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ANIMAL BEHAVIOUR, 1999, 58, 287–294

Article No. anbe.1999.1149, available online at http://www.idealibrary.com on

Operational sex ratio influences female preference and male–male


competition in guppies

MULLICA JIROTKUL
Department of Ecology, Evolution and Marine Biology, University of California, U.S.A.

(Received 21 May 1998; initial acceptance 6 August 1998;


final acceptance 4 April 1999; MS. number: 5858R)

Manipulation of the operational sex ratio (OSR) in guppies, Poecilia reticulata, causes changes in
male–male competition and female mate choice. In this study OSR is defined as the number of sexually
active males divided by the total number of sexually active adults of both sexes. The rate of male
courtship displays decreased, and interference behaviours between males increased, at male-biased OSRs.
The OSR influenced both copulatory tactics and postcopulatory guarding. All copulations followed
sigmoid displays, except at an OSR of five males to one female where 60% of copulations occurred during
sneak attempts. Compared with copulations that followed sigmoid displays, successful sneak copulations
were followed by a shorter period of postcopulatory mate guarding and a shorter refractory period before
males resumed courtship activities. Females preferred males with more orange colour whenever they had
a choice, and the preference for orange colour was stronger with more male-biased OSRs. The OSR thus
influences the presence, absence or relative importance of both female mate choice and male–male
competition which, in turn, should affect the evolution of secondary sexual traits.
 1999 The Association for the Study of Animal Behaviour

The evolution of elaborate sexual characters can be more potential mates. In sex-role-reversed species,
explained by both competition for mates and mate females compete more intensely at female-biased OSRs
choice (Darwin 1871). The impact and intensity of these (Colwell & Oring 1988; Berglund 1994; Vincent et al.
two modes of sexual selection may vary in a given 1994). Thus, changes in OSR seem to trigger changes in
time and place, and can occur either independently or competition intensity and may cause changes in male
simultaneously (Höglund 1989; Andersson 1994). The mating tactics (Verrell 1983; Krupa & Sih 1993; Berglund
operational sex ratio (OSR) is thought to determine, 1994; Wearing-Wilde 1996) and prolong copulations
in part, the relative importance of mate choice and (Clark 1988; Souroukis & Murray 1995; Vepsäläinen &
sexual competition (Clutton-Brock & Vincent 1991; Savolainen 1995; Alonso-Pimentel & Papaj 1996; Dick &
Clutton-Brock & Parker 1992; Owens & Thompson 1994). Elwood 1996; Yamamura & Jormalainen 1996).
Here I define OSR as the ratio of the number of sexually The OSR is influenced by three main factors: the differ-
active males to the sum of sexually active males and ence in the potential reproductive rate of each sex; the
receptive females at a given time in a population adult sex ratio; and the distribution of sexually mature
(Kvarnemo & Ahnesjö 1996). adults in a population (Clutton-Brock & Parker 1992;
A bias in OSR can alter both male–male competition Kvarnemo et al. 1995). Potential reproductive rate is
and female mate choice. Many studies have shown that defined as the potential rate at which each sex
in male-biased OSRs, males tend to compete more contributes offspring to the next generation. In species
intensively for mates because they have more potential where males do not provide parental care, the male
competitors (e.g. Lawrence 1986; Enders 1993; Krupa & potential reproductive rate is limited only by the number
Sih 1993; Jormalainen et al. 1994; Dick & Elwood 1996; of receptive females. The OSR will be biased towards
Clutton-Brock et al. 1997), while a study of St Peter’s fish, the more abundant sex with the higher potential
Saratherodon galilaeus, showed that females become more reproductive rate.
selective because of a greater opportunity to choose mates I investigated experimentally the quantitative effects of
(Balshine-Earn 1996). At female-biased OSRs, males of the OSR as a continuous variable on female mate choice,
many species become more selective because there are and male–male competition. Guppies, Poecilia reticulata,
Correspondence and present address: M. Jirotkul, Institute of Science, are appropriate test subjects for these effects for several
Walailak University, 222 Thasala District, Nakhon Si Thammarat, reasons: the OSR and female preference for orange colora-
80160, Thailand (email: jmullica@praduu2.wu.ac.th). tion in males can be measured quantitatively; the adult
0003–3472/99/080287+08 $30.00/0 287  1999 The Association for the Study of Animal Behaviour
288 ANIMAL BEHAVIOUR, 58, 2

sex ratio fluctuates considerably within and between were reared with other females in their aquaria. This
guppy populations (Seghers 1973); the local sex ratio also allowed males to gain sexual experience prior to being
varies between pools within the same river, from 0.2 to observed for this study. Fish were reared at 22–24C on a
0.9 (G. Grether, unpublished data); females are ready to 12:12 h light:dark photoperiod, and were used in the
mate only when they are virgin, or just after giving birth experiments when they were 3 months old. All fish were
which occurs about once a month, so the number of fed twice daily on newly hatched brine shrimp in the
receptive females at any given time is very low, resulting morning, and Tetramin flake food and liver paste food,
in a strongly male-biased OSR (Houde 1987); and finally alternated, in the afternoon.
guppies are one of the most thoroughly studied species The female’s response to male courtship displays varies
with respect to sexual selection (Andersson 1994; Houde depending on male traits, male behaviours and female
1997). Understanding the effect of the OSR on sexual receptivity. Houde (1987) ranked female responses on
selection in guppies may help in developing a general a scale of 0–5: (0) no response; (1) stopping current
model of sexual selection for other species. activities and looking at a courting male; (2) un-
ambiguously following the displaying male; (3) a gliding
METHODS motion with stiffened body and arched towards the male;
(4) both the focal male and female spinning in circles;
Guppies are small poeciliid fish native to the coastal and (5) rapid spinning, copulating and the focal male
mountain ranges of northeastern South America and the jerking backwards. Explicit sexual behaviour was defined
adjacent islands of the Caribbean (Haskins et al. 1961; as a female response ranked 2 or above.
Rosen & Bailey 1963). Males have very complex colour The relative area of orange coloration on the body and
patterns and courtship behaviours. Females are dull and tail of males from the Paria population is 103%
usually bigger than males. Males provide no parental care, (N=200) and I used these values as the meanSD
nor do they defend territories. They spend most of their percentage orange area of the laboratory Paria popu-
time following females, courting and attempting to mate. lation. I manipulated the OSR by changing the number of
After a successful copulation, which occurs in nature only sexually active males and receptive females while keeping
after many copulatory attempts, some males guard their the total population density constant. Five replicates
mate by swimming with her and fending off other males. of each treatment were run with new groups of fish at
Male guppies use three mating tactics: sigmoid displays; each of the following OSRs: 1:5; 2:4; 3:3; 4:2 and 5:1
sneak attempts; and interference with other courting males:females (i.e. 0.17, 0.33, 0.50, 0.67 and 0.83 OSR).
males (Liley 1966). Sigmoid display behaviours include Prior to observations in a given trial, I sketched each
males bending their bodies into a sigmoid shape, an male’s colour pattern and used these sketches to identify
oscillating movement along the body axis, and moving each male during the trial. Male guppies chosen for the
up and down in front of the females (Baerends et al. 1955; experiment averaged 10% orange body area but all fell
Liley 1966; Houde 1987). Sneak attempt behaviours within a range of 5–15% which is similar to the natural
include swimming behind females and thrusting of the Paria population. To measure male colour patterns, I used
gonopodium without initial courtship (Farr 1980). Inter- the photographic method developed by Endler (1978).
ference behaviours include cutting in front of another At 0655 hours, each replicate of six fish was placed in
courting male and attempting to court the female. This an aquarium measuring 1.00.3 m0.45 m high and lit
interference behaviour is not an aggressive behaviour and by a fluorescent light and several incandescent lights
is very brief and noninjurious. Both sigmoid displays and which yielded an irradiance of 10 ìmol/m2 per s at the
sneak attempts can lead to successful mating (Liley 1966). water surface. The latter is similar to natural habitat light
Interference also increases the probability of the interfer- conditions (Endler 1991). In each replicate, I allowed fish
ing male mating successfully because he increases his to acclimatize for 5 min before starting to record behav-
chances of courting females. iours at two times: (1) 0700 hours (time (t)=0), and (2)
I used the first- or second-generation offspring of 0700 hours the day after introduction (t=24).
guppies descended from wild guppies collected in April During the first period of observation (t=0), I observed
and October 1996 from the Paria River, Trinidad. The all behaviours for 40 min, changing focal males about
Paria females prefer males with a large degree of orange every 10 s which allowed me to record all behaviours by
coloration (Houde 1987, 1988, 1997). Pregnant females all males including copulations and postcopulatory
were obtained from the laboratory stock (stock tanks guarding. Each male in a replicate had different colour
measured 1.00.45 m0.6 m high, with ca. 100–120 patterns in order to avoid misidentification. At t=24, each
fish per tank) and placed into individual holding tanks, male was chosen at random as the focal male for 5 min.
measuring 0.30.2 m0.15 m high, until they gave After all males had been observed for three 5-min inter-
birth. Offspring were collected within 24 h of birth, vals, observation continued for 30 min more during
and reared in aquaria measuring 0.60.25 m0.3 m which time I changed focal males about every 10 s. This
high with 40–50 fish per tank. Males and females were additional 30-min observation period increased the
separated when they were 4–5 weeks old and kept in opportunity to observe each male interacting with more
tanks measuring 0.60.25 m0.3 m high, with 20–25 females in the tank.
fish per tank. Females were reared to maturity and com- During each observation period, I recorded the
pletely isolated from mature males. Males were kept numbers of sigmoid displays, female responses to
isolated from females that were used in this study but each sigmoid display, male sneak attempts, male–male
JIROTKUL: OPERATIONAL SEX RATIO IN GUPPIES 289

interference behaviours and copulations, total courtship longer courtship (F2,22 =300.03, P<0.001, r 2 =0.97,
time (at t=24 only), mate-guarding duration, refractory Y=0.42–1.92X+4.68X2, t 2X =7.60, P<0.001; Fig. 2b).
duration, and the number of sigmoid displays and sneak Females also received more interferences at male-biased
attempts that ended in copulation. At t=24, I also OSRs (F2,22 =120.70, P<0.001, r 2 =0.92, Y=23.20–
recorded how often males switched between females. 176.69X+3001.00X2, t 2X =6.56, P<0.001; Fig. 2c) but there
Total courtship time could not be recorded at t=0 was no significant trend in the number of sneak attempts
because, during this observation period, I did not do a that females received (Fig. 2d).
5-min male focal observation. For each replicate, all
behaviours were calculated as average rates per male per
15 min. Female Preference
Female preference for orange colour in males in each At female-biased OSRs, males received more total
replicate was the sum of all female preferences in that female responses (all females in the replicate combined;
replicate because females could not be identified. I esti- F1,23 =17.53, P<0.005, r 2 =0.43, Y=78.16–41.07X; Fig. 1e),
mated male attractiveness by using the fraction of female but females responded more to courting males at male-
responses (FFR; Houde & Endler 1990; Endler & Houde biased OSRs (F2,22 =43.77, P<0.001, r 2 =0.80, Y=69.97–
1995): FFR was the number of male displays that elicited 388.49X+691.52X2, t 2X =3.76, P<0.005; Fig. 2e). The
a female response of 2 or more on the 0–5 scale divided by slopes of female preference for orange coloration in males
the total number of male displays. The strength of female had a significantly positive relationship with the OSR
preference for orange coloration was estimated as the (F1,18 =5.16, P< 0.05, r 2 =0.22, Y= 0.76+6.58X; Fig. 3).
regression of FFR on the orange area of males in each
replicate.
For two reasons, only the behavioural data at t=24 were Copulatory Tactics and Postcopulatory Guarding
used in analyses. First, during the first observation period,
virgin females are highly receptive and tend to mate At male-biased OSRs, individual males copulated less
indiscriminately. These indiscriminate matings do not often (F1,23 =43.45, P<0.001, r 2 =0.65, Y=3.26–3.47X; Fig.
usually occur in nature and these females become more 1f), but females copulated more frequently (F1,23 =24.43,
selective 24 h after introduction (Houde 1987, 1988, P<0.001, r 2 =0.52, Y= 0.15+3.23X; Fig. 2f ). All copu-
1997). Therefore, the behavioural data at t=24 were more lations followed sigmoid displays, except at 0.83 OSR
likely to represent what happens in nature. Second, after where only 40% of copulations followed sigmoid displays.
successful copulations, which usually happened within The remaining 60% occurred during sneak attempts (nine
24 h of introduction, males usually stopped all courting sneak copulations out of 15 copulations). Also at 0.83
activities for up to 20 min (personal observation). Using OSR, sneak copulations were followed by a shorter guard-
the behavioural data at t=24 is a common practice in ing period (Kruskal–Wallis test: H1 =9.65, P<0.005) and
guppy sexual selection studies (Houde 1987, 1988; Houde males resumed courting females sooner (one-way
& Endler 1990; Endler & Houde 1995). The FFR and ANOVA: F1,13 =7.44, P<0.05) than males who mated suc-
orange colour were arcsine transformed to achieve nor- cessfully after sigmoid displays. The postcopulatory
mality. For each replicate, I used each individual male’s guarding duration and refractory period were constant at
behaviour to calculate a mean of that behaviour per other OSRs (one-way ANOVAs: guarding time effect:
replicate per 15 min. I tested all variables for normality F4,20 =0.78, NS; refractory time effect: F4,20 =2.56, NS).
using Lilliefors’ test, and used nonparametric tests where
normality or other assumptions of parametric tests were DISCUSSION
not met. If the coefficient (c) of OSR2 was significant, I
used quadratic partial regressions. This means that the The average adult sex ratio of wild guppies from various
regression can be explained by a quadratic curve. All streams in Trinidad, West Indies is 0.42 (Seghers 1973).
significance tests were two tailed. Seghers (1973) showed that the adult sex ratio from the
Paria River varies spatially from 0.35 to 0.40 and tem-
porally from 0.40 to 0.53. Clearly, the adult sex ratio in
RESULTS the wild tends to be female biased. However, females are
Male Mating Tactics rarely receptive. Consequently, the OSR is strongly male
biased (Houde 1988); thus, only the 0.67and 0.83 OSRs of
With a more male-biased OSR, males displayed less this experiment may reflect the real OSRs that occur in
(F1,23 =14.01, P=0.001, r 2 =0.38, Y=32.77–27.90X; Fig. 1a) the Paria River. Nevertheless, the adult sex ratios of wild
and courtship was shorter (F1,23 =107.51, P<0.001, guppies in other streams in Trinidad vary from 0.20 to
r 2 =0.82, Y=1.03–0.72X; Fig. 1b) but they interfered with 0.85 (G. Grether, unpublished data), so general conclu-
other males more (F1,23 =94.62, P<0.001, r 2 =0.80, sions from other OSR treatments in my study may apply
Y= 5.67+30.47X; Fig. 1c). Males sneaked the most at to other wild guppy populations.
0.5 OSR (F2,22 =5.95, P<0.01, r 2 =0.35, Y=1.19+17.78X– A male-biased OSR would be expected to increase male–
22.35X2, t 2X = 2.38, P<0.05; Fig. 1d). male competition for mates because there are proportion-
At male-biased OSRs, females received more sigmoid ally fewer females. This could affect male mating tactics,
displays (F2,22 =26.82, P<0.001, r 2 =0.71, Y= 17.28+ male reproductive success, and female mate choice. This
132.55X–95.74X2, t 2X = 3.24, P<0.005; Fig. 2a), and is the second species in which males are known to use
290 ANIMAL BEHAVIOUR, 58, 2

50 1.2
45 (a) * 1.1 (b) **

Courtship time (min)/male


40 1
Sigmoid displays/male

35 0.9
30 0.8
25 0.7
20 0.6
15 0.5
10 0.4
5 0.3
0 0.2
0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9

30 10
(c) ** 9 (d) *
25
8
Interferences/male

7
20
Sneaks/male
6
15 5
4
10
3
2
5
1
0 0
0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9

100 5
90 (e) ** 4.5 (f) **

80 4
70 3.5
Copulations/male

60 3
FFR/male

50 2.5
40 2
30 1.5
20 1
10 0.5
0 0
0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9

Operational sex ratio


Figure 1. Relationship between the operational sex ratio (males/(males+females)) and (a) the number of sigmoid displays per male, (b) the
total courtship time (min) per male, (c) the number of interference behaviours per male, (d) the number of sneak attempts per male, (e)
fraction of female responses (FFR) per male, and (f) the total number of copulation events per male. Each point represents the average of male
behaviour in each replicate. +indicates points with overlapping data. Linear or quadratic regressions were used: *P<0.01; **P<0.001.

different combinations of alternative mating tactics rate, copulation duration, postcopulatory guarding and
depending on the OSR. Studies of water striders (Aquarius mating activity (Krupa & Sih 1993; Sih & Krupa 1995;
remigis and Gerris lacustris) have shown that the OSR Vepsäläinen & Savolainen 1995). In my study, a lack of
affects the movement strategies of males (i.e. changing females to approach using sneak behaviours may explain
sites or remaining stationary at hot spots), harassment why fewer sneak attempts occurred at more male-biased
JIROTKUL: OPERATIONAL SEX RATIO IN GUPPIES 291

50 2.5
45 (a) * (b) *

Courtship time (min)/female


40 2
Sigmoid displays/female

35
30 1.5
25
20 1
15
10 0.5
5
0 0
0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9

150
12
(c) * (d)

10
Interferences/female

100
Sneaks/female

50 4

0 0
0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9

400 5
(e) * 4.5 (f) *
350
4
300
Copulations/female

3.5
250
FFR/female

3
200 2.5

150 2
1.5
100
1
50 0.5
0 0
0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9

Operational sex ratio


Figure 2. Relationship between the operational sex ratio (males/(males+females)) and (a) the number of sigmoid displays per female, (b) the
total courtship time (min) per female, (c) the number of interference behaviours per female, (d) the number of sneak attempts per female, (e)
fraction of female responses (FFR) per female, and (f) the total number of copulation events per female. Each point represents the average of
female behaviour in each replicate. +indicates points with overlapping data. Linear or quadratic regressions were used; *P<0.001.

OSRs. On the other hand, at female-biased OSRs, there In general, male–male competition for mates among
were females available for males to court, reducing the guppies should increase at male-biased OSRs (Emlen &
need for sneaking behaviour. The plasticity in mating Oring 1977) as in other species (e.g. Lawrence 1986;
tactics with varying OSR may be an adaptive response of Enders 1993; Krupa & Sih 1993; Jormalainen et al. 1994;
males to a fluctuating sex ratio. Dick & Elwood 1996; Clutton-Brock et al. 1997). In
292 ANIMAL BEHAVIOUR, 58, 2

12 this may be an adaptive response by females to local


changes in the OSR.
10 Several studies have examined the effect of the OSR on
postcopulatory guarding (McLain 1980, 1989; Sillén-
8 Tullberg 1981; Clark 1988; Telford & Dangerfield 1990;
Female preference

Rowe 1992; Vepsäläinen & Savolainen 1995). They


6 showed that males guarded for longer at male-biased
OSRs. This reduced sperm competition by preventing the
4 females from remating prior to oviposition. Moreover,
postcopulatory mate guarding would also be expected to
2 be more important at more male-biased OSRs since it
would decrease the likelihood of sperm displacement,
0 which is another form of male–male competition (i.e.
interference after copulation). In my study, sigmoid
–2 displays that led to copulations occurred in all treat-
0.3 0.4 0.5 0.6 0.7 0.8 ments and the duration of postcopulatory mate guarding
Operational sex ratio and refraction were not significantly different across
Figure 3. Relationship between the operational sex ratio (males/ treatments. Surprisingly, sneak attempts that led to
(males+females)) and the slopes of FFR. Each point represents the copulations occurred only at 0.83 OSR. These sneak
slope of the regression of FFR on the orange area of males in each copulations happened very quickly; therefore, few sperm
replicate (i.e. the strength of female preference for orange colora- might have been transferred. When compared with copu-
tion). FFR was the ratio of male displays that elicited a female lations that followed sigmoid displays, successful sneak
response of 2 or more on a scale of 0–5 divided by the total number copulations were followed by shorter periods of postcopu-
of male displays. Linear regression was used: *P<0.05.
latory guarding and refraction. The mate-guarding period
may be short because it is more costly to guard than to
guppies, males interfered more with other males at male- find a new female. By contrast, copulations that followed
biased OSRs. Interference behaviours may result in higher sigmoid displays were longer, possibly increasing the
mating success for certain individuals in a population. For probability of successful sperm transfer. In this case, the
example, dominant male boat-tailed grackles, Quiscalus benefits of longer mate guarding may outweigh the ener-
major, increase their mating success by interfering with getic cost because the probability of fathering offspring is
other courting males (Poston 1997). Male guppies spent relatively high.
less total time courting females because their sigmoid Empirical studies of sexual selection commonly show
displays were frequently interrupted but these inter- that either female mate choice or male–male competition
rupted males stayed with the females for longer when drives the evolution of secondary sexual characteristics
given the opportunity. Thus at some OSRs, male–male (reviewed in Andersson 1994). In my experiments, the
interference may increase male mating success and play modes of sexual selection did not shift from active female
an important role in guppy sexual behaviour. choice to male–male competition with more male-biased
My study supports the hypothesis (Thornhill 1984; OSRs. In fact, both modes were present and the intensity
Lawrence 1986; Souroukis & Murray 1995) that the avail- of both modes increased as the OSR became male biased.
ability of mates influences both the sexual response and More studies that examine the effect of the presence of
mating strategy of females and the mating strategy of both modes of sexual selection are needed before conclu-
males. While each male receives fewer female sexual sions about their role in the evolution of secondary sexual
responses, on average each female responds more at characters can be made.
male-biased OSRs. In addition, females mated more often In conclusion, this study has demonstrated the relative
at higher OSRs. This indicates that when mate availability importance of different modes of sexual selection with
was high (i.e. at male-biased OSRs), females responded changing OSR. My findings show that at male-biased
more strongly to a courting male and were more willing OSRs, both modes of sexual selection are important
to mate. mechanisms generating sexual selection but only mate
Females should become more selective when OSR choice causes an increase in the rate of change of orange
increases because choosing a mate is less costly at male- coloration in males. However, the rate of change in
biased OSRs with more males from which to choose orange coloration in males may also depend on other
(Souroukis & Murray 1995). In this study, I was able to factors, such as gene flow, predation intensity, geographi-
measure the degree of female preference quantitatively by cal variation in female preference for colour patterns and
measuring the slopes of female response to orange area in density (Endler & Houde 1995).
males. I found that the degree of female preference for
orange colour increased at male-biased OSRs, which Acknowledgments
means that males with a higher percentage of orange area
received a stronger female sexual response, and suggests I thank John Endler, Armand Kuris, Robert Warner, Steve
that females became more selective. This indicates Rothstein, Cristina Sandoval, Gregory Grether, Molly
strongly that mate choice is not genetically fixed in this Cummings, Will McClintock, Chris Farmer, Sherie Zahn,
species but changes when the OSR changes. In nature, Lynda Holland, Robert Brooks, Ralph V. Park and two
JIROTKUL: OPERATIONAL SEX RATIO IN GUPPIES 293

anonymous referees for their useful comments on the Höglund, J. 1989. Pairing and spawning patterns in the common
manuscript. Many thanks to Krisanadej Jaroensutasinee toad, Bufo bufo: the effects of sex ratios and the time available for
for technical support. This study was supported in male–male competition. Animal Behaviour, 38, 423–429.
part by the Development and Promotion of Science and Houde, A. E. 1987. Mate choice based upon naturally occurring
colour pattern variation in a guppy population. Evolution, 41,
Technology Talents, Thailand, and by NSF grants DEB
1–10.
9317391 and 9508198. Houde, A. E. 1988. The effects of female choice and male–male
competition on the mating success of male guppies. Animal
Behaviour, 36, 888–896.
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