.8% of all observations, females only assigned 3.7 – 13.8%
of their time to vigilance in the collected scan sample. The focal data used in the present study reveal a
similar picture for the Lajuma population. While males exhibited a median vigilance of 16.2%, female me-
dian vigilance reached only 6.1% (Supplementary figure 1). The Wilcoxon rank-sum test yields a significant
difference between male and female time spent vigilant at the 5% level (W = 0, p = 0.024, r = -0.753).
Proportion of time vigilant

depending on sex
0.30
0.25
Proportion of time vigilant
0.20
0.15
0.10
0.05
W = 0
p = 0.024
r =-0.753
Female Male
Sex
Supplementary figure 1 Boxplot of the proportions of focal sample points in
which adult females and males were vigilant in the Lajuma study population
including results from the Wilcoxon rank-sum test comparing male and female
proportion of time spent vigilant.
ii
Inter-individual differences in spatial proximity to opposite-sex adults

A comparison of the 5m proximity association indices for each male in both study populations shows that
most males did not distribute their tolerance evenly across females (Supplementary figure 3). Exceptions
are AM 2 at Lajuma and Govu at Mawana. Both of these males were non-dominant during data collection
and spent almost equal amounts of their time with all females in their group. Furthermore, visual inspection
of the total amount of times a male was found within 5m vicinity of a female reveals a difference between
males in the frequency in which they were found near females in both study populations and the total amount
of time a female had a male neighbour was variable between females as well.
Male proximity within 5m to females
Lajuma
4
AM 1* AM 2 AM 3*
Standardized AI
3
2
1
-2 -1 0
1 7 6 5 3 2 1 7 6 5 3 2 1 7 6 5 3 2
Female
Mawana
-1 0 1 2 3 4
Gov Twe* Ati* Wol Can

Standardized AI
-3
Up Ge Ga Pr Ro Re Up Ge Ga Pr Ro Re Gu Nk Gu Nk Gu Nk
Female
Supplementary figure 2 Standardized values of the association index for male proximity to females within 5m in both study
populations. Female are ordered according to their rank (highest ranking females on left side) on the x-axis. Differently coloured
bars represent different males and asterisks next to male names in the legend indicate dominant males.
iii
Demography-corrected mating indices

Supplementary table 1 shows how corrected mating indices were calculated with the example of MI male in
the Lajuma population to evaluate the role of demography on male mating success. The results of the mating
index calculations as described in the main text were multiplied with a correction factor (the average number
of male group members during each dyad’s co-residence) to take into account potentially enhanced diffi-
culties to obtain a high share of female matings. The correction factor varied slightly (between 5.7 and 6.0)
and thus the differences in ranks between the corrected and uncorrected mating index are minor.
Supplementary table 1 Illustration of how corrected mating indices were calculate with the example of MImale .
Female Male Mating Index Correction factor Corrected MI Rank Rank

(F) (M) (MImale) (CF) MIcorr MI MIcorr.
As used in the statistical Average number of males MImale * CF Ordered by value Ordered by value
analysis in group during co-resi- 1: highest MImale 1: highest MIcorr
dence of F and M
AF 1 AM 1 0.111 6.0 0.666 10 10

AF 2 AM 1 0.500 6.0 3.000 02 02
AF 3 AM 1 0.000 6.0 0.000 12 12
AF 5 AM 1 0.000 6.0 0.000 12 12
AF 6 AM 1 0.250 6.0 1.500 08 08
AF 7 AM 1 0.500 6.0 3.000 02 02
AF 1 AM 2 0.364 6.0 2.184 07 07
AF 2 AM 2 0.000 6.0 0.000 12 12
AF 3 AM 2 0.200 5.7 1.140 09 09
AF 5 AM 2 0.000 5.7 0.000 12 12
AF 6 AM 2 0.400 5.7 2.280 06 06
AF 7 AM 2 0.100 5.7 0.570 11 11
AF 1 AM 3 0.000 6.0 0.000 12 12
AF 2 AM 3 0.000 6.0 0.000 12 12
AF 3 AM 3 0.000 5.7 0.000 12 12
AF 5 AM 3 0.500 5.7 2.850 02 04
AF 6 AM 3 0.500 5.7 2.850 02 04
AF 7 AM 3 0.636 5.7 3.625 01 01
Analysis of situations with a potentially higher value of male presence

ADDITIONAL ASSOCIATION INDICES
Proximity to males is hypothesized to be useful to females regarding predation risk. One goal of this study
was to identify and analyse situations in which male spatial association is particularly relevant to females.
Apart from feeding situations (see main text), females with a depending infant could be more vulnerable to
predation risk or relying more on high quality food sources and hence evaluate male tolerance to be more
valuable. Moreover, situations in which individuals are not located in a refuge could also require more
vigilance and lead to females assigning additional value to male proximity. To analyse the effect of tolerance
in these potentially sensitive situations, two additional association indices were calculated analogous to the
method described in the main text for both study populations: one taking into account only females who had
an infant of less than one year of age and one including only situations outside of refuges. In the calculation
of these indices, both general proximity and feeding situations in particular were taken into consideration
iv
which resulted in four additional association indices. Using the same method as in the main analysis, all
association indices were correlated with the mating indices for female copulations to investigate whether
males who spent more time in the proximity of mothers with a depending infant or provided tolerance out-
side of refuges were gaining a higher mating success. Refuges were operationally defined to be trees with a
minimum height of 5m at both field sites.
The results of the correlation analysis show that in the Lajuma population, a male’s copulation success was
not related to the amount of times he was recorded to be within 5m of a female. This holds for both the
analysis restricted to females with infants as well as the analysis of situations outside of refuges.
Males who spent more time in the proximity of mothers with depending infants received a significantly
lower share of those females’ matings at Mawana. In situations outside of refuges, the correlation between
a male’s 5m spatial association with a female and the proportion of that female’s matings did not reach
statistical significance (Supplementary table 2).
Supplementary table 2 Kendall correlation coefficients and bootstrap confidence intervals of the statistical relationship be-
tween male proportion of female matings and spatial association in 5m with mothers of depending infants as well as associa-
tion outside of refuges.
Study population Parameter Restriction Effect size (τ) 95% CI

Lajuma AIoverall Females with infants -0.023 -0.501 0.419
AIfeeding Females with infants -0.091 -0.499 0.336
AIoverall Situations outside of refuge 0.064 -0.365 0.456
AIfeeding Situations outside of refuge -0.080 -0.491 0.317
Mawana AIoverall Females with infants -0.524 -0.813 0.125
AIfeeding Females with infants -0.598 -0.910 -0.063
AIoverall Situations outside of refuge -0.024 -0.328 0.296
AIfeeding Situations outside of refuge -0.160 -0.454 0.162
According to the results above, neither the presence of an infant nor the location outside of a refuge was a
relevant parameter for females to reward male tolerance with higher mating access. The negative correlation
between mating success and male spatial association with mothers of depending infants in the Mawana
population indicates that females with infants might even assign a negative value to male presence. One
explanation for this finding could be that females in some cases regard male presence to be a threat rather
than a benefit, e.g. concerning infanticide.
It should be noted that the data collection on the Mawana population started during a time where the young-
est infants were already about 6 months old and ended before the onset of the next birth season. It is possible
that male-female proximity is not a reliable indicator of male-infant proximity due to a more frequent spatial
separation between mother and infant with progressing infant age. Moreover, imposing further restrictions
on the analysed data leads to a calculation based on a strongly reduced amount of data points and these
results should hence not be regarded as statistically meaningful.
PROPORTION OF TIME SPENT FEEDING PER FEMALE
If male presence leads to a higher feeding efficiency, the proportion of a female’s time allocated to feeding
could be increased in situations where a male is in proximity. Supplementary figure 2 indeed shows that
females at Lajuma on average spent a higher share of their time feeding when the closest individual in
proximity was an adult male compared to those situations in which the nearest neighbour was an adult
female. This finding holds in the overall data set as well as in a separate analysis of females with infants as
well as situations outside of refuges. However, the variation between female time allocation to feeding was
considerable in all three analysed data subsets and not all females increased their proportion of time feeding
when the nearest neighbour was male rather than female.
v
Male proximity within 5m to females
a) Overall b) Females with infants

0.6
0.6
median Female median Female
mean AF 1 mean AF 1
AF 7 AF 7
0.5
0.5
AF 6 AF 6
Proportion of time feeding

AF 4 AF 4
AF 3 AF 3
AF 2
0.4
0.4
0.3
0.3
0.2
0.2
0.1
0.1
p  0.115 p  0.091
0.0
 2  4.333 df  2 2  4.8 df  2
0.0
Female Male None Female Male None
Neighbour Neighbour
c) Situations outside of refuges

0.6
median Female
mean AF 1
AF 7
0.5
AF 6
AF 4
AF 3
AF 2
0.4
0.3
0.2
0.1
p  0.847
0.0
 2  0.333 df  2
Female Male None

Neighbour
Supplementary figure 3 Female focal individual’s time allocation to feeding depending on nearest neighbour’s sex in the Lajuma
population. The y-axis shows the proportion of focal sample points in which each female individual’s activity was feeding in the
overall data set (a) as well as restricted to sample points featuring females with infants (b) and situations outside of refuges (c). For
each data subset, proportion of time feeding was calculated for three categories of sample points: 1) the nearest neighbour was an
adult female, 2) the nearest neighbour was an adult male and 3) there was no neighbour within 5m of the focal individual. Females
are listed in order of their rank with the highest ranking female on top. The plots include results of a Friedman’s ANOVA analysing
the differences between the proportions of time feeding regarding each of the three neighbour conditions.
Moreover, females overall spent a higher proportion of their time feeding when there was no neighbouring
individual in 5m compared to situations in which a female was located within 5m. This finding was found
also for females with a depending infant and situations outside of refuges. The increased feeding activity in
vi
the absence of neighbours indicates that rather than a promoting effect of male neighbours on feeding ac-
tivity, females in vicinity might impose restrictions on a focal female’s food intake. The results of Fried-
man’s ANOVA reveal no statistically significant differences between the proportion of female’s time spent
feeding when there was a female neighbour, a male neighbour or no neighbour in the overall data set (χ2 =
4.333, df = 2, p = 0.115), when analysing females with infants (χ2 = 4.8, df = 2, p = 0.091) and in situations
outside of refuges (χ2 = 0.333, df = 2, p = 0.847).
Reference
Baldellou M, Henzi PS, 1992. Vigilance, predator detection and the presence of supernumerary males in
vervet monkey troops. Animal Behaviour 43:451-461.
vii
Declaration of authorship
I declare that I have used no sources and aids other than those indicated. All passages quoted from publica-
tions or paraphrased from these sources are indicated as such, i.e. cited and/or attributed. This thesis was
not submitted in any form for another degree or diploma at any university or other institution of tertiary
education.
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Male services and mating success

Sexual selection as instigator of male

Material and Methods

DETERMINANTS OF FEMALE MATING SUCCESS

Study population Parameter Effect size (τ) 95% CIBCa

Study population Parameter 1 Parameter 2 Effect size (τ) 95% CI BCa

LOW-COST MALE SERVICES AND MATING SUCCESS

SPATIAL ASSOCIATION AND SOCIAL DOMINANCE

Male services and reproductive success

Figure 1 The provision of male services could promote male repro-

Sex differences in vigilance

Proportion of time vigilant

Inter-individual differences in spatial proximity to opposite-sex adults

Male proximity within 5m to females

Gov Twe* Ati* Wol Can

Demography-corrected mating indices

Female Male Mating Index Correction factor Corrected MI Rank Rank

AF 1 AM 1 0.111 6.0 0.666 10 10

Analysis of situations with a potentially higher value of male presence

Study population Parameter Restriction Effect size (τ) 95% CI

Male proximity within 5m to females

a) Overall b) Females with infants

Proportion of time feeding

c) Situations outside of refuges

Female Male None

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