This document describes a study that investigated whether maternal mongooses preemptively alter testosterone levels in their offspring based on predicted post-birth conditions. The study found that pup mongooses born into larger groups had significantly longer ano-genital distances (AGD), a marker for testosterone, compared to pups from smaller groups, even after controlling for body size. Pups with higher AGD were also more likely to survive past 6 months. This provides evidence that maternal mongooses can preemptively prepare offspring for future competitive conditions by altering testosterone levels before birth.

1. Early life testosterone as pre-emptive
preparation for post-birth conditions in a
cooperative mammal
Running Title:- Predictive Maternal Effects
Author:- Katherine Suzanne Appleby, University of Exeter, ksa208@exeter.ac.uk
Type of Publication:- Ecol. Lett.
Keywords:- Adaptive, Cooperative, Pre-emptive Conditioning, Maternal Effects, Testosterone, Intraspecific
Competition
Word Count:- Abstract: 147, Text: 3859, Text Boxes: 167
References:- 50
Figures:-4
Tables:-1
Text Boxes:-5
Correspondence:- Author for Correspondence, K.S.Appleby, Penryn Campus, University of Exeter, Cornwall
Statement of authorship:-
KA performed all data manipulation, analysis and report writing, subsequent to a primary data
collection courtesy of the Banded Mongoose Research Project.

2. 1
Abstract.
The concept that maternal parents can prepare unborn offspring for predicted post-natal conditions by
manipulating their phenotypic expressions, is an exciting, far-reaching idea. Previous studies have
highlighted testosterone levels as a phenotypic trait that may be specifically manipulated by mothers in
anticipation of future conditions. Here, I explored the possibility that pre-emptive, maternally controlled
testosterone variation features within the cooperative mammal, Mungos mungo.
Using an associated morphological testosterone marker, Ano-Genital Distance (AGD,) I attempted to
determine the existence of a correlation between marker and the changing post-birth condition of group
size. Testosterone, measured as AGD, was noted in relation to group size. Analysis showed that pups born
into larger groups reported significantly increased AGD’s irrespective of body-size, and that these individuals
were more likely to survive post 6 months. These findings provide evidence that adaptive, pre-emptive
maternal preparation of unborn juveniles, can be witnessed in mammalian structures.

3. 2
Introduction.
Scientists and naturalists alike have long documented the juvenile stages of life across a range of
taxa. These juvenile stages of life are typically characterised by intense competition and high
mortality [Sulivan 1989; Clutton Brock et al 1985]. Substantial parental investment means that
ensuring the survival of young, particularly through these hazardous times, is crucial to parents in
order to maximise their own fitness. The importance of this insurance gives rise to various
mechanisms designed to maximise offspring survival, increasing the likelihood of eventual
reproduction in a second generation, as well as providing fitness benefits to a parental lineage
[McGraw et al 2005, Sargent 1988].
Maternal effects have been established as one process by which juvenile safety can be enhanced.
Defined as a phenotypic effect within an organism, transferred from parent to offspring, irrespective
of genotype, this “conditioning” can provide much needed protection from the threats presented to
juveniles [Wells 2007; Altmann & Alberts 2005]. These enhanced conditions can provide long lasting
benefits to an organism, continuing beyond infancy and into adult life [Wu et al 2004; Bernardo
1996]. Maternal effects can be adaptive, resulting in the divergence of offspring away from the
phenotypic average that would have arisen as a consequence of a parent’s typical genetic
contribution to a developing zygote. This divergence will potentially facilitate increased fitness,
reflected in both current and future generations via increased safety to juvenile organisms [Allen et
al 2008; Lindström 1999; Bernardo 1996; Mather & Jinks 1971]. In species with a particularly high
level of infant mortality, and/or a heightened risk of threat, this early advantage would seem to be
especially beneficial, to both parent and offspring alike.

4. 3
Maternal effects occur within four distinctly classified phases of offspring development; pre-
reproductive, early reproductive, late reproductive and post-reproductive [Russel & Lummaa 2009].
Evidence for the existence of maternal effects is unevenly established across all four stages,
however, an inequality extending to taxa-specific knowledge [Räsänen & Kruuk 2007; Bernado
1996]. This imbalance is in part, due to the scale of clarity with which maternal effects within these
different phases present themselves. Effects occurring within post-natal phases can be observed
with greater ease than those pre-natal. Organisms within developmental stages of pre-natal life, are
frequently inaccessible, hidden from predators either internally within a mother’s womb or within
complex burrows, nests and so on. Observation requires the use of internal, penetrative techniques,
potentially hazardous to the unborn.
Variation in ease of observations of maternal effects also differs with taxonomic classification. The
clarity with which oviparous species present their young, in comparison to viviparous companions,
causes inequality in the ease with which data can be collected. [Räsänen & Kruuk 2007; Bernardo
1996]. Parental provisioning of resources to a nest of bird chicks is a clearly visible maternal effect,
whereas observation of increased placental flow of oxygen to an unborn foetus is one requiring the
use of possibly invasive equipment, a potentially more time and resource costly process. In recent
times however, these difficulties have been somewhat addressed, due in part to the advancement of
technologies, allowing for comprehensive, detailed examination of even the most impenetrable and
illusive species [Cowley 1991]
These advancements in the ease with which maternal effects can be studied, has led to increasing
interest in the field, specifically those within pre-natal phases, which may prove valuable in
maximising juvenile survival. Prenatal maternal alterations to developing offspring phenotypes have

5. 4
been chosen, based on the conditions that the mother anticipates to arise after birth. Pre-emptive
conditioning has the potential for adaptive value across a wide variety of systems. However, there
are a number of conditions under which its use could be arguably, most valuable. Firstly, it would
seem reasonable to predict that any system so reliant on predictions, would perform optimally
under stable, predictable environmental conditions, with minimal variation [Reed et al 2010; Tufto
2000; Mosseau & Fox 1998]. Any system that experiences large deviations from this optimum, would
likely be beyond the capability of this pre-emptive maternal effect mechanism. The phenotypic
plasticity required would almost certainly be unable to keep pace with any system that experienced
such a dynamic, ever changing environment. Despite research suggesting that maternal effects may
be capable of fast adaption in the face of change, parents would simply be unable to make the
quantity of alterations required to successfully “tune” its unborn young to the conditions [Heath et
al 2003]. Any alterations made in response to these changes, would likely be rapidly outdated.
A second scenario may be considered in those species exhibiting limited generation times. Such
situations give rise to restrained variation in the range of conditions which may be witnessed during
an organism’s lifetime. While such systems are usually characterised by rapid reproductive rates,
reducing the need for maximised survival rate per individual, this decreased variation would seem an
appropriate condition under which such a mechanism could best operate.
The next scenario, where pre-emptive conditioning could be considered as an especially high value
adaptation, is in those K-selected species systems, where reproductive litters are limited in both size
and frequency. In such systems, where reproductive efforts are limited by high levels of parental
investment, the adoption of a strategy that would ensure safe voyage into adulthood would seem

6. 5
most prudent. In this way, reproductive fitness may be maximised, a necessity considering the
exceptional cost incurred to respective parents of losing offspring before their development.
The last condition under which such a mechanism seems naturally well-suited, is where the juvenile
mortality threat imposed by the surrounding community and its conditions, is considerably higher
than typical background rates. Increased mortality may translate itself via threats in the form of
extreme environmental conditions, competition, predation and disease transmission. In such
circumstances, where mortality rates are excelled, enforcement of pre-emptive conditioning could
lead to an increase in survival and related fitness of parents who possess this capability.
Previous research regarding the existence of pre-emptive maternal effects have frequently cited
hormonal variation as one method of maternal alteration of an offspring’s phenotypic development.
This alteration is in light of anticipated conditions, mothers predict to occur post-birth. A range of
species, including the Crocuta crocuta (spotted hyena) and Pomacentrus amboinensis (tropical
damsel fish) have been shown to exhibit maternally controlled hormonal variation [Dloniak et al
2006; McCormick 1999]. Hormones, specifically testosterone have a proven variety of links to
physical traits such as strength and competing ability. Variation in these qualities could dramatically
alter an organisms suitability to its surrounding conditions. Consideration to the competitive, harsh
environment juveniles face post-birth, therefore highlights maternal alteration of testosterone, as a
valuable outlet of predictive maternal effects. [Tschirren et al 2009; Groothuis & Schwabl 2007]
Understanding of pre-emptive hormonal preparation will enable more comprehensive
understanding of the flexibility and adaptability of populations faced by changing environmental
conditions, in addition to respective limitations. The existence of inter-disciplinary links this system

7. 6
provides to other bio-scientific fields would place any findings into significance within the realms of
evolutionary, physiological and conservation biology [Wolf & Wade 2009]. By defining these
relationships, deeper comprehension of the natural world’s tolerance of change might be achieved,
fundamental in aiding our understanding of how species might with-stand environmental instability
[Vissel 2008; Galloway 2005].
Here, I expand upon the potential role of testosterone variation as an effective way of mothers
increasing the suitability of offspring phenotype to environmental conditions present. Testosterone
measured by way of an associated morphological marker, Ano-Genital Distance (AGD,) was
measured in relation to changing pre-natal condition, group size. A data set was drawn from an
existing study site of wild M. mungo, taken over a period of 6 years and containing over 400
individuals. Alongside this response variable, survival after 6 months was noted per individual, in
order to determine whether any adaptive advantage in any variation in AGD existed. Additionally,
levels of rainfall over the 6 months prior to gestation, were also noted, to account for variation in
environmental conditions. Head width per organism was also recorded, by way of controlling for the
effect of body size, along with individual sex.

8. 7
Materials & Methods.
472 individual M. mungo’s from 85 litters across 9 groups were sampled across 5 years, from 2007 to
2012 from the study site. All animals were captured, and individually identified after first observing
from the den (aged approx. 4 weeks.) Data was compiled into one set before being analysed
between September and December 2014. Statistical analysis was performed using the statistical
software programme, R. A mixed effects GLMM and an LM model, using the package lm4e were
used, given the potential for over-lapping effects in pack and group. All data was examined for
normality, linearity and heterogeneity, after which no alterations or transformations were deemed
necessary.
1) Study Species - All individuals recorded as part of these observations, came from existing data
originating from the Banded Mongoose Research Project, a wild population under long term
observation. The project, based on the Mweya peninsula, Uganda (0 12'S; 27 54'E), has been
established over twenty years and features an amalgamation of scientific contribution from several
European institutions. M. mungo are a highly cooperative species, found living in communal groups
of around 15-20 individuals, a figure highly variable under changing conditions [Gilchrist & Otali
2002; De Luca & Ginsberg 2001; Pienaar 1964]. The study site featured 12 differing social groups,
scattered across a 4.95km2
area, of which 9 groups and their corresponding individuals were
observed. Primarily inhabiting savannah and woodland ecosystems endemic to Africa, the
organism’s cooperative breeding tendencies, are one reason why they have become a highly
researched study system [Cant et al 2013; Gilchrist & Russel 2007].

9. 8
2) Data-AGD- AGD was used as an associative measure of testosterone for the purposes of this
study. This was due to the limitations of the existing data set and the implications that invasive mass
testosterone sampling would involve. Previous literature has demonstrated the applicability of AGD
as a viable, indicative marker of an organism’s reproductive hormonal levels, specifically
testosterone [Fouqueray et al 2014; Eisenberg et al 2011; Wolf et al 2002]. During original capture,
still images were taken of the ano-genital region of all individuals. Pups were held a set distance
away from the camera for each photograph taken. A centimetre ruler was aligned alongside the
creature, and the image was taken, to include both ruler and organism. Care was taken to ensure
good quality, well focused and lit images were collected. Immediate repeat images of the same
individual were captured in nearly all cases, in order to ensure optimum visual quality.
AGD Measurement- Collected images were analysed in Autumn 2014, using the imagery software
package, Imagej. Using the cm ruler in shot, Imagej was calibrated to a set measurement of 10mm.
This calibration was repeated every 10th
measurement, with data analysis only taking place between
the hours of 9 am and 12pm daily. These boundaries aimed to control for human error, as a result of
variation in concentration. Each image was measured three times, after which a mean mm length
average was calculated. Where repeat images of an individual existed, each was measured as noted
before, after which a mean average was taken. Despite AGD’s use as an associated testosterone
marker across a wealth of previous literature, its quantifiable definition varies. This dilemma appears
to be accountable, firstly to interpretation on behalf of the measurer, and secondly, the physiology
of the study subject in hand. Varying anatomical limitations mean achieving an accurate ratio,
without resorting to intrusive methodology, is challenging. Such difficulties requires the adoption of
a flexible definition. A densely furred appearance and the posture in which animals were captured,
meant that for the purposes of this investigation, the definition appropriated was as follows:- “mid-
anus to the base of the scrotum in males and from the mid-anus to the base of the posterior

10. 9
fourchette in females” [Eisenberg et al 2011; Sathyanarayana et al 2010]. Collected data was then
statistically analysed. Due to the existence of the repeated measures pack and group, they were
included as random effects within a Linear Mixed Model, which was used as base for statistical
analysis. A model was generated, in order to establish effects differing variables had upon AGD. This
model was composed of the variables Rainfall, Group-size, Head-width and Sex. AGD was fitted as
the continuous response variable, before an Anova and drop1 function were completed.
3) Data-Survival- During original data collection, recapture of pups was attempted at a stage 6
months post birth. Those pups repeatedly absent from any recaptured groups, as well as those
known to be deceased, were recorded as dead. In order to investigate for any possible adaptive
advantage in AGD variation, a second statistical model, a GLMM was produced, with random effects
pack and group included. Survival was fitted as a binomial response variable, and categorised
numerically. Those survived were allocated a 1, while those deceased were represented by 0.
Survival was then contrasted against AGD, Rainfall, Group-size, Head-width and Sex. Following this,
an Anova and drop1 function were completed.

11. 10
4) Other Response variables- The following were used as response variables in both the AGD based
LMM model, and the Survival based GLMM model.
Group Size-Variation in group size represented the predominant changing post-natal condition. Each
pup born into a group was noted, and the total group size was recorded. Group size was then set as
a binomial variable within both models.
Head width-Head width was measured, via the use of digital callipers, to the nearest closest 0.1 mm.
Recordings were standardized by measuring the width of the widest point of the skull. This point is
clearly visible by the presence of M. mungo scent glands, sited directly below this point on the
cheek. Head width was included as a binomial variable in both models, by way of controlling for
differing body size contributing to variations in AGD.
Rainfall-Rainfall was measured across gestation (60 days prior to birth,) utilising the mm
measurements recorded by weather station Mweya, central to the study site. These recordings were
accounted for by way of a binomial variable in both linear models.
Group sex proportions-Sex was denoted by F = female, M=male. Variation in the categorical data set
was tracked throughout the entire collection, included in both models.

12. 11
Results.
Testosterone via AGD and group size.
Pups born into larger groups were found to have significantly increased AGD measurements, and by
association, testosterone levels, in comparison to pups from smaller groups (X2
1=13.022, df=1,
p<0.001, Fig 1.)
Testosterone via AGD and head width.
In keeping with previous research, pups of a larger head width, and associated body size, reported
statistically significant increased AGD measurements across both sexes (X2
1=14.773, df=1, p< 0.001,
Fig 1 & 2.) This control measure ensured variations in body size were not soley responsible for
variation within AGD.
Testosterone via AGD and Group sex proportions.
Concurrent with known information regarding inter-sexual Testerone variation, variation in
proportion of the sexes within groups had a significant effect upon individual pup AGD
measurements (X2
1=148.252, df=1, p< 0.001). Males reported an average increase in AGD of 11.6%,
in comparison to their female conterparts. Total sex proportion across the data set of 472 individuals
equalled 231:241, female:male.

13. 12
Testosterone via AGD and rainfall.
Variation in total rainfall witnessed during gestation (60 days prior to birth,) was found to have no
significant effect upon AGD (X2
1=0.06, df=1, p= 0.807,) despite a variation of 502% in the amounts
received over the 5 year period
Survival and Testosterone via AGD.
Pups which reported increased AGD recordings, and therefore associated testosterone levels, were
found to have a significantly elevated chance of surviving beyond 6 months of life (X2
1=3.89, df=1 p=
0.0486, Fig.2.)
Survival and Others-Head Width.
A significant, positive correlation was witnessed between survival and individual head width (X2
1=
25.9592, df=1 p<0.001, Fig 3.) Pups found alive post 6 months of their life were to found to have
significantly increased head widths.
Group size.
Group size was found to have no significant effect on survival, despite variation of 200% observed
between largest and smallest groups (X2
1=0.005, df=1 p= 0.944.)
Rainfall
Variation in rainfall was found to have no effect on survival (X2
1=3, df=1, p= 0.0832)

14. 13
Group sex proportions
Differing sex proportions within groups were found to no have statistically significant effect upon
survival (X2
1=0.126, df=1, 0.723)
Additionally, a table of summary statistics regarding the data set was generated (Table 1).

15. 14
Discussion.
M. mungo pups born into larger groups reported significantly increased Ano-genital distance, and by
association, testosterone levels, than conspecifics born into smaller groups. Pups reporting increased
AGD measurements and associated testosterone levels, experienced significant increases in survival
rates beyond the first 6 months of life. The fact that the effects of group-size upon AGD were found
to be strongly significant, even when body size was controlled, is imperative. Such findings would
suggest that this change in AGD is at least partially independent of individual body size, and gives
rise to the scenario of equal sized conspecifics, with differing AGD due to respective group sizes
(Fig.1, Fig.2).
These findings suggest pre-emptive maternal conditioning via increased testosterone input is
present within certain mammals at least, concurrent with limited, relevant available research
[Fouqueray et al 2014; Correa et al 2013]. The presence of this system here can be explained via
reference to the earlier mention operant conditions for predictive maternal effects. The costs
imposed by large-scale cooperative living, as observed in the M. mungo, act to increase mortality
rates from birth onwards. This risk matches conditions described by the third scenario, where pups
are threatened by exposure to heightened danger. Social living is rarely witnessed to the extent
observed in the M. mungo. The scale of this aggregation presents costs, one of which being the
disproportional relationship between group size and resource pay off available per head [Hixon &
Jones 2005; Polis 1984]. This scenario will inevitably lead to intense density-dependent intraspecific
competition. Pups will often be presented with the necessity to engage in physical battle with
conspecifics, in order to access essential resources, and consequently suffer from associated
increased mortality rates [Rood 2014; Johnson & Hubbell 1974]. Any mechanism or substance
capable of enhancing competitive performance would appear adaptive. Testosterone has been

16. 15
previously linked to higher levels of aggression within individuals, outlined as the “Challenge
Hypothesis,” and its intensified abundance can be attributed to increased growth, strength and
dominance [Muller & Wrangham 2004]. Such attributes can greatly increase an individual’s fitness,
via increased resource share, elevating their likelihood of survival and subsequent breeding
[Wingfield et al 1990; Marler & Moore 1988]. Maternally-controlled testosterone variation therefore
seems a suitable, adaptive and pre-emptive method of offspring preparation.
This variation in both associated resource share and fitness, translates into the elevated post six
month survival rates witnessed in those pups with increased AGD’s. These pups, reporting increased
testosterone and associated AGD, appear to have been more successful in avoiding the risks
presented, in comparison to conspecifics from smaller groups, with lower testosterone and smaller
AGD measurements. This enhanced preparation as a result of pre-natal maternal conditioning within
the womb, has allowed these pups to develop superior phenotypes compared to their conspecifics.
This maternal effect clearly increases the fitness of both the original parent and that of future
generations, via the extended survival rates afforded by an individual’s enhanced performance. This
provides evidence for increased testosterone levels, especially when present within larger group
sizes, being an adaptive mechanism. Such a system could be assumed to be present among many
other species, where intra-specific competition challenges members throughout their lifetime.
Positive correlation was also reported present between survival and head width. Those pups with
increased head width, and assumed overall body size, were more likely to survive beyond 6 months,
than those with smaller measurements. Such findings are indicative of widely established
relationships between size and traits such as strength and defence, highlighting the value of
increased size in the light of limited resources [Allen et al 2008; Hendrickx et al 2003]. In accordance
to similar results within other taxa such as a variety of both crustaceans and insects, larger pups are

17. 16
more likely to possess improved physical capabilities, enabling successful encounters with intra, as
well as interspecific competitors [West-Eberhard 1979].
Increased rainfall was found to lack any significant correlation with neither variations in AGD, nor
survival, conflicting with expectations founded from previous research, from a variety of societies
including avian and human [Hinde et al 2009; Gluckman et al 2005; Welberg et al 2001]. Initial
investigation identified this variable as a possible indicator of environmental conditions, such as food
availability. It was hypothesised that increased levels of rainfall would positively correlate to food
availability. This would allow for a maternal prediction to be made concerning the post-natal
conditions that may face her offspring, resulting in a changed AGD. However, statistical analysis
failed to provide any evidence suggesting this was the case. While rainfall may indeed be an
indicator of pre-natal conditions, it is possible that these do not provide an accurate forecast of post-
birth conditions. If this were the case then pre-emptive preparation of unborn young, based on the
conditions experienced prior to birth, could result in an out-dated, ill-fitting phenotypic set up. A
second explanation lies in the possibility that the effect of varying conditions are displayed
elsewhere in the pup’s geno/phenotype, aside from hormonal components as investigated here.
Distinguishing between genetically inherited traits triggered in response to environmental variation,
and deliberately maternally-inflicted pre-emptive phenotypic alteration is challenging, however.
[Räsänen & Kruuk 2007].
Group sex ratio was found to have significant effect upon AGD and associated testosterone. Males
witnessed an average increase in AGD ratios of 11.6% in comparison to females, a point explained
via hormonal levels as a sex controlled trait [Hau 2007; Clotfelter et al 2004]. Variations in group sex
ratio were found to have no effect upon the survival rate of individuals during the first 6 months of
life. While variation in male/female group proportions may be thought to increase in-group fighting,

18. 17
this a rare occurrence, with group size appearing to be a larger influence for intra-group aggression,
as shown in a variety of species including, Neolamprologus pulcher, (Cichlid) [Balshine et al 2001].
In summary, these results highlight the effect that intraspecific competition can have at a
fundamental level, and provide some of the first mammalian based evidence for pre-emptive
conditioning in unborn young, based on pre-natal conditions. This highlights the potential for future
work establishing the extent to which this system is present within the natural world. Full
appreciation will enable a wide variety of benefits and expansions in ability across multiple areas of
animal behaviour and beyond, within physiological, evolutionary and genetic contexts. By
establishing maternal effects as a key cause of variation within organisms, a more accurate
understanding of the mechanisms upholding speciation and diversification may be afforded
[Bernardo 1996.] This is fundamental in aiding wider understanding of the flexibility, plasticity and
adaptability that species possess. Application of this knowledge within a conservation context will
enable enhanced population modelling and predictions in the face of globally changing ecosystems.
By establishing the capabilities and limitations of such mechanisms, researchers may hope to gain an
ever increasingly realistic picture of how the natural world will evolve to these dramatic changes,
and as importantly, why.

19. 18
Acknowledgments.
The author would like to thank the following for their valuable contributions to this piece:- Prof. M
Cant for his much appreciated supervision, along with Dr E Vitikainen who provided scrupulous
assistance and F Thompson, whose contribution to statistical analysis was invaluable. The data set
from which this piece of work originated comes from the Banded Mongoose Research Project,
Uganda, and as such all members of team must be thanked for their thorough data collection. This
piece was fully funded as part of the University of Exeter’s undergraduate programme.

20. 19
Figures.
6
7
8
9
10
11
12
13
20 25 30 35 40
AGD(mm)
Head width (mm)
Female
Large Group, (17+)
Small Group (0-16)
Fig 1. The scatter plot demonstrates the significant positive
correlation between AGD and head width in female, M. mungo
(X2
1= 14.773, df=1, p<0.001.) Positive correlation between
individual AGD measurements originating from differing group
sizes is also shown. (X2
1= 13.022, df=1, p<0.001)
(17+)
(0-16)

21. 20
Fig 2. The scatter plot demonstrates the significant positive
correlation between AGD and head width in male, M. mungo
(X2
1= 14.773, df=1, p<0.001.) Positive correlation between
individual AGD measurements originating from differing group
sizes is also shown. (X2
1= 13.022, df=1, p<0.001)
6
7
8
9
10
11
12
13
14
15
20 25 30 35 40
AGD(mm)
Head width (mm)
Males
Large Group, 17 +
Small group, 0-16
(17+)
(0-16)

22. 21
Fig 3. The bar chart displays the significant variation in survival status of M.
mungo pups in comparison to AGD (X2
1=3.89, df=1 p= 0.0486) (SEM;
deceased after 6 months, 0.0835, survived after 6 months, 0.0754.)
9.7
9.8
9.9
10
10.1
10.2
10.3
10.4
10.5
10.6
Deceased After 6 Months Survived After 6 Months
AvergeAGD(mm)
Survival Status
27
27.5
28
28.5
29
29.5
30
Deceased After 6 Months Survived After 6 Months
HeadWidth(mm)
Survival Status
Fig 4. The bar chart displays the significant variation in survival status of M.
mungo pups in comparision to Head Width (X2
1= 25.9592, df=1 p<0.001)
(SEM, deceased after 6 months 0.134, survived after 6 months 0.124.)

23. 22
Category Measurement Result
AGD Average Female (mm) 9.73
Average Male (mm) 10.87
Average Total (mm) 10.31
Survival Average Female (%) 50.22
Average Male (%) 49.58
Average Total (%) 49.90
Rainfall Average during gestation (mm) 145.93
Variation (%) 502
Head Width Average Female (mm) 28.97
Average Male (mm) 29.23
Average Total (mm) 29.10
Group Size Average Size (no. individuals) 9.85
Variation (%) 950
Table 1. A table displaying key summary statistics taken from the
data set.

24. 23
Reference List
1) Allen, R. M., Buckley, Y. M., & Marshall, D. J. (2008). Offspring Size Plasticity in Response to
Intraspecific Competition: An Adaptive Maternal Effect Across Life‐History Stages. Am Nat, 171, 225-
237.
2) Altmann, J., & Alberts, S. C. (2005). Growth Rates in a Wild Primate Population: Ecological Influences
And Maternal Effects. Behav Ecol Sociobiol, 57, 490-501.
3) Balshine, S., Leach, B., Neat, F., Reid, H., Taborsky, M., & Werner, N. (2001). Correlates of Group Size
in a Cooperatively Breeding Cichlid Fish (Neolamprologus pulcher), Behav Ecol Sociobiol, 50, 134-140.
4) Bernardo, J. (1996). Maternal effects in animal ecology. Am Zoo, 36, 83-105.
5) Cant, M. A., Vitikainen, E., & Nichols, H. J. (2013). Demography and social evolution of banded
mongooses. Adv Stud Behav, 45, 407-445.
6) Clotfelter, E. D., O'Neal, D. M., Gaudioso, J. M., Casto, J. M., Parker-Renga, I. M., Snajdr, E. A., et al.
(2004). Consequences of elevating plasma testosterone in females of a socially monogamous
songbird: evidence of constraints on male evolution? Horm Behav, 46, 171-178.
7) Clutton-Brock, T. H., Albon, S. D., & Guinness, F. E. (1985). Parental investment and sex differences in
juvenile mortality in birds and mammals. Nature, 313, 131 – 133.
8) Correa, L. A., Frugone, M. J., & Soto-Gamboa, M. (2013). Social dominance and behavioral
consequences of intrauterine position in female groups of the social rodent. Physiol behav, 119, 161-
167
9) Cowley, D. E. (1991) Prenatal effects on mammalian growth: embryo transfer results. 4th ICSEB
Proceedings, Vol. II. Dioscorides Press, Portland, pp. 762-779.
10) De Luca, D. W., & Ginsberg, J. R. (2001). Dominance, reproduction and survival in banded mongooses:
towards an egalitarian social system, Anim Behav, 61, 17-30.
11) Dloniak, S. M., French, J. A., & Holekamp, K. E. (2006). Rank-Related Maternal Effects of Androgens on
Behaviour in Wild Spotted Hyaenas. Nature, 440, 1190-1193.
12) Eisenberg, M. L., Hsieh, M. H., Walters, R. C., Krasnow, R., & Lipshultz, L. I. (2011). The Relationship
Between Anogenital Distance, Fatherhood, and Fertility in Adult Men, PLoS One, 6.
13) Fouqueray, T. D., Blumstein, D. T., Monclús, R., & Martin, J. G. (2014). Maternal Effects on Anogenital
Distance in a Wild Marmot Population. PloS One, 9.
14) Galloway, L. F. (2005). Maternal effects provide phenotypic adaptation to local environmental
conditions. New Phytologist, 166, 93-100.
15) Gilchrist, J. S., & Otali, E. (2002). The effects of refuse-feeding on home-range use, group size, and
intergroup encounters in the banded mongoose. Can J Zool, 80, 1795-1802.
16) Gilchrist J. S., Russell A. F.(2007). Who cares? Individual contributions to pup care by breeders versus
non-breeders in the cooperatively breeding banded mongoose (Mungos mungo). Behav. Ecol.
Sociobiol, 61, 1053–1060.
17) Gluckman, P. D., Hanson, M. A., Spencer, H. G., & Bateson, P. (2005). Environmental influences during
development and their later consequences for health and disease: implications for the interpretation
of empirical studies. Proc R Soc Lond B Biol Sci, 272, 671-677.
18) Groothuis, T.G.G., Schwabl, H., (2007). Hormone-mediated maternal effects in birds: mechanisms
matter but what do we know of them? Phil. Trans. Royal Soc. B, 363,1647–166.
19) Hau, M. (2007). Regulation of male traits by testosterone: implications for the evolution of vertebrate
life histories. BioEssays, 29, 133-144.
20) Heath, D. D., Heath, J. W., Bryden, C. A., Johnson, R. M., & Fox, C. W. (2003). Rapid evolution of egg
size in captive salmon. Science, 299, 1738-1740.

25. 24
21) Hendrickx, F., J. P. Maelfait, M. Speelmans, & N. M. Van Straalen. (2003). Adaptive reproductive
variation along a pollution gradient in a wolf spider. Oecologia, 134, 189–194.
22) Hinde, C. A., Buchanan, K. L., & Kilner, R. M. (2009). Prenatal environmental effects match offspring
begging to parental provisioning. Proc R Soc Lond B Biol Sci, 276, 2787-2794.
23) Hixon, M. A., & Jones, G. P. (2005). Competition, predation, and density-dependent mortality in
demersal marine fishes. Ecology, 86, 2847-2859.
24) Johnson, L. K., & Hubbell, S. P. (1974). Aggression and competition among stingless bees: field studies.
Ecology, 55, 120-127.
25) Lindström, J. (1999). Early development and fitness in birds and mammals. TREE, 14, 343-348.
26) Marler, C. A., & Moore, M. C. (1988). Evolutionary costs of aggression revealed by testosterone
manipulations in free-living male lizards. Behav Ecol Sociobiol, 23, 21-26.
27) Mather,K., & Jinks, J.L., (1971). Biometrical Genetics. The study of continuous Variation. Chapman and
Hall, London, UK.
28) McCormick, M. I. (1999). Experimental test of the effect of maternal hormones on larval quality of a
coral reef fish. Oecologia, 118, 412-422
29) McGraw, K. J., Adkins-Regan, E., & Parker, R. S. (2005). Maternally derived carotenoid pigments affect
offspring survival, sex ratio, and sexual attractiveness in a colorful songbird. Naturwissenschaften, 92,
375-380.
30) Mousseau T. A., & Fox C. W. (1998). Maternal effects as adaptations. Oxford University Press, Oxford,
UK.
31) Muller, M. N., & Wrangham, R. W. (2004). Dominance, aggression and testosterone in wild
chimpanzees: a test of the ‘challenge hypothesis’. Anim Behav, 67, 113-123.
32) Piennar, U.V . (1964). The small mammals of the Kruger National Park - A systematic list and
Zoogeography. Koedoe, 7, 1-25
33) Polis, G. A. (1981). The evolution and dynamics of intraspecific predation. Annu Rev Ecol Syst, 225-
251.
34) Räsänen, K., & Kruuk, L. E. B. (2007). Maternal effects and evolution at ecological time‐scales. Funct
Ecol, 21, 408-421.
35) Reed, T. E., Waples, R. S., Schindler, D. E., Hard, J. J., & Kinnison, M. T. (2010). Phenotypic plasticity
and population viability: the importance of environmental predictability. Phil. Trans. Royal Soc. B, 277,
3391–3400.
36) Rood, J. (2014). Ecology and Social Evolution in the Mongooses.In: (Ecological aspects of social
evolution: birds and mammals), { [Eds] [Rubenstein, D., Wrangham, R.W] }. Princeton University Press,
Princeton, pp. 131-153.
37) Russell, A. F., & Lummaa, V. (2009). Maternal effects in cooperative breeders: from hymenopterans to
humans. Philosophical Transactions of the Royal Society B: Biological Sciences, 364, 1143-1167.
38) Sathyanarayana, S., Beard, L., Zhou, C., & Grady, R. (2010) Measurement and correlates of ano‐genital
distance in healthy, newborn infants. Int J Androl, 33, 317-323.
39) Sargent, R. C. (1988) Paternal care and egg survival both increase with clutch size in the fathead
minnow, Pimephales promelas. Behav Ecol Sociobiol, 23, 33-37.
40) Sullivan, K. A. (1989) Predation and starvation: age-specific mortality in juvenile juncos (Junco
phaenotus). The J Anim Ecol, 275-286.
41) Tschirren, B., Sendecka, J., Groothuis, T. G., Gustafsson, L., & Doligez, B. (2009) Heritable variation in
maternal yolk hormone transfer in a wild bird population. Am. Nat, 174, 557-564.
42) Tufto J. 2000 The evolution of plasticity and nonplastic spatial and temporal adaptations in the
presence of imperfect environmental cues. Am. Nat. 156, 121–130.
43) Visser, M. E. (2008). Keeping up with a warming world; assessing the rate of adaptation to climate
change. Proc R Soc Lond B Biol Sci , 275, 649-659.

26. 25
44) Welberg, L. A. M., & Seckl, J. R. (2001) Prenatal stress, glucocorticoids and the programming of the
brain. J. Neuroendocrinol, 13, 113-128.
45) Wells, J. C. (2007). The thrifty phenotype as an adaptive maternal effect. Biol Rev, 82, 143-172.
46) West-Eberhard, M. J. (1979). Sexual selection, social competition, and evolution. Proc. Am. Phil. Soc,
222-234.
47) Wingfield, J. C., Hegner, R. E., Dufty Jr, A. M., & Ball, G. F. (1990). The" challenge hypothesis":
theoretical implications for patterns of testosterone secretion, mating systems, and breeding
strategies. Am Nat, 136, 829-846.
48) Wolf, C. J., Hotchkiss, A., Ostby, J. S., LeBlanc, G. A., & Gray, L. E. (2002). Effects of prenatal
testosterone propionate on the sexual development of male and female rats: a dose-response
study. Toxicol Sci, 65, 71-86.
49) Wolf, J. B., & Wade, M. J. (2009). What are maternal effects (and what are they not)? Philos T Roy Soc
B, 364, 1107-1115.
50) Wu, G., Bazer, F. W., Cudd, T. A., Meininger, C. J., & Spencer, T. E. (2004). Maternal nutrition and fetal
development. J Nutr, 134, 2169-2172.

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Appendix
A. Meeting Schedule:-
I met with project supervisor, Prof Michael Cant, and/or secondary supervisor Dr
Emma Vitikainen on the following occasions:-
 September 17th 2014-Project Outline
 September 22nd 2014-Additional Thoughts
 September 26th 2014-Data Set
B. Risk Assessment
Below is the accepted risk assessment for this project.

28. 27
WORKSTATION RISK ASSESSMENT FORM
HEALTH AND SAFETY (DISPLAY SCREEN EQUIPMENT)
REGULATIONS 1992
Assessment Form Reference: office use only:
User’s Full Name: Katherine Suzanne Appleby
College/Division/Dept/Unit: CLES Corwall
Line Manager/Supervisor: Mike Cant
Building & Room Number:
Site: Cornwall Campus
Date: 19/09/14
WORKSTATION ASSESSMENT CHECKLIST
(PLEASE USE THE GUIDANCE NOTES WHEN COMPLETING THIS FORM)
Risk Factors
Tick
answer
Yes / No
Comments
1. HISTORY
Any history of musculo-skeletal disorders
including back, neck and wrist pain?
Y Lupus and associated conditions
Any problems with vision (e.g. headaches,
focusing, reading screen, dry eyes)?
Y Lupus and associated conditions
Eyesight test for DSE in the last two years? Y
Has the user been advised of their entitlement
to eyesight testing/eye care voucher scheme?
Y

29. 28
www.admin.ex.ac.uk/personnel/occ_health/dse.shtml
Has any Health & Safety training been
undertaken?
N
2. USAGE
Average daily use (hours)? 2
Left or right handed? L
Activities other than using DSE? N
3. DISPLAY SCREEN (monitor)
Are the characters clear and readable? Y
Is the screen’s specification suitable for its
intended use?
Y
Is the text size comfortable to read? Y
Are the brightness and/or contrast adjustable? Y
Does the screen swivel and tilt? Y
Is the screen free from glare and reflections? Y
Is the user facing the screen? Y
Are adjustable window coverings provided and
in an adequate condition (e.g. blinds)?
Y

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4. KEYBOARD
Does the keyboard tilt? N
Are the characters on the keys easy to read? Y
Is there support for the user’s hands / forearms
infront of the keyboard?
N
Is the user a touch typist
(i.e. not a ‘hunt and pecker’)?
Y
5. MOUSE, TRACKBALL, MAT
Is the device positioned close to the user? Y
Does the device work smoothly at a speed that
suits the user?
Y
6. CHAIR
Is the chair suitable for the user i.e:-
Does the chair have a working:
• back height and tilt adjustment?
• lumbar support?
• seat height adjustment so that forearms
can be positioned horizontal to keyboard?
• swivel mechanism?
N
7. DESK

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Is the work surface large enough for all
necessary equipment, papers etc?
Y
Are surfaces free from glare and reflection? Y
Is there adequate leg room? Y
Can both feet be placed flat on the floor? Y
8. ENVIRONMENT
Is there enough room to change position and
vary movement?
Y
Is the lighting suitable, e.g. not too bright
or too dim to work comfortably?
Y
Are levels of heat comfortable? Y
Does the air feel comfortable? Y
Are levels of noise comfortable? Y

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C. Ethics
Below is the accepted, and ethics application for this project.