Folia Primatologica 93 (2022) 97-105 brill.

com/ijfp

Brief Report

A case of polygyny in the Bornean white-bearded

gibbon (Hylobates albibarbis)
Carolyn Thompsona,b,c,∗ , Eka Cahyaningruma , Hélène Birota , Abdul Aziza,d and
Susan M. Cheynea,e
a
The Borneo Nature Foundation (BNF), Jalan Bukit Raya, Palangka Raya, Central Kalimantan,
74874, Indonesia
b
Genetics Evolution and Environment, University College London, Gower St, Bloomsbury, London
WC1E 6BT, UK
c
Institute of Zoology, Zoological Society of London, Regent’s Park, London, NW1 4RY, UK
d
Centre for International Cooperation in the Management of Tropical Peatlands (CIMTROP),
University of Palangka Raya, Palangka Raya, Central Kalimantan, 74874, Indonesia
e
Faculty of Humanities, Social Sciences and Law, Oxford Brookes University, Headington Road,
Headington, Oxford, OX3 0BP, UK
* Corresponding author; e-mail: carolyn.thompson.17@ucl.ac.uk
Received 8 October 2020; accepted 27 November 2021; published online 1 March 2022;
published in print 12 May 2022

Abstract – Gibbons (family Hylobatidae) typically form groups that encompass a single breeding pair. Here,
we present the first evidence of polygyny (where a single male has more than one female mate) in the Bornean
white-bearded gibbon (Hylobates albibarbis). In July 2014, an adult female yet to have emigrated from her
natal group gave birth to an infant, bringing the total group size to six individuals (one adult male, two adult
females, one subadult female, and two infant females). Forty months later in November 2017, the same female
gave birth to a second infant. Between July 2014 and April 2018, the two breeding females within the group
remained mutually tolerant of each other, often singing the characteristic female vocalisation, the great call,
in unison, until the eldest adult female dispersed in November 2018. We explore possible reasons behind this
group’s mating system flexibility by examining dispersal limitation due to environmental constraints, factors
associated with a large home range size, mutual tolerance between females, and a lack of mating opportunities.
Keywords – extra-pair copulations, Indonesia, mating system, monogamy, multi-female group, social system.

Introduction between conspecifics), social organisation (i.e.,

Primate social systems vary between species,
within-species and even on a population level
(Fleagle, 2013; Koenig et al., 2013). These
variations are the result of specific ecological
characteristics relating to access to resources
(e.g., food and mates) and potential risks (e.g.,
predators) (Koenig et al., 2013). Different social
systems are defined by looking at associated
components, such as social structure (i.e., dis-
persal patterns and quality of relationships
© CAROLYN THOMPSON ET AL., 2022
This is an open access article distributed under the terms of the CC BY 4.0 license.
size, composition and cohesion), and mating
system (i.e., how individuals mate and breed
within their social system) (Dunbar, 1988; Kap-
peler and van Schaik, 2002; Kappeler, 2019).
Four main mating systems have been identi-
fied in primate species: monogamy (exclusive
mating relationship between one male and one
female until disrupted, e.g., death, intruder, part-
ner disperses etc.), polygyny (a single male
has more than one female mate), polyandry (a
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DOI 10.1163/14219980-20200801
via free access
C. Thompson et al.
single female has more than one male mate)
and polygynandry (both males and females
have multiple mates) (Strier, 2016; Huck et al.,
2020). It is important to note that mating sys-
tems can be genetic (i.e., related to breeding,
where an infant is conceived) or social (i.e.,
related to mating, which does not necessarily
lead to conception). Terms associated with mat-
ing systems are often used inconsistently in the
literature (Kappeler, 2019; Huck et al., 2020).
For clarity, when discussing different “mating
systems” in this paper, we are referring to mat-
ing and reproductive behaviour and not group-
ing patterns.
Gibbons (family Hylobatidae) are small arbo-
real apes found in east and southeast Asia.
They typically live in small family units con-
sisting of a single breeding pair and their off-
spring (O’Brien and Kinnaird, 2011). Both
sexes of offspring usually emigrate from their
natal group, often being pushed out by the
same-sex parent once they have reached sex-
ual maturity around eight years (Brockelman et
al., 1998; Bartlett, 2011; Kappeler et al., 2013;
Guan et al., 2013). Emigrating offspring move
to a new home range to mate with a lone individ-
ual or displace a previously-mated individual in
a “takeover” event (Mackinnon and Mackinnon,
1977; Brockelman et al., 1998). By dispersing,
individuals avoid the risk of inbreeding (Wolff,
1993; Perrin and Mazalov, 2000), reduce local
resource competition (Greenwood, 1980) and
intra-group competition between same-sexed
individuals (Dobson, 1982; Moore and Ali,
1984).
Monogamy is evident in only 14% of pri-
mates (Rutberg, 1983; Clutton-Brock, 2016)
and can have associated benefits such as extra
paternal investment (Borries et al., 2011). Early
research suggested gibbons were strictly monog-
amous (Carpenter, 1940; Chivers, 1974;
Srikosamatara, 1980). However, long-term stud-
ies have reported partner changes and extra-pair
copulation events (EPCs) (a mating behaviour
where an individual copulates furtively with
a group outsider) (e.g., white-handed gibbon,
Hylobates lar: Sommer and Reichard, 2000;
Reichard, 2009), as well as polygynous mating
systems (pileated gibbons, Hylobates pileatus:
98
Srikosamatara and Brockelman, 1987; black
crested gibbon, Nomascus concolor: Fan et
al., 2006, Huang et al., 2013; Hainan gibbon,
Nomascus hainanus: Liu et al, 1989; Zhou et al.,
2005; 2008; Cao vit gibbon, Nomascus nasu-
tus: Fan et al., 2010) and polyandrous mating
systems (H. lar: Barelli et al., 2008; Reichard
et al., 2012; siamang, Symphalangus syndacty-
lus: Lappan, 2007; western hoolock, Hoolock
hoolock: Siddiqi, 1986; Ahsan, 1995).
Observations of polygyny in gibbons were
first published in 1987 (H. pileatus: Skrikosa-
matara and Brockelman, 1987). Of the four gen-
era of gibbons, Nomascus species have the high-
est incidence of polygyny (Malone and Fuentes,
2009; N. nasutus: Fan et al. 2010; N. concolor:
Fan and Jiang, 2009; N. hainanus: Zhou et al.,
2005, 2008). Interestingly, some species of gib-
bon have flexibly adopted both monogamous
and polygamous (i.e., polygyny or polyandry)
mating systems, possibly as a response to exter-
nal ecological and social factors (e.g., food and
mate availability) (Kappeler et al., 2013; Som-
mer and Reichard, 2000). Polygyny has been
hypothesised to be caused by either a lack of
available habitat to disperse into upon reaching
sexual maturity (Liu et al., 1989; Huang et al.,
2013), larger home ranges requiring more indi-
viduals to defend (Rutberg, 1983; Jiang et al.,
1999; Fan et al., 2015), and/or a mutual toler-
ance between sexually mature females within
the same group (Srikosamatara and Brockel-
man, 1987; Liu et al., 1989; Jiang et al., 1999).
Polygyny in many gibbon groups has been
reported as being short-lived (i.e., lasting less
than two years) (Fuentes, 1998; Fan et al.,
2010). Gibbon groups exhibiting this mating
system for longer have only been documented
in Nomascus species (Zhou et al., 2005; Fan and
Jiang, 2009; Fan et al., 2010).
This brief report presents the first evidence
of polygyny recorded in Bornean white-bearded
gibbons (Hylobates albibarbis). With current,
long-term gibbon field studies running beyond
15 years, more reliable observations of group
dynamics can now be made. Herein we give
possible explanations for this rare occurrence
observed in this species.
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Folia Primatologica
 A case of polygyny in the Bornean white-bearded gibbon
Materials and methods
STUDY LOCATION
These observations were documented by the
Borneo Nature Foundation (BNF) in the Natu-
ral Laboratory of Peat Swamp Forest (NLPSF),
Central Kalimantan, Indonesia (2°19 S and
113°54 E). The NLPSF is located 20 km south-
west of the provincial capital, Palangka Raya,
and is operated by the Centre for the Interna-
tional Cooperation in the Management of Trop-
ical Peatlands (CIMTROP).
STUDY SPECIES
H. albibarbis is endemic to south-west Kali-
mantan between the Kapuas and Barito Rivers
(Buckley et al., 2006). The species is found in
primary and disturbed secondary tropical forest,
as well as montane and lowland habitats, includ-
ing peat-swamp forest (Nijman et al., 2008).
H. albibarbis is currently listed as Endangered
on the International Union for Conservation of
Nature’s Red List of Threatened Species due
to illegal trade, hunting and deforestation (Nij-
man et al., 2008; Cheyne et al., 2019). Previous
research suggests this species displays a monog-
amous mating system (Gittins and Raemaek-
ers, 1980; Cheyne and Chivers, 2006) with no
records of polygamy to date. The average group
size is 4.45 individuals (range: 3-5 individu-
als) (Cheyne et al., 2008; Cheyne, 2010). The
inter-birth interval is 2.4 years and the gestation
period lasts for ca. 210-270 days (7-9 months)
(Cheyne, 2010).
BEHAVIOUR MONITORING
Since 2005, BNF has been monitoring four
habituated H. albibarbis groups with overlap-
ping home ranges. More than 4449 h (1898
follow days on 44 individuals) have been col-
lected (usually between 04:00 and 14:00). Of
the groups studied, 2065 h (608 days) were
spent observing group K between May 2005
and July 2019. The group was followed for
up to eight days every month. We monitored
group composition (including dispersal pat-
terns, births and deaths) and copulation events,
and recorded these observations ad libitum.
Vol. 93(1), 2022
Individuals were identified through detailed
physical cues (e.g., eyebrows, beards, develop-
ment of genitalia, body size) and behavioural
criteria (e.g., singing, foraging dynamics, rela-
tive proximity to other group members). Indi-
vidual identification guides (with photographs)
were created to aid field researchers.
Results
Group K has been observed since 2005,
enabling us to monitor life histories, changes in
demographics, dispersal events, and the death
and arrival of new infants. In 2005, group K
consisted of an adult mated pair (a female called
CL and a male called BL), the formation of
which was before the study period, and two
female offspring (ZZ and JL) (fig. 1). We esti-
mated that ZZ was born in 2000 and dispersed
from the group in 2011. JL was estimated to
be born in 2003 and remained within her natal
group despite being more than 10 years old.
Between 2005 and 2012, CL gave birth to three
other infants (KK, BRL and an unnamed infant).
KK and the unnamed infant died of unknown
causes in 2007 and 2012, respectively, whilst
BRL dispersed from her natal group in 2016.
In 2014, the adult females within group K,
CL and JL, each gave birth to a female infant;
CL to EL, and CRL to JL (figs 1 and 2). EL
was CL’s fourth infant, born in April 2014. CRL
was JL’s first infant, born in July 2014. There
were numerous copulations recorded between
CL and BL in the run-up to the births between
December 2012 and January 2014 (fig. 2). An
EPC involving CL was also recorded between
an adult male (CA) in neighbouring group C
in July 2013. No copulation attempts were
observed between JL and any outside males.
The arrival of the two infants, EL and CRL,
increased the family group size to six individ-
uals: one adult male, two adult females, one
subadult female and two infant females (fig. 1).
Between the births in 2014 and 2015, both adult
females duetted with BL, singing their great
calls in unison (fig. 2; see supplementary video
S1).
In May 2016, a copulation was observed
between JL and BL. In November 2017, JL gave
birth to her second infant female, JNL (figs 1
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99
C. Thompson et al.

Figure 1. Family tree of group K. Males are represented by a rectangle, females by an oval. Offspring that
dispersed or died during the study period are highlighted in grey. The dotted line indicates the unknown mate
of JL.

Figure 2. Timeline of events for group K between December 2012 and November 2018, highlighting
copulations, shared singing behaviour and births.

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100 Folia Primatologica
 A case of polygyny in the Bornean white-bearded gibbon
and 2). Between 2016 and 2017, CL appeared
to travel further away from the rest of her group,
but still joined the duet and great call with
BL and JL. In September 2018, there was an
aggressive encounter between BL and CL, when
he attacked her. The context was unknown.
No aggressive or displacement behaviours have
ever been recorded between CL and JL. On
25 November 2018, CL was observed travel-
ling farther away from the group before being
chased away by her own group. CL then disap-
peared and was not observed up until July 2019.
Based on anecdotal evidence since this publi-
cation, we believe that CL was forced out of
the group permanently while her youngest off-
spring, EL, remained.
Discussion/conclusion
Monogamy was thought to occur in about 14
to 15% of primates (Rutberg, 1983; Clutton-
Brock, 2016), but our definition of monogamy
has more recently been subject to reassessment,
owing to more detailed observations and the
availability of genetic information. With numer-
ous observations of partner changes and EPCs
occurring in many “monogamous” mammals
(Hughes, 1998) including gibbons (Palombit,
1994; Reichard, 1995; Barelli et al., 2013), the
traditional definition of “monogamy” has since
been challenged (Tecot et al., 2016). As evi-
dent in this study, an EPC involving the adult
female, CL, was observed during an intergroup
encounter. Although no EPCs were recorded
for JL, it is possible that these events occurred
but were not recorded due to the difficulty of
identifying individuals during brief intergroup
encounters, or occurred on non-follow days.
In the genus of Hylobates, cases of polyg-
yny are rare (Malone and Fuentes, 2009) and
have been short-lived (i.e., lasting less than two
years) (Fuentes, 1998; Sommer and Reichard,
2000). Even Nomascus species which demon-
strate a higher incidence of polygyny (Zhou et
al., 2005; Fan and Jiang, 2009; Fan et al., 2010),
have cases of mature subadult gibbons eventu-
ally replacing the adult male or female in their
natal groups (N. concolor: Huang et al., 2013).
Although we recorded no aggressive encoun-
ters between CL and JL, the departure of CL
Vol. 93(1), 2022
in November 2018 resulted in JL eventually
replacing her mother. Our long-term observa-
tional study of a H. albibarbis group verifies
that they exhibited a polygynous mating sys-
tem for approximately four years, making it the
longest observation in Hylobates. Reasons for
its occurrence are not fully understood, but we
explore several possible explanations below: (1)
lack of optimal quality habitat to disperse to; (2)
mutual tolerance between females and/or short-
age of mating opportunities; and (3) home range
size indicators.
LACK OF OPTIMAL QUALITY HABITAT
Polygyny in gibbons has previously believed
to have occurred in response to a lack of opti-
mal quality habitat for individuals to disperse
to and find mates (Liu et al., 1989; Huang
et al., 2013). Group K resided in a degraded
peat-swamp forest which, although protected
since 2004, has faced many years of distur-
bance from illegal logging and forest fires (Har-
rison et al., 2007; Cheyne et al., 2019). Fur-
thermore, the peat-swamp forest is recognised
as a low-productivity habitat (Morrogh-Bernard
et al., 2003) with important seasonal fluctua-
tions of fruit availability (Cheyne, 2010; Har-
rison et al., 2016). In 2015, forest fires devas-
tated 53.8 km2 of habitat in NLPSF (Cheyne et
al., 2019). Although this forest loss has reduced
available gibbon habitat in the region, group
K’s home range did not appear to be directly
impacted. In fact, group K maintained a large
home range of 154 ha (Cheyne et al., 2019).
Although the peat-swamp forest is considered to
be a low-productivity habitat, food is available
year round (Harrison et al., 2010). We, there-
fore, discount a lack of available habitat as a
possible enabler for the observed polygyny.
MUTUAL TOLERANCE AND MATING
OPPORTUNITIES
Although not necessarily a driver of polyg-
yny, mutual tolerance between breeding female
gibbons would support it, and also benefit group
cohesion when defending food resources and
reducing predation risk. Nevertheless, subadult
female gibbons approaching sexual maturity
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101
C. Thompson et al.
would not normally be tolerated by the adult
female within the same group (Jiang et al.,
1999). During a 20-month study on N. con-
color, no displacement or aggressive encounters
were recorded between two adult females how-
ever, suggesting that mutual tolerance allowed
more than one breeding female to reside in
the group (Jiang et al., 1999). Similar observa-
tions have been recorded in N. hainanus (Liu
et al., 1989) where the females fed amicably,
and in H. pileatus where the great call was
sung in unison (Srikosamatara and Brockelman,
1987). No bouts of aggression or displacement
were observed between JL and CL, and they
were recorded regularly singing the great call
together, often in proximity with BL, thus sug-
gesting a mutual tolerance between the adult
females.
Mutual tolerance alone does not explain
polygyny, however. Although dispersal patterns
of gibbons in NLPSF are still poorly under-
stood, there does appear to be a limited number
of available breeding males in the region (for
reasons unknown). Furthermore, other groups
that are followed regularly have subadult males
who are more than 12 years of age and yet to
emigrate from their natal group, indicating a
delayed dispersal.
HOME RANGE SIZE INDICATORS
Generally, home ranges across gibbon species
vary between 7 and 58 ha (Leighton, 1987).
Gibbon groups exhibiting polygynous mating
systems have been known to occupy up to
500 ha however, demonstrating a possible rela-
tionship between mating system, group size and
habitat area (N. hainanus: Liu et al., 1989).
Larger home ranges have the resources to sus-
tain two or more breeding females and their
offspring (Jiang et al., 1999; Fan et al., 2015).
Females have been speculated to benefit from
these larger home ranges by having greater
access to resources, therefore improving their
overall chances of reproductive success and fit-
ness (Guan et al., 2017).
Drivers behind group K’s large home range
size could be due to a lack of intergroup com-
petition in the NLPSF, and/or the “Resource
Dispersion Hypothesis” (where resources are
102
patchily distributed across a large area so a
group of animals will share the same space
with little to no feeding competition) (Macdon-
ald, 1983; Kruuk and Macdonald, 1985; Carr
and Macdonald, 1986). Whilst some groups in
NLPSF are limited by their home range abut-
ting the forest edge and therefore are unable
to expand, group K have few neighbours to
the west (possibly due to a population decline
in the early 2000’s associated with logging in
the region) (Cheyne, pers. obs.). The “Resource
Dispersion Hypothesis” can be discounted in
the NLPSF, however, as food is available
throughout the year (Harrison et al., 2010) and
the gibbons have little variation in their home
range use (Singh et al., 2018; Cheyne et al.,
2019).
In summary, we suggest that a lack of neigh-
bouring groups and competition for food and
space could have facilitated group K’s large
home range size. This, coupled with delayed
male dispersal and a female skew in the pop-
ulation, as well as intragroup female tolerance,
could have enabled the polygyny observed in
group K. Genetic testing would confirm pater-
nity of all offspring and help to explain some
of the intragroup relationships. It would also
improve our knowledge of gibbon dispersal pat-
terns, an area of research that is still understud-
ied.
In this brief report, we have not only high-
lighted the first recorded observation of polyg-
yny in this species, but also the value of long-
term monitoring to improve our understand-
ing of gibbon mating systems. The implica-
tions of a polygynous mating system in this
region (e.g., inbreeding and overcrowding) are
unknown and therefore there is huge benefit in
continued long-term monitoring. Furthermore,
with anthropogenic activities and habitat loss
increasing in this region, we may see more inter-
group interactions and conflicts, EPCs, and less
available optimal habitat for the formation of
new gibbon groups, thus impacting mating sys-
tems with unknown consequences.
Acknowledgements
We wish to thank RISTEK for supporting this
research, as well as our partners, CIMTROP,
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Folia Primatologica
 A case of polygyny in the Bornean white-bearded gibbon
and all the BNF staff, volunteers, students and
interns who have worked on the gibbon project
since 2005. We also thank Axel Martinez Ruiz
and Jenny Grib for their technical knowhow.
Finally, we wish to thank the reviewers for their
constructive comments and for strengthening
our manuscript.
Statement of ethics
This field study has no ethical conflicts to
disclose. The research was approved locally
by RISTEK (HB: 225/SIP/FRP/SM/VI/2013,
CT: 043/SIP/FRP/SM/II/2014 and SMC: 217-
A/SIP/FRP/SM/I/2014) and complies with the
Association for the Study of Animal Behaviour
guidelines for the treatment of animals in be-
havioural research.
Conflict of interest statement
The authors have no conflicts of interest to
declare.
Funding sources
Arcus Foundation; US Fish and Wildlife
Great Ape Fund.
Author contributions
All authors assisted with data collection in
Indonesia under SMC’s project management
and CT’s field supervision. As first and corre-
sponding author, CT led on the writing of the
manuscript. EC, HB and SMC contributed to
the writing and editing of the manuscript. AK
and EC prepared the data. CT, HB and SMC
prepared the figures. All authors edited the final
manuscript.
Supplementary material
Supplementary material is available online at:
https://doi.org/10.6084/m9.figshare.18709058
Vol. 93(1), 2022
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