How many gamergates is an ant queen worth? - ResearchGate

Naturwissenschaften (2008) 95:109–116
DOI 10.1007/s00114-007-0297-0
ORIGINAL PAPER
How many gamergates is an ant queen worth?
Thibaud Monnin & Christian Peeters
Received: 13 September 2006 /Revised: 14 May 2007 /Accepted: 2 August 2007 / Published online: 25 August 2007
# Springer-Verlag 2007
Abstract Ant reproductives exhibit different morphological
adaptations linked to dispersal and fertility. By reviewing
the literature on taxa where workers can reproduce
sexually (i.e. become gamergates) we show that (1) species
with a single gamergate generally have lost the winged
queen caste, whereas only half of the species with several
gamergates have, and (2) single-gamergate species have
smaller colonies than multiple-gamergate species. Comparison
with “classical” ants without gamergates, where
having one vs having several winged queens are two
distinct syndromes, suggests that having one vs having
several gamergates are not. Gamergate number does not
affect the success of colony fission, but retention of the
queen caste permits the option of independent foundation.
Keywords Queenless ants . Monogyny. Polygyny.
Colony size . Dispersal . Fission
Introduction
Colony-founding strategies of ants are largely shaped by
selective pressures on the survival and dispersal of sexuals
(e.g. Bourke and Franks 1995; Peeters and Ito 2001). At
one extreme, species favour dispersal and produce many
winged queens that fly away from the colony to mate and
start new colonies alone. This strategy of independent colony
foundation is characterised by the production of many
T. Monnin (*) : C. Peeters
Fonctionnement et évolution des systèmes écologiques
CNRS UMR 7625, Université Pierre et Marie Curie,
7 quai Saint Bernard,
75005 Paris, France
e-mail: Thibaud.Monnin@snv.jussieu.fr
C. Peeters
e-mail: cpeeters@snv.jussieu.fr
reproductives with high dispersal but low success. In contrast,
other species produce few queens dispersing on foot
accompanied by many workers. This strategy of colony
fission (also termed budding) is characterised by the production
of few reproductive propagules [queen(s) and accompanying
workers] with low dispersal but high success.
Some species mix both strategies by producing queens
dispersing alone on the wing as well as reproductives dispersing
on foot with workers (e.g. Peeters 1997; Heinze and
Keller 2000; Peeters and Ito 2001). This is the case of some
ants where the workers are capable of mating and sexual
reproduction. Some of these species have lost the queen
caste and exclusively reproduce by colony fission, with one
or more mated workers (called gamergates, Peeters and
Crewe 1984) establishing new colonies with the help of
nestmate workers. However, others have retained the queen
caste and can thus reproduce by both independent colony
foundation and colony fission.
Species with gamergates are restricted to three subfamilies
characterized by a low degree of dimorphism between
winged queens and workers (Amblyoponinae, Ectatomminae,
Ponerinae). We review the literature and investigate
the relationships between occurrence of the queen caste,
gamergate number and colony size. We discuss this in
the light of the evolution of multiple-breeders societies
and the transition from independent colony foundation to
colony fission, and we draw comparisons with the monogynous
and polygynous “syndromes” often used to categorise
“classical” ants with winged queens (reviewed in e.g.
Bourke and Franks 1995).
Materials and methods
Although gamergates were first described in species that
lack a queen caste, hence the now familiar “queenless ant”

110 Naturwissenschaften (2008) 95:109–116
label, gamergates also occur in some species that have
retained winged queens. In the present paper, “queenless”
refers to species that always lack the queen caste, and we
explicitly state when we refer to species that have both
queens and gamergates.
We compare single- vs multiple-gamergate species for the
occurrence of queens and colony size. Thirty-five species
(11 genera) for which 6 to 129 colonies were collected are
included in our analysis. Eleven species with five or less
colonies collected are excluded because gamergate number
and colony size cannot be determined reliably. We cannot
control for phylogenetic relationships because there are no
species-level phylogenies in the genera investigated. We limit,
to some extent, the potential problems of non-independence
of species by working at the genus level in addition to species
level. To do this, we pool congeneric species with one vs
several gamergates and compare genera for the occurrence of
the queen caste and colony size. Working at the genus level
is justified because each genus where gamergates occur represents
at least one independent evolution of gamergates.
Indeed, all the genera with gamergates included in recent
molecular phylogenies are more related to one or several
genera without gamergates than to another genus with gamergates
(Moreau et al. 2006; Brady et al. 2006). For instance,
Rhytidoponera and Gnamptogenys (with gamergates) are
more closely related to Ectatomma and Typhlomyrmex (without
gamergates), respectively, than to each other. Additionally,
8 of the 11 genera with gamergates include “classical”
species with queens and no gamergates. The hypothesis that
gamergates evolved independently within each of these
genera is more parsimonious than the hypothesis of evolution
in a common ancestor. Only two genera (Pachycondyla and
Platythyrea) include both single- and multiple-gamergate
species, and both also have “classical” species with queens
and no gamergates. For these genera, we pooled species
according to gamergate number (either single-gamergate or
multiple-gamergate species).
Our genus-level approach is validated by recent molecular
phylogenies that have established the monophyly of
currently recognized genera in subfamilies Ponerinae and
Ectatomminae (Brady et al. 2006; Moreau et al. 2006). A
notable exception is Pachycondyla in which several genera
have been synonymized (Bolton 2003). We take a conservative
approach and consider Pachycondyla as one unit,
although gamergates may have evolved repeatedly in its
constituent taxa. In contrast, we consider Dinoponera and
Streblognathus as unrelated because new molecular data
show that they are not sister genera (J. Schmidt, personal
communication), unlike previously thought.
We define colony size as the number of workers plus
dealated queens (i.e. queens who shed their wings), irrespective
of whether these queens were mated and laying eggs or
not, thereby excluding winged gynes (young virgin queens)
and males. This is because dealated queens will stay in the
colony, whereas alate queens and males will leave. For species
where some colonies have queens and others have gamergates,
average colony size is given for both types of colonies, but all
colonies were pooled for inter-specific comparisons.
Results
Gamergate number and occurrence of the queen caste
Gamergate number strongly correlates with the presence
of queens. All single-gamergate species in our sample
lack queens (n=19, Table 1), whereas 8 of 16 multiplegamergate
species have queens (Table 2, Yates’ corrected
χ 2 =9.64, df=1, p=0.0019). This difference remains true
when pooling species by genus: none of the six genera with
single-gamergate species has species with both gamergates
and queens, whereas five of the seven genera with multiplegamergate
species have (Tables 1 and 2, Yates’ corrected
χ 2 =4.27, df=1, p=0.0387). Importantly, queens are unknown
in Diacamma, Dinoponera and Streblognathus, three
genera where all species have a single gamergate, whereas
queens exist in various species belonging to the other genera
with gamergates.
Multiple-gamergate species form a heterogeneous group and
no pattern emerges regarding the occurrence of queens
(Table 2). The numbers of gamergates and queens vary
widely, and either caste of female can dominate reproduction.
Several species have colonies with multiple gamergates and/or
asinglequeen(n=7), and one has multiple gamergates or
several functional queens (Gnamptogenys striatula). Queens
and gamergates can cohabit in the same colonies in three
species, namely, Pachycondyla (Bothroponera) tridentata
(Sommer and Hölldobler 1992; Sommer et al. 1994), Platythyrea
quadridenta (Ito 1994) andPlatythyrea sp.1(F.Ito,
personal communication). In P. tridentata and P. quadridenta,
some dealated queens do not lay eggs, and in
P. tridentata this is because queens can be behaviourally
dominated by gamergates and consequently suppressed
from reproducing. Queens and gamergates never occur
simultaneously in the other five species. In Harpegnathos
saltator, queens found new colonies independently after a
nuptial flight, and some of their daughter workers mate
and reproduce after their death. Unlike other ants with
gamergates, colonies do not propagate by fission, but
queens perform independent colony foundation, with
colonies headed by gamergates producing new queens
annually. Thus each colony goes through a queen phase
followed by a gamergate phase (Peeters and Hölldobler
1995; Peetersetal.2000). In Gnamptogenys menadensis,
colonies founded by a queen also become headed by
gamergates after the death of the queen. However, unlike in

Naturwissenschaften (2008) 95:109–116 111
Table 1 Average colony size of single-gamergate species
Species Colony size mean±SD
(range)
H. saltator, most colonies are produced by fission and do not
go through a queen phase. Indeed, only 5% of colonies are
headed by a queen (Gobin et al. 1998). This contrasts with
G. striatula where 96% of colonies have queens and no
gamergates. This species has numerous queens (14±18,
mean±SD) so that it is unlikely that a colony ever loses all
of them. However, workers start to perform sexual calling in
experimentally orphaned colonies (Blatrix and Jaisson 2000),
and in one population gamergates reproduced in two
colonies lacking queens (Blatrix 2000; Blatrix and Jaisson
2000). In Rhytidoponera chalybea and Rhytidoponera confusa,
workers also differentiate into gamergates after queen
death (Ward 1983, C. Peeters, unpublished data). Last,
Rhytidoponera metallica occasionally produce queens. These
are capable of starting colonies in the laboratory (Ward
1986), but they have never been found to head colonies in
nature, suggesting that this species is in the process of losing
the queen caste.
Gamergate number and colony size
Our considerably expanded sample of species confirms
Peeters’ (1993) remark that single-gamergate species have
Excavated
colonies
smaller colonies than multiple-gamergate species, with 83±
83 workers in the former vs 169±154 in the latter (mean±
SD, single-gamergate species: n=19, median=54, quartiles=
15 and 124, Table 1; multiple-gamergate species: n=16,
median=136, quartiles=41 and 237, Table 2; colonysizes
are not normally distributed and were transformed in square
roots, which are normally distributed and show homogeneity
of variance; t test for independent samples: t=−2.069,
df=33, p=0.046, Fig. 1a). Pooling species by genus shows
the same trend, albeit not statistically significant (Fig. 1b):
genera with single-gamergate species have colonies of 64±61
workers, whereas genera with multiple-gamergate species
have colonies of 133±107 workers (single-gamergate genera:
n=6, median=56, quartiles=23 and 68, Table 3; multiplegamergate
genera: n=7, median=102, quartiles=65 and 251,
Table 3; Mann–Whitney U: z=−1.500, p=0.134).
Discussion
References
Diacamma australe 129±56 9 Peeters and Higashi 1989
Diacamma ceylonense “nilgiri” 267±109 10 Cournault and Peeters, submitted for publication
Diacamma cyaneiventre 214±80 6 André et al. 2001
Diacamma ceylonense 247 (91–499) 39 Cuvillier-Hot et al. 2001, Karpakakunjaram et al. 2003
Diacamma sp. from Japan 124±50 29 Fukumoto et al. 1989, Kawabata and Tsuji 2005
Diacamma sp. from Malaysia 86±41 51 Sommer et al. 1993
Dinoponera australis 15±7 66 Fowler 1985, Paiva and Brandão 1995, Paiva 1997,
Monnin, unpublished data
Dinoponera gigantea 54±27 10 Overal 1980, Fourcassié and Oliveira 2002, Monnin,
unpublished data
Dinoponera quadriceps 73±39 70 Monnin and Peeters 1999, Monnin, unpublished data,
Vasconcellos et al. 2004
Pachycondyla (crassa-group) krugeri 45±22 19 Peeters and Crewe 1986, Wildman and Crewe 1988, Villet
and Wildman 1991
Pachycondyla (Hagensia) havilandi 25±9 13 Villet 1992
Pachycondyla (Bothroponera) sp.
8 from West Java
11±6 8 Ito 1993a
Pachycondyla (Bothroponera) 9(2–18) 53 Peeters et al. 1991, Ito and Higashi 1991, Higashi et al.
sublaevis
1994
Platythyrea lamellosa 115±83 25 Villet et al. 1990
Platythyrea schultzei 21±10 7 Villet 1991
Streblognathus aethiopicus 35 (9–51) 24 Ware et al. 1990, Cuvillier-Hot 2002
Streblognathus peetersi 95±45 44 Cuvillier-Hot 2002
Thaumatomyrmex atrox 4(1–9) 52 Jahyny et al. 2002
Thaumatomyrmex contumax 2.5 (1–9) 8 Jahyny et al. 2002
When several studies were pooled the SD could only be calculated when the authors provided the raw data. Otherwise, the range is given
(minimum–maximum). All the species belong to the subfamily Ponerinae (Bolton 2003).
Our review reveals two clear patterns: single-gamergate
species mostly lack queens (a few exceptions exist; F. Ito,
personal communication), whereas multiple-gamergate spe-

112 Naturwissenschaften (2008) 95:109–116
Table 2 Average colony size and occurrence of the queen caste in multiple-gamergate species
Species Colonies with gamergates only Colonies with queen(s) References
Excavated
colonies
Colony
size
Gamergate
number
Queen
number
Colony size Excavated
colonies
Gamergate
number
Amblyopone reclinata 6±5 102±50 22 Ito 1993b
Gnamptogenys menadensis 5±4 111±77 35 1 0 141±43 2 Gobin et al. 1998
Gnamptogenys striatula 25/565 and 445±226 2 14±18
31/266
a
0 386±313 47 Blatrix 2000, Blatrix and Jaisson
2000
Harpegnathos saltator b
3 to 6 83±58 16 1 0 58±30 43 Peeters et al. 2000
Leptogenys peuqueti 4±4 30±24 13 Ito 1997
Leptogenys schwabi 6/260 and 184±76 17 Davies et al. 1994
7/146
Leptogenys sp. 24 3±4 13±7 8 Ito 1997
Pachycondyla (Ophthalmopone) 13±16 158±160 37 Peeters and Crewe 1984, 1985, 1986,
berthoudi
Sledge et al. 1996, 1999, 2001
Pachycondyla (Bothroponera) 81 to 100% 42±16 3 1
tridentata
c
86 to 100% 58±28 4 Sommer et al. 1994
Platythyrea quadridenta 2±2 18±15 14 1 d
2±3 31±30 2 Ito 1994, personal communication
Platythyrea sp.1 3±2 17±6 10 1 4±2 19±3 3 F. Ito, personal communication
Rhytidoponera aurata e
22±9 182±98 13 Komene et al. 1999
Rhytidoponera chalybea f
4±3.5 181±154 48 1 0 440±171 21 Ward 1983
Rhytidoponera confusa f
4±3.5 164±208 64 1 0 264±144 65 Ward 1983
Rhytidoponera metallica 5 to 15% 381 (79–1,092) 16 Chapuisat and Crozier 2001, Thomas
and Elgar 2003
Rhytidoponera mayri (sp. 12) 20±19 528±269 12 Pamilo et al. 1985, Peeters 1987, Tay
and Crozier 2000
Gamergate number, colony size and dealated queen number are given as mean±SD (range is given when SD could not be determined). The fraction of gamergates (e.g. 25 gamergates out of 565
workers in G. striatula) is given when only few colonies were dissected or when colonies were not dissected in their entirety. Gnamptogenys and Rhytidoponera belong to the subfamily
Ectatomminae, Amblyopone belongs to Amblyoponinae, and the other genera belong to Ponerinae (Bolton 2003).
a
Alate queens were included because many were mated and laying eggs. Most queens reproduce; 66±41% of 100 queens from 18 colonies were mated and laying eggs (84±29% for dealate
queens and 43±44% for alate queens, Blatrix 2000).
b
The population from Bangalore is atypical (Peeters et al. 2000) and therefore excluded from these data.
c
Only 1 out of 5±9 dealated queens (47% of which were inseminated) had developed ovaries and laid eggs in one colony, simultaneously with several gamergates (Sommer et al. 1994).
d
This colony contained a second dealated but non-laying queen (Ito 1994).
e
In R. aurata, 11 additional colonies were found to have only one gamergate and to be smaller, with only 75±38 workers (Komene et al. 1999). This bimodal distribution differs from other
multiple-gamergate species, where gamergate number varies continuously, and Komene et al. (1999) suggest that these small monogynous colonies are new colonies founded by fission.
f
Ward (1983) gives the average gamergate number for R. chalybea and R. confusa colonies pooled together but not for each species.

Naturwissenschaften (2008) 95:109–116 113
Colony size
a species level b genus level
250
200
150
100
50
0
Single Multiple
cies often retain them, and the former species have smaller
colonies than the latter. These patterns are also true at the
genus level. We estimate that ca. 200–300 species of ants
have gamergates, and thus future studies will enrich this
comparative approach.
Gamergates cannot found new colonies independently,
for both proximate (e.g. no metabolic reserves) and ultimate
Table 3 Colony size at the genus level, calculated by averaging the
mean colony size of the species of the given genus
Genera Colony
size
350
300
250
200
150
100
Number of
species
Single Multiple
Gamergate number
Fig. 1 Colony size of single- and multiple-gamergate species and
genus. a Colony size at the species level. The graph shows the backtransformed
means±95% confidence intervals of the squared root
data. Single-gamergate species have smaller colonies than multiplegamergate
species. b Colony size at the genus level. Box plots give
median, quartiles and range. Single-gamergate genera tend to have
smaller colonies than multiple-gamergate genera, but the difference is
not significant. See results for statistics
Single gamergate
Diacamma 178 6 144
Dinoponera 47 3 146
Pachycondyla 23 4 93
Platythyrea 68 2 32
Streblognathus 65 2 68
Thaumatomyrmex 3 2 60
Multiple gamergates
Amblyopone 102 1 22
Gnamptogenys 251 2 86
Harpegnathos 65 1 59
Leptogenys 76 3 38
Pachycondyla 105 2 44
Platythyrea 19 2 29
Rhytidoponera 313 5 239
Excavated
colonies
Pachycondyla is the only genus with both single- and multiplegamergate
species, and this reflects its polyphyletic status
50
0
(fitness advantages of staying in natal colony, given limited
dispersal on ground) reasons (Liebig et al. 1998). Accordingly,
colony fission is obligate in gamergate species without
winged queens. This is true regardless of gamergate
number which has no known impact on the success of fission.
Therefore, with respect to colony dispersal, having
one vs several gamergates may not be two distinct strategies
analogous to having one vs several queens in
“classical” ants. Indeed, in the latter, the transition from
monogyny to polygyny is often associated with a transition
from independent colony foundation to colony fission (e.g.
Solenopsis invicta, Ross and Keller 1995; Myrmica rubra,
Walin et al. 2001; Formica, Sundström et al. 2005). Bourke
and Franks (1995) reviewed various factors that favour the
evolution of multiple-queen societies, such as high predation
on queens during mating and nest initiation, habitat
patchiness (dispersing queens may fail to find a suitable
habitat) and nest site limitation. Clearly, these factors also
favour a shift from independent colony foundation to fission.
Habitat saturation (i.e. incipient colonies may be
outcompeted) favours large multiple-queen colonies reproducing
by colony fission (e.g. Crozier and Pamilo 1996;
Tschinkel 2006). Nonetheless, gamergate number affects
nestmate relatedness, so that having one vs several gamergates
is similar to having one vs multiple queens with respect
to colony kin structure.
The fact that multiple-gamergate societies often have
winged queens (Table 2) complicates the comparison with
single-gamergate societies, which typically lack queens
(Table 1). Thus we need to consider separately species that
have lost the queen caste from those that have retained it.
Among species that lack queens, gamergate number is not
associated with clear differences in ecological requirements
of colonies. Colony fission occurs irrespective of whether
there are one or multiple gamergates. Gamergate colonies
are potentially immortal because any young worker can
replace a dying gamergate, provided that foreign males are
present. In contrast, monogynous colonies of “classical”
ants will often go extinct upon the death of the founding
queen, whereas polygynous colonies will not as they typically
readopt young queens. Hence, single- and multiplegamergate
societies do not differ in terms of long-term
stability or capacity to reproduce by fission, and do not
seem to represent distinct “syndromes”.
It is not surprising that some species with gamergates
have retained winged queens. Although independent colony
foundation is risky, it can yield large benefits such as
colonising a distant patch of suitable habitat. Species in
which both queens and gamergates reproduce are comparable
to “classical” ants having two queen phenotypes, i.e.
macrogynes and microgynes, or queens with and without
stored energetic reserves (e.g. Rosengren et al. 1993;
Heinze and Keller 2000; Rüppell et al. 2001). Presumably,

114 Naturwissenschaften (2008) 95:109–116
gamergate species in which queens were eliminated have
experienced stronger selection against independent colony
foundation relative to species that retained queens. One
hypothesis for the retention of the queen caste is that
species having both queens and gamergates live in more
unstable or patchier environments, where the capacity to
disperse over long distances is strongly favoured. Indeed,
restricted dispersal has been shown in a species without
queens (Doums et al. 2002). However, our knowledge of
the biology and ecology of ants with gamergates is too
limited to test this hypothesis.
Bourke (1999) argues that colony size and social complexity
are key determinants of social insect biology. All
species with gamergates have small colonies (Tables 1 and 2)
compared to that of higher ants, which can reach hundreds of
thousands of workers. However, single-gamergate species
(or genera) have smaller colonies than multiple-gamergate
species (genera). Why? Gamergates generally have low
fertility (because they belong to the worker caste), and this
may limit colony size in single-gamergate species. However,
intra-specific comparisons suggest that this is not the whole
story. Among species where gamergate number varies
among colonies, single-gamergate colonies are smaller than
multiple-gamergate colonies in Rhytidoponera aurata
(Komene et al. 1999), but there is no difference in
Amblyopone sp. (Ito 1993b), G. menadensis (Gobin et al.
2001), Leptogenys peuqueti (Ito 1997) andPachycondyla
(Ophthalmopone) berthoudi (Peeters and Crewe 1984;
Sledge et al. 1996, 1999, 2001). This lack of a correlation
between gamergate number and colony size may be explained
by a decrease in individual fertility as breeder number rises
(P. berthoudi, Sledge et al. 1996).
Based on what is known about the life history of 46
species with gamergates, we propose that the evolution of
gamergate number and the loss of the queen caste follows
a general pattern. Firstly, reproduction by multiple gamergates
evolves due to the adaptive benefits of secondary
polygyny, i.e. increased colony lifespan and resource inheritance
(Peeters and Ito 2001), and gamergates coexist
with queens. This is likely the case in species where colonies
are obligatorily founded by a queen (i.e. H. saltator,
Peeters and Hölldobler 1995; Peeters et al. 2000). Secondly,
winged queens are lost whenever independent colony
foundation is no longer selected for, giving rise to species
where multiple gamergates are the sole reproductives.
Thirdly, gamergate number is reduced, and this gives rise
to single-gamergate species. This pattern is supported by
the lack of a queen caste in almost all single-gamergate
species vs only half the multiple-gamergate species. One or
more of these three stages are found in each genus having
gamergates. Future studies must investigate whether reduction
in gamergate number increases colony efficiency, e.g.
by reducing the number of hopeful reproductive workers
(Molet et al. 2005) and by reducing their probability of
becoming gamergate and thus increasing their helping
effort (Field et al. 2006). To conclude, how many gamergates
is a queen worth? With respect to long-range
dispersal, winged queens are irreplaceable, which is why
many gamergate species have retained the queen caste.
Acknowledgements We thank Claudie Doums, Fuminori Ito and
Jürgen Heinze for constructive discussions and comments, as well as
two anonymous referees for useful suggestions.
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