Familiarity breeds progeny: so
t m
*
ian C
sonia
tate
e on
contain an adult male year round, although most males live solitarily. We compared
Introducti
Animals ar
of group living exceed costs (Krause and Ruxton 2002).
adult females live in groups while adult males typically
pper 1996,
e Central
rican ring-
tailed coati (Nasua nasua) social groups typically contain
2009). The presence of adult males in social groups
mating season (Kaufman 1962, Booth-Binczik et al. 2004,
Hirsch 2007a). If an adult male is already associated with
Correspondence: Ben Hirsch;
E-mail: hirschb@si.edu
Molecular Ecology (2011) 20, 409?419 doi: 10.1111/j.1365-294X.2010.04940.xlive alone (Kaufman 1962; Smythe 1970; Russell 1982;
Gompper & Krinsley 1992; Gompper 1995). By living
alone, adult male coatis increase their foraging success
but live under increased predation risk and in some
could have important effects on the mating system and
distribution of reproduction within and between groups
of ring-tailed coatis.
In both coati species, adult males violently fight each
other for access to receptive females during the shortThese costs and benefits of sociality can differ according
to sex, leading to sexual segregation in some species
(Conradt 1998; Ruckstuhl & Neuhaus 2002). Coatis
(Nasua spp.) show strong patterns of sexual segregation:
one adult male throughout the year, while other adult
males live alone after dispersing from their natal group
at 2 years of age (Alves-Costa et al. 2004; Resende et al.
2004, Hirsch 2007a,b; Costa et al. 2009; Olifiers et al. 2010 Blackand groups of female coatis come into oestrus during the same 1?2 week period. During
the mating season, solitary adult males followed groups and fought with the group living
male. This aggression was presumably to gain access to receptive females. We expected
that high reproductive synchrony would make it difficult or impossible for the one group
living male to monopolize and defend the group of oestrous females. However, we found
that group living males sired between 67?91% of the offspring in their groups. This
reproductive monopolization is much higher than other species of mammals with
comparably short mating seasons. Clearly, living in a group greatly enhanced a male?s
reproductive success. At the same time, at least 50% of coati litters contained offspring
sired by extra-group males (usually only one offspring per litter); thus, resident males
could not prevent extra-group matings. The resident male?s reproductive advantage may
reflect female preference for a resident male strong enough to fend off competing males.
Keywords: coati, extra-pair paternity, mating success, Nasua nasua, paternity, reproductive
skew, reproductive synchrony, sociality
Received 28 May 2010; revision received 18 October 2010; accepted 21 October 2010
on
e predicted to live in groups if the benefits
cases have higher ectoparasite loads (Gom
2004; Hass & Valenzuela 2002). Unlike th
American species (Nasua narica), South Amedegree to which sociality affects reproductive success. Coati mating is highly seasonal
reproductive success of group living and solitary adult male coatis to determine thereproductive success in adul
(Nasua nasua)
BEN T. HIRSCH*?? and JESUS E. MALDONADO
*Center for Conservation and Evolutionary Genetics, Smithson
3001 Connecticut Ave., Washington, DC 20008, USA, ?Smith
AA 34002-9898, Barro Colorado Island, Panama, ?New York S
Abstract
The ring-tailed coati (Nasua nasua) is thwell Publishing Ltdciality increases
ale ring-tailed coatis
onservation Biology Institute, National Zoological Park,
n Tropical Research Institute, Unit 9100 Box 0948, DPO
Museum, CEC 3140, Albany, NY 12230, USA
ly coati species in which social groups
a social group, it may gain priority access to receptive males coexisting in a coati group outside the mating
do not exhibit a mating system that exactly replicates
410 B. T . HIRSCH and J . E . MALD ONADOfemales during the mating season. Because groups of
adult female coatis are able to exclude individual adult
males from dense food patches, females should be able
to evict unwanted adult males from their group (Gomp-
per 1996; Hirsch 2007b). Despite this ability, females in
most ring-tailed coati groups allow a male to enter their
group, even when male sociality does not appear to ben-
efit the group as a whole. No examples of adult male
parental care, such as protection from predators or food
provisioning, have been observed in ring-tailed coatis
(Di Blanco & Hirsch 2006; Hirsch 2007a,b). It is plausible
that male sociality is a function of female choice. If this is
true, one could predict that within-group males should
have higher reproductive success compared to solitary
males.
During the short 1?2 week mating season, an influx of
solitary male ring-tailed coatis typically enters and
follows the social group (Hirsch 2007a). Similar social-
mating pattern has been found in some primate species
such as patas (Erythrocebus patas) and blue monkeys
(Cercopithicus mitus stuhlmanni) (Ohsawa et al. 1993;
Cords 2000, 2002; Mugatha et al. 2006). Within-group
male coatis often spend several hours per day chasing
and fighting; these solitary males and adult males usually
lose weight and receive numerous injuries during the
mating season (Binzcik 2006; Hirsch 2007a). Because all
within-group adult female coatis are simultaneously in
oestrus, sequential mate guarding should be difficult or
impossible; thus, reproductive skew should be low (Cant
1998; Reeve et al. 1998; Nunn 1999). High reproductive
synchrony has been linked to low reproductive skew in
multi-male mating systems and to increase extra-pair
paternity in single-male mating systems (Stutchbury &
Morton 1995; Stutchbury 1998; Westneat & Stewart 2003;
Isvaran & Clutton-Brock 2007; Ostner et al. 2008). It has
been hypothesized that reproductive synchrony functions
to increase female mate choice and reduce the possibility
that males can monopolize female reproduction (Emlen
& Oring 1977; Stutchbury 1998; Ostner et al. 2008).
The social system of ring-tailed coatis differs with
respect to many studies of reproductive skew and
extra-pair paternity because coatis have one adult male
per group and several females. In many mammals, the
number of adult males in a group is highly correlated
with the number of adult females and their degree of
reproductive synchrony (Nunn 1999). Ring-tailed coati
groups in Iguazu, Argentina, contain one adult male
and 1?10 adult females year round; therefore, there is
no possible link between the number of adult males
and females in a group during most of the year. In this
respect, ring-tailed coatis resemble harem groups (Hec-
kel et al. 1999; Pemberton et al. 2002; Fabiani et al.
2004). To our knowledge, no examples of multiple adultthese extensively modelled systems, studies of other
species can be used to construct hypotheses concerning
reproductive success in ring-tailed coatis. The primary
goal of this paper is to test the extent to which three
nonmutually exclusive factors determine male repro-
ductive success in ring-tailed coatis.
1 Do social males have higher reproductive success
than solitary males?
If adult male coatis live in social groups year round
to increase access to mates during the mating season,
the number of offspring sired by social males should
be higher than solitary males.
2 Do high levels of reproductive synchrony lead to low
reproductive skew?
Reproductive synchrony should limit the degree to
which a group living male can monopolize mating.
Judging by comparable studies of other mammal spe-
cies, the ring-tailed coatis? short mating season should
prevent group males from siring most of their group?s
young (Isvaran & Clutton-Brock 2007; Ostner et al.
2008). To determine predicted values of within-group
paternity in ring-tailed coatis, we looked at previous
studies of mammal species that live in one-male or
harem groups and exhibit short mating seasons
(<2 months). Species that formed temporary harem
breeding groups had between 40% and 75% within-
group paternity (Le Boeuf and Reiter 1988, Hoelzel
et al. 1999, Fabiani et al. 2004, JM Pemberton et al.
unpublished data cited in Isvaran & Clutton-Brock
2007). In species that form more permanent social
groups, within-group males sired between 30 and 50%
of offspring (Heckel et al. 1999, Heckel et al. 2003,
Ohsawa et al. 2003, Dechmann et al. 2005, Hatcher
2007). We therefore predict that the resident male
should sire roughly 30?50% of its group?s offspring.
3 Does the ratio of males to group living females influ-
ence reproductive skew?
As the number of females in a coati group increases,
the number of satellite adult males following the group
during the mating season should increase, and the abil-
ity of the lone within-group male to defend females
and monopolize matings is predicted to decline.
Methods
The study was conducted in Iguazu National Park,
Argentina (54W, 26S), between July 2002 and Decem-season have been reported. When group living males
encounter solitary males, they usually fight them off;
thus, coati groups may be limited to one adult male
because of male?male aggression. Even though coatis 2010 Blackwell Publishing Ltd
ber 2004. A total of 150 coatis were captured in groups and were never observed residing in their natal
FA MILIARITY B REEDS PROGENY 41132 ? 10 ? 12 inch Tomahawk or similar traps, immobi-
lized with Ketamine and Xylazine and fitted with
unique combinations of multicoloured ear tags for indi-
vidual identification (Rototag ear tags, Dalton Co.). A
small plug of skin tissue was punched out during ear
tagging, and the tissue was stored in 10% DMSO saline
solution. Samples were kept at room temperature
between the date of capture and January 2005. The sam-
ples were stored in a )80 C freezer from January 2005
until DNA extraction in August 2007.
Genetic sampling focused on four habituated social
groups with overlapping home ranges (Hirsch 2007a).
Demographic data and group censuses were typically
taken at least once a month per group between June
2002 and December 2004. In this population, group
sizes ranged from 8 to 65 individuals and group per
years included in the paternity analyses ranged from 8
to 54; PQ 2002 = 8, 2003 = 15, GR ?PSG 2002 = 54, PSG
2003 = 12, 2004 = 29, SF 2002 = 25 (Hirsch 2007a,b). In
Iguazu, pregnant coatis gave birth to an average of
approximately 4.5 offspring per year (range 2?7; Hirsch
2007a). All adult females in the study groups were sex-
ually receptive during the mating season and appeared
pregnant before the groups disbanded in the nesting
season. Differences between the number of mated
females and number of females with offspring were
typically because of 100% mortality within a litter or
the death of the mother.
During some years, it was possible to trap and sam-
ple every individual in a group, while in other years
extensive sampling of some groups was not possible.
All adult females and living offspring were captured in
a total of five group years (PQ 2002, 2003, PSG 2002,
2003, SF 2002). The juveniles present in these groups
were all born in October, but were typically not trapped
until January to March of the following year (at 4?
5 months of age); thus, some of the offspring died
before we were able to sample them. Late in 2002, five
adult females and their juvenile offspring split off from
the GR group and formed a new group (PSG group).
The 2002 PSG juveniles were sired while the mothers
were still part of the larger GR group. During 2004,
only 9 of 31 juveniles from the PSG group were cap-
tured and sampled.
Adult males, adult females and offspring were sam-
pled in a total of five group mating seasons (PQ 2003,
PSG 2002, 2003, 2004, SF 2002). Some males that were
temporarily group members outside of the mating sea-
son were also sampled. In most cases, social adult
males were not related to any adult females in their
social group (B. T. Hirsch and J. E. Maldonado unpub-
lished data). Males that were captured and tagged in
their natal groups were observed entering neighbouring 2010 Blackwell Publishing Ltdgroup at adulthood (Hirsch 2007a). During the mating
season (early-mid August), additional satellite adult
males were observed following groups. The number of
satellite adult males per coati group during the mating
season was defined as the total number of adult males
seen within 15 m of the social group while adult
females were in oestrous. Because we were not able to
simultaneously monitor all social groups and it was dif-
ficult or impossible to distinguish between unmarked
solitary males, the number of males per group recorded
during the mating season was likely an underestimate.
We often discovered extra-group males when they
fought with other adult males; thus, the number of
observed males should have correlated closely with the
amount of male?male aggression and presumably with
the actual number of males attempting to mate. The
maximum number of recognizable males observed
within a group during the mating season was five
males (SF 2003). Solitary males were also seen outside
the mating season. Some of these solitary males were
trapped and sampled, while others were not. A total of
11 adult males classified as putative fathers were cap-
tured and sampled. In some cases (N = 4), males were
trapped as subadults (12?23 months old) in their natal
group and later observed to be members of another
group after they were fully mature males (typically
when 3 years of age or older).
DNA purification was carried out using a Qiagen Bio-
Sprint 96 workstation following the protocol for DNA
extraction from animal tissues as supplied by the manu-
facturer. All individuals were genotyped at 15 microsat-
ellite loci (Ma3, Davis & Strobeck 1998; Pfl2, Pfl8, Pfl9,
Kays et al. 2000; PLOT-01, PLOT-04, Fike et al. 2007;
PLM12, PLM13, Siripunkaw et al. 2007; F03, H03, E05,
H07, A08, F02, D03, Molecular Ecology Resources pri-
mer development consortium 2010). The polymerase
chain reaction mixtures (25 lL) were composed of
1.5 lL template DNA, 2.5 lL Gold PCR Buffer, 3 lL
MgCl2, 2.5 lL dNTP?s, 2 lL BSA, 2 lL Betaine, 1 lL
fluorescently labelled forward and reverse primers,
0.15 lL AmpliTaq Gold DNA polymerase (Applied Bio-
systems) and 9.35 lL H2O. In cases where reactions did
not yield a product, the reaction was repeated with lar-
ger quantities of DNA (2?3 lL per reaction). Because of
poor amplification, all PLM12 primer reactions con-
tained 3 lL of stock DNA. Primers specifically designed
for Nasua nasua (F03, H03, E05, H07, A08, F02, D03)
with the same annealing temperature were multiplexed
in the same PCR using 0.2?0.6 lL of each primer. PCRs
began with an extended denaturation of 96 C for
9 min, followed by 95 C denaturation cycle for 45 s, a
45 s annealing cycle and then a 72 C extension cycle
for 45 s (annealing temperatures in Table 1). The last
Table 1 The 15 microsatellite loci used to determine paternity. A total of 149 individuals were typed. Temp equals annealing tem-
pecte
lusio
Ho
0.55
0.15
0.49
0.58
0.41
0.68
0.71
0.66
0.42
0.51
0.04
0.15
0.71
0.56
0.58
412 B. T . HIRSCH and J . E . MALD ONADOthree 45 s steps were then repeated 34 times, followed
by a final 72 C extension cycle of 10 min. For PLM12,
we used a modified touchup PCR program. The initial
9-min 96 C denaturation cycle was followed by a 1-
min 95 C denaturation cycle, then a 1-min annealing
cycle, followed by a 72 C extension cycle for 1 min.
The annealing cycle started at 54 C, then was raised
one degree each step until the last step reached 57 C.
This cycle was repeated 15 times and then ended with
perature in the PCR. Hobs and Hexp represent observed and ex
berg equilibrium are indicated by asterisks. Locus-specific exc
column
Temp
No. of
alleles Size
Ma3 58 3 153?157
Pfl2 56 2 148?150
Pfl8 57 5 194?202
Pfl9 53 5 205?217
PLOT-01 64 3 155?159
PLOT-04 64 5 333?347
PLM12 54?57 7 217?229
PLM13 58 3 102?108
F03 60 5 109?119
H03 59 4 114?126
E05 60 2 148?150
H07 59 5 159?173
A08 59 7 213?225
F02 58 2 203?205
D03 58 5 259?269a final 72 C extension cycle of 10 min.
Products were electrophoresed through an ABI 3130xl
genetic analyzer (Applied Biosystems, Inc., Foster City,
CA.). Alleles were sized by comparison with concur-
rently run dye-labelled DNA size standards. Fragment
size analysis was performed using the GeneMapper
software (Applied Biosystems), and each genotype was
confirmed by visual inspection of the electrophero-
grams. All samples were amplified and genotyped at
least two times for each locus. Replicate genotyping
was carried out to minimize problems associated with
allelic dropout and misclassification of genotypes. If the
initial two genotypes derived from a sample did not
match, that sample was run two more times and the
genotype was determined using a consensus of all four
samples. We used the CERVUS 3.0 (Kalinowski et al.
2007) computer program to calculate whether the 15
loci conformed to Hardy?Weinberg equilibrium
(Table 1). Two alleles that were not in Hardy?Weinberg
equilibrium were included in the analysis because it
was determined that their inclusion would not lead to
an overestimation of within-group male paternity.Given that most alleles deviate from Hardy?Weinberg
because of allelic dropout or null alleles and that these
two alleles (Pfl2 and Pfl9) had null frequency values of
0.17 and 0.067, respectively, we believe there were no
major issues with including these alleles in the analyses
(Dakin & Avise 2004). In addition, there were no allele
mismatches at these two loci between known mother?
offspring pairs, which is evidence for a low frequency
of allelic dropout at these loci. Tests for linkage disequi-
d Heterozygosity. Significant deviations from the Hardy?Wein-
n probabilities for the second parent are reported in the last
He HW Fnull Exclusion
7 0.498 NS )0.058 0.806
4 0.218 ** 0.170 0.903
7 0.492 NS )0.026 0.723
4 0.663 ** 0.067 0.614
6 0.414 NS )0.005 0.833
5 0.698 NS )0.003 0.546
1 0.744 NS 0.023 0.500
4 0.559 NS )0.089 0.705
3 0.396 NS )0.038 0.777
0 0.520 NS 0.014 0.733
0 0.065 NS 0.213 0.969
4 0.145 NS )0.031 0.927
8 0.736 NS 0.017 0.309
4 0.501 NS )0.061 0.813
4 0.587 NS 0.003 0.655librium were implemented in GENEPOP 4.0.10, and no
statistically significant evidence for linkage disequilib-
rium between pairs of loci was found.
We used CERVUS 3.0 to determine paternity and mater-
nity assignments of 76 juvenile coatis. Maternity was
assigned using a total evidence approach, combining
behavioural data (grooming rates between mothers and
potential offspring) and genotyping (Prodohl et al.
1998; Slate et al. 2000). In the three cases where geneti-
cally assigned maternity did not match the mother pre-
dicted by grooming data, we examined LOD scores
(combined likelihood ratios of parental assignment) and
allele mismatches between the offspring and the pre-
dicted mother. In all three cases, the mother predicted
using grooming data had a positive LOD score and no
allele mismatches; so, we used the behavioural data to
determine maternity in these cases, and not the mother
assigned by CERVUS.
Paternity was calculated using trio LOD scores with
known mothers. To simulate the expected probability of
correctly determining the father at random from the Ig-
uazu population, it was assumed that half of the adult
 2010 Blackwell Publishing Ltd
coatis in the population were sampled. This estimate
was derived from the literature on adult sex ratios in
Nasua narica, where an average of 5.63 males per social
group has been reported (range 2.5?14)(Kaufman 1962;
Russell 1979, Gompper et al. 1997; Hass 2002; Hass &
Valenzuela 2002; Booth-Binczik et al. 2004; McColgin
2006). Given that a total of four groups and 11 adult
males were captured and sampled in Iguazu, a 50%
adult male capture rate was determined to be a reason-
able estimate. The combined exclusion probability for
the second parent was 0.993. Both relaxed and strict trio
LOD estimates were calculated based on a simulation
of 10 000 offspring with 15 loci and 2.75% mistyped
loci (80% critical trio LOD = 0.00, strict 95% critical trio
LOD = 2.12). The percentage of mistyped loci entered
into the model was based on the observed frequency
derived from CERVUS. In cases where the within-group
male was known and no father was assigned, it was
tiple paternity litters as the dependent variable. Because
spring: AK, AE, AM, CU, and c10). If offspring were
sired by adult males that were never sampled, using
only assigned offspring would lead to an overestimate
of within-group paternity (40 of 42 = 95.2% within-
group male paternity). If all offspring who were not
assigned to the within-group male using the 80% or
95% probability are classified as extra-group offspring,
the percentage of within-group paternity ranged
between 66.7% and 91.3% using 80% confidence and
44.4?66.7% with 95% confidence. We regard the 95%
confidence results as the absolute minimum level of
within-group paternity. Because males assigned at the
80% level but not at the 95% level often had zero allele
mismatches between father and offspring (N = 5) and
had at most one mismatch, we regard the 80% criteria
as the most robust measure of paternity for this study.
The proportion of within-group paternity (using the
80% assignments) was not correlated with the number
ales
ing s
r gro
the
es
FA MILIARITY B REEDS PROGENY 413the dependent variables were percentages, we logodds
transformed the data before running the analyses to
better conform to assumptions of normality.
Results
Likelihood analyses resulted in paternity assignments
for 42 of the 74 typed offspring with 95% confidence
(56.8% assignment) and 59 of 74 at 80% confidence
(79.7% assignment)(Table 2). Of these assigned off-
spring, five were assigned to an extra-group male (off-
Table 2 The percentage of offspring fathered by within-group m
sents the number of adult females in the group during the mat
females whose offspring survived the nesting season. Males pe
group males observed within 15 m of the social group during
sampled for DNA were not included in the table
Group Year # Mated $ # Moms # Mal
PQ 2003 5 5 1
PSG 2002 9 2 2
PSG 2003 5 3 1
PSG 2004 7 5 3
SF 2002 ?6 6 2
Totalassumed that the offspring was sired by an extra-group
male. Original least square (OLS) regressions were cal-
culated to determine whether the percentage of within-
group paternity per year was related to the number of
adult females, number of pregnant females, number of
males and male ? female ratio (JMP 5.1.2; SAS Institute).
We also ran these analyses with the percentage of mul- 2010 Blackwell Publishing Ltdof adult females or adult females with living offspring
from that group year (OLS regression: number of
females; F1,5 = 0.215, slope = )0.096, P = 0.675, number
of mothers; slope = )0.073, F1,5 = 0.118, P = 0.754).
Within-group paternity was also not significantly corre-
lated with the number of males observed with the
group during the mating season or the ratio of observed
males to within-group females, but the effect slopes
were in the predicted negative direction in both cases
(OLS regression: number of males; slope = )0.577,
F1,5 = 4.650. P = 0.120, male-female ratio; slope = )5.067,
F1,5 = 0.537, P = 0.083).
Multiple paternity was found in at least 9 of 17 litters
(53%). During some group years, juveniles were
trapped several months after the birth season and sev-
eral juveniles probably died before being sampled
(details in: Hirsch 2007a). If paternity analyses are
restricted to 2003, when almost all of the litters were
completely sampled, four of eight litters had more than
versus extra-group males. The number of mated females repre-
eason that could have mated. The number of mothers includes
up were calculated based on the number of social and extra-
mating season. Juveniles that died before being captured and
# Offspring
# Fathered
within-group
% Within-group
paternity
95% 80% 95% 80%
23 13 21 0.565 0.913
6 4 5 0.667 0.833
15 9 12 0.600 0.800
9 4 6 0.444 0.667
12 9 9 0.750 0.750
65 39 55 0.600 0.815
Table 3 Paternity assignments for six coati group years. Year indicates the year of birth of the offspring
Offspring Mother
Pair
AM
Pair
LOD
Candidate
father
Group
male
Pair
AM
Pair
LOD
Trio
AM
Trio
LOD
Trio
D LOD
Within-group
male
PQ 2002
AA AY 2 )3.961 VI ? 1 )1.245 4 )2.967 5.135 ?
AK AY 0 1.661 VI ? 1 )0.444 1 1.810 11.644 No
LW AY 0 2.617 VI ? 1 )0.823 2 )2.981 5.801 ?
SB AY 1 )0.721 OB1 ? 2 )3.882 3 )3.470 0.417 ?
PB GZ 1 )0.843 VI ? 1 )2.090 3 )3.420 5.287 ?
PU GZ 0 6.070 TV ? 1 )2.494 2 )2.082 1.089 ?
CC MA 0 7.535 TV ? 2 )6.636 2 )4.448 0.112 ?
CL MA 0 7.736 VI ? 1 )0.706 1 )2.251 5.369 ?
TC MA 0 3.590 EH ? 3 )6.234 4 )8.604 0.406 ?
PQ 2003
AD (AN) 0 0.343 OB1 OB1 0 3.876 0 4.264 9.378 ***
JK (AN) 0 1.402 OB1 OB1 0 4.602 0 6.367 13.518 ***
OV (AN) 0 )0.840 OB1 OB1 0 2.676 1 0.645 1.177 *
RR (AN) 0 )0.430 OB1 OB1 0 4.049 1 4.077 11.118 ***
DI AY 0 2.163 OB1 OB1 0 2.388 2 0.125 2.118 *
GD AY 0 3.290 OB1 OB1 0 2.715 0 5.644 10.815 ***
RQ AY 0 1.976 MD OB1 2 )6.194 2 )2.927 1.886 No
SV AY 0 4.661 OB1 OB1 0 0.677 1 0.724 3.219 *
ED DA 0 2.428 OB1 OB1 0 0.275 0 4.829 9.674 ***
OZ DA 0 2.172 OB1 OB1 0 1.551 1 4.218 10.894 ***
RY DA 0 1.512 OB1 OB1 1 )1.652 2 0.191 8.705 *
SZ DA 0 4.969 OB1 OB1 0 0.275 0 4.876 9.725 ***
AL GZ 0 )0.283 OB1 OB1 1 )0.032 2 0.822 13.843 *
JF GZ 0 4.483 OB1 OB1 0 )0.282 1 0.689 5.162 *
LO GZ 0 )0.226 OB1 OB1 0 1.383 1 4.534 14.645 ***
MM GZ 0 1.864 OB1 OB1 0 1.716 1 4.164 11.901 ***
OG GZ 0 0.341 OB1 OB1 0 2.898 1 5.234 15.489 ***
RX GZ 0 3.108 OB1 OB1 0 1.383 0 6.757 14.460 ***
SN GZ 0 3.383 OB1 OB1 0 2.222 1 4.307 7.734 ***
AE MA 0 5.022 DaMale OB1 1 )1.745 1 2.242 3.646 No
GL MA 0 4.937 OB1 OB1 1 )2.455 1 0.686 5.867 *
TL MA 0 5.507 OB1 OB1 0 1.324 0 4.738 11.138 ***
VL MA 0 3.586 OB1 OB1 0 3.479 0 7.313 12.640 ***
PSG 2002
JS GH 0 4.228 VI VI 0 2.172 0 5.228 13.062 ***
SX GH 0 3.369 VI VI 2 )4.760 2 )2.350 2.562 No
TM GH 0 3.067 VI VI 0 1.593 0 4.482 12.925 ***
BS PS 0 0.879 VI VI 0 1.902 0 5.779 12.097 ***
DM PS 0 2.413 VI VI 0 1.848 0 5.239 10.599 ***
KG PS 0 2.713 VI VI 1 )1.660 1 1.113 11.009 *
PSG 2003
AS CM 0 5.447 VI VI 0 3.045 0 3.833 12.355 ***
CV CM 1 0.396 VI VI 1 )0.557 1 3.424 11.121 ***
IB CM 0 4.313 VI VI 1 1.461 1 2.552 0.243 ***
KH CM 0 4.880 VI VI 0 0.621 1 1.155 0.443 *
ZS CM 0 3.674 VI VI 0 3.852 0 6.373 16.337 ***
AM GH 0 7.093 VV VI 1 )0.022 1 1.211 5.739 No
BJ GH 0 2.818 VI VI 0 0.140 1 2.910 12.320 ***
BO GH 0 2.718 VI VI 0 2.245 0 5.570 13.461 ***
DH GH 0 3.382 VI VI 0 2.773 0 5.938 16.409 ***
ES GH 0 4.319 VI VI 1 )2.761 1 0.599 3.849 *
BK NY 0 1.483 VI VI 1 )0.752 1 0.497 3.226 *
BM NY 1 1.847 VI VI 1 )0.638 2 )0.476 0.272 No
IP NY 0 1.245 OB1 VI 1 0.345 2 )1.866 4.030 No
414 B. T . HIRSCH and J . E . MALD ONADO
 2010 Blackwell Publishing Ltd
Table 3 (Continued)
0 1.955 0 2.464 3.790 ***
c10 JW 0 6.033 BF VI
FA MILIARITY B REEDS PROGENY 415c2 JW 0 6.416 VI VI
c5 PS 0 1.532 VI VI
c6 PS 0 2.395 MD VI
c7 PS 0 0.792 VI VI
c3 SX 0 3.160 VI VI
c8 SX 0 1.519 VI VI
SF 2002
KK BR 0 3.629 WW WW
MH BR 0 0.956 WW WW
CU EK 0 3.889 IK WW
LS EK 1 0.222 MD WW
BT LD 0 2.317 WW WW
RW MN 0 4.494 MD WW
SR MN 1 1.105 WW WW
JL MS 0 0.377 WW WW
MG SL 0 2.071 WW WW
PK SL 0 5.579 WW WW
TR SL 0 2.072 WW WWOffspring Mother
Pair
AM
Pair
LOD
Candidate
father
Group
male
RO NY 1 1.830 VI VI
RS NY 0 3.811 VI VI
PSG 2004
c1 BS 0 2.907 VI VI
c9 CM 0 2.521 VI VIone father. In seven cases where multiple paternity was
detected, only one offspring in the litter was fathered
by a solitary male (Table 3). In one case, two of five off-
spring in a litter were sired by extra-group male(s)
(female NY during 2003). We were not able to deter-
mine whether the same solitary male fathered more
than one offspring in the same group or any other
group during the same mating season. This might have
occurred, but this was not possible to confirm without a
complete sampling of potential fathers. There were no
significant correlations between group size, the number
of females or the male ? female ratio on the percentage
of multiple litters in a group (all P values 0.888). The
relatively high levels of multiple paternity are evidence
that most females were mating with multiple males.
Discussion
Do social males have higher reproductive success than
solitary males?
Groups of ring-tailed coatis generally had low levels of
extra-group paternity, even though many litters had
one offspring fathered by an extra-group male. Within-
ZO SL 1 2.694 WW WW
AM, number of allelic mismatches. Group males were observed to be
***Strict 95% confidence, *relaxed 80% confidence. ? indicates unknow
 2010 Blackwell Publishing Ltd0 5.212 0 7.920 9.588 No
0 2.459 0 5.435 16.184 ***
1 )2.757 1 0.189 2.369 *
2 )4.266 2 )3.388 0.178 No
0 1.845 0 5.369 10.813 ***
2 )4.157 2 )3.027 3.990 No
1 )0.918 1 0.473 0.700 *
0 4.046 1 3.928 7.482 ***
0 4.931 1 4.726 7.106 ***
1 )1.813 2 0.181 3.992 No
2 )4.130 3 )3.782 1.652 No
0 6.930 1 5.613 12.078 ***
2 )2.194 4 )7.480 0.492 No
0 3.691 2 3.600 6.641 ***
0 6.579 2 5.005 14.791 ***
0 6.696 1 6.372 8.858 ***
0 2.333 0 4.361 2.352 ***
0 4.457 1 4.166 7.664 ***Pair
AM
Pair
LOD
Trio
AM
Trio
LOD
Trio
D LOD
Within-group
male
0 0.218 1 2.771 8.009 ***
0 0.894 0 2.772 1.669 ***
0 3.917 0 2.301 1.037 ***group paternity was between 66.7% and 91.3% per
group ?year using the relaxed 80% probability. No soli-
tary male sampled in this study came close to
approaching the reproductive success of the social
males. All five offspring assigned to extra-group males
at the 80% confidence level were assigned to different
fathers (VI, DaMale, VV, IK and BF) and there was no
evidence that any one extra-group male sired large
numbers of offspring (Table 3). Because of the high
degree of breeding synchrony between groups at the
study site, it would be difficult or impossible for a soli-
tary male to mate with enough females from different
groups to achieve similar or greater mating success than
social males. This result is strong evidence that social
males have higher reproductive success than solitary
males.
Do high levels of reproductive synchrony lead to low
reproductive skew?
Levels of mating success for the social males were
higher than predicted for a species with high reproduc-
tive synchrony and ample opportunities for extra-group
copulations (we predicted values ?50%). Two recent
0 2.647 2 2.207 4.538 ***
incorporated into the social group during the mating season.
n individual.
studies using phylogenetic comparisons found that
dominant or within-group males must defend more
females engaged in extra-group copulations, suggesting
average for carnivores (Binzcik 2006; Iossa et al. 2008).
416 B. T . HIRSCH and J . E . MALD ONADOthat the resident male could not prevent such matings.
Yet these matings did not lead to high levels of extra-
group paternity: did adult females choose to mate with
resident males when they were most likely to conceive?
Mechanisms leading to high mating success in social
male coatis
The mechanisms that led to high reproductive success
in social male coatis are not clear. Social males may try
to outcompete solitary males using sperm competition,
which should result in coatis having large testes com-
pared to other mammals (Parker et al. 1997; Soulsbury
2010). Even after considering that male coatis exhibit
larger testicular volumes during the mating season, the
relative testicular volume in coatis is not larger thanthan one female at the same time (but see Wimmer &
Kappeler 2002). Current models of reproductive skew
cannot explain the reproductive success of group living
male ring-tailed coatis.
Does the number females and males influence
reproductive skew?
An increase in the number of females that are simulta-
neously in oestrus should lead to a decrease in male
monopolization and reproductive skew (Nunn 1999;
Isvaran & Clutton-Brock 2007, Ostner et al. 2008). A
trend was found that within-group paternity decreased
as the number of male to female coatis increased, but
this was not statistically significant, which was likely
because of the low sample size (n = 5 group years).
During the mating season, adult males often engaged in
vicious fights for access to adult females. At the end of
the mating season, adult males typically had noticeable
weight loss and severe cuts and wounds on their body.
Even though within-group males sired the majority of
offspring, multiple paternity was found in a minimum
of 50% of litters. These litters, however, normally con-
tained only one offspring of an extra-group male. Manyreproductive skew was lower and extra-group paternity
higher in populations with greater reproductive syn-
chrony (Isvaran & Clutton-Brock 2007, Ostner et al.
2008). This relationship is consistent with incomplete
control models of reproductive skew which predict that
a main factor driving reproductive skew is the ability of
dominants to monopolize access to females and prevent
other males from mating with them (Altmann 1962;
Cant 1998; Clutton-Brock 1998; Reeve et al. 1998). Short
reproductive seasons generally lead to a high degree of
overlap in female oestrous periods, which means thatIn general, coati copulations were not brief and pairs
were observed to mate for more than 55 min, which is
similar to Nasua narica (Hass & Roback 2000). This
behaviour could be related to induced ovulation,
although it is not known if coatis are induced or spon-
taneous ovulators (Hass & Roback 2000; Lariviere &
Ferguson 2002, Lucero et al. 2007; Iossa et al. 2008).
These lengthy copulatory bouts likely make sneaky
mating more difficult and increase the probability of
mating interruption. If within-group males are able to
mate with females at will, they should have a lower
risk of being interrupted and can copulate with females
for the sufficient length of time needed to induce ovula-
tion. Alternatively, extra-group males that need to fight
the social male for access to females may not be able to
mate at will. This hypothesis is complicated by obser-
vations that females often leave their group to mate
with extra-group males, which has also been observed
in N. narica (Hass & Roback 2000; Booth-Binczik et al.
2004). By leaving their group, females are able to mate
with extra-group males and lower their risk of being
interrupted. If ring-tailed coatis are spontaneous ovula-
tors, within-group males may be able to determine the
time of ovulation from olfactory cues. Because within-
group males have more contact with adult females, it is
possible that they have better knowledge about the
ideal time for reproduction and are able to outcompete
extra-group males using these cues. Alternatively,
females may be able to choose when they mate with
various males and the high reproductive success of
social males could result largely from female mate
choice.
There are strong reproductive benefits to sociality in
male ring-tailed coatis, although the exact mechanisms
for this are unclear. It is also uncertain why adult males
are found in groups year round. Presumably, adult
males can only enter coati groups if females allow it.
When the mating season starts, adult females may have
already chosen a highly desired adult male to enter
their group. The exact traits that are desired or selected
for by adult females are unclear, but it appeared that
within-group males were larger and better fighters than
extra-group males. Because adult males typically fight
when they meet, social males may have their competi-
tive ability tested several times a year outside the mat-
ing season. If resident social males are chiefly
responsible for excluding other adult males from social
groups, this aggression would limit the number of
social males in the population. It is notable that no coati
groups have been observed with more than one adult
male simultaneously living in the group outside the
mating season. It seems plausible that adult male broth-
ers or other close kin could form a coalition to enter 2010 Blackwell Publishing Ltd
and remain in a social group, but this has never been
of ovulation. Social males may be more knowledgeable
Lori Eggert and Jenny Fike for primer aliquots. Frank Hailer,
Binzcik GA (2006) Reproductive Biology of a Tropical Proyconid,
FA MILIARITY B REEDS PROGENY 417Emily Latch and Nancy Rotzel provided BTH valuable advice
on primer optimization and laboratory techniques. Mirian
Tsuchiya-Jerep graciously provided us with details for the
primers she developed and without which this study could
not have been done. We are particularly thankful to Mirian
and Eduardo Eizirik. BTH thank Charles Janson for his
advice and support during the course of the research. This
paper benefited tremendously thanks to comments on earlier
drafts by Egbert Leigh Jr., Roland Kays, Christie Riehl, Brandt
Ryder, Jon Slate and two anonymous reviewers. This study
complied with all institutional, national and ASAB ? ABS
guidelines for animal welfare. This study was funded by an
NSF doctoral dissertation improvement grant (BCS-0314525),
the Smithsonian Institution postdoctoral fellowship pro-
gramme, the Smithsonian Undersecretary for Science
restricted endowment funds, and the Smithsonian Conserva-
tion Biology Institute.
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B.T.H. is a behavioural ecologist who uses mammalian study
systems to address questions related to the evolution of social-
ity, group formation and geometry, predation, competition,
and the ecology of tropical forests. This fieldwork was con-
ducted as part of B.T.H.?s Ph.D. project and the genetic ana-
lyses were conducted at the Smithsonian Center for
Conservation and Evolutionary Genetics during a post-doctoral
fellowship. J.E.M. is a research geneticist interested in using
molecular genetic tools to answer basic and applied questions
dealing with conservation genetics, systematics, behavioural
ecology and the evolution and genetic diversity of a variety of
mammals.
FA MILIARITY B REEDS PROGENY 419 2010 Blackwell Publishing Ltd