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Assortative mating by aggressiveness type in orb weaving spiders


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Animals within a population differ consistently in behavior over time and/or across conditions. A general question is how such differences referred to as personalities are maintained through evolution. One suggested mechanism is a nonrandom mate choice, which has been supported in species in which mate choice associates with direct material benefits. Much less is known about mating patterns and personality in species where males provide only sperm and in which the benefits of female choice are based only on good and/or compatible genes. The bridge spider Larinioides sclopetarius Clerck (Araneidae) exhibits heritable between-individual differences in intrasex aggressiveness. We studied mating probabilities by aggressiveness type of both sexes, and success in sperm competition of aggressive versus nonaggressive males. We staged trials that resemble field conditions: 4 males (2 aggressive and 2 nonaggressive) had simultaneous choice between an aggressive and a nonaggressive female. Although there were no differences in initial approaches of male types toward female types, aggressive males mainly mated with aggressive females, and nonaggressive males more likely mated with nonaggressive females. Female aggressiveness type was not related to fecundity, which may be a consequence of equal food supply in the laboratory. However, in double-mating trials using the sterile-male technique to measure paternity of aggressive versus nonaggressive males, we found that sons of aggressive parents fathered relatively more offspring. We conclude that assortative mating by aggressiveness type might maintain between-individual differences in aggressiveness in L. sclopetarius.
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Original Article
Assortative mating by aggressiveness type in
orb weaving spiders
SimonaKralj-Fišer,a,b Graciela A.Sanguino Mostajo,b OnnoPreik,b StanoPekár,c and Jutta
aInstitute of Biology, Scientific Research Centre, Slovenian Academy of Sciences and Arts, Novi trg
2, PO Box 306, SI-1001 Ljubljana, Slovenia, bBiozentrum Grindel, Zoological Institute & Museum,
University of Hamburg, Martin-Luther-King Platz 3, D-20146 Hamburg, Germany, and cDepartment of
Botany and Zoology, Faculty of Sciences, Masaryk University, Kotlá
ská 2, 611 37 Brno, Czech Republic
Animals within a population differ consistently in behavior over time and/or across conditions. Ageneral question is how such differ-
ences referred to as personalities are maintained through evolution. One suggested mechanism is a nonrandom mate choice, which
has been supported in species in which mate choice associates with direct material benefits. Much less is known about mating pat-
terns and personality in species where males provide only sperm and in which the benefits of female choice are based only on good
and/or compatible genes. The bridge spider Larinioides sclopetarius Clerck (Araneidae) exhibits heritable between-individual dif-
ferences in intrasex aggressiveness. We studied mating probabilities by aggressiveness type of both sexes, and success in sperm
competition of aggressive versus nonaggressive males. We staged trials that resemble field conditions: 4 males (2 aggressive and 2
nonaggressive) had simultaneous choice between an aggressive and a nonaggressive female. Although there were no differences in
initial approaches of male types toward female types, aggressive males mainly mated with aggressive females, and nonaggressive
males more likely mated with nonaggressive females. Female aggressiveness type was not related to fecundity, which may be a conse-
quence of equal food supply in the laboratory. However, in double-mating trials using the sterile-male technique to measure paternity of
aggressive versus nonaggressive males, we found that sons of aggressive parents fathered relatively more offspring. We conclude that
assortative mating by aggressiveness type might maintain between-individual differences in aggressiveness in L. sclopetarius.
Key words: aggression, competition, female choice, personality, reproductive success, sexual selection. [Behav Ecol]
Optimality approaches in behavioral ecology assume that selec-
tion favors the convergence of behavioral types/strategies toward
a single one with the highest fitness (Dall et al. 2004). Individual
dierences in behavior have been traditionally considered as noise
around an adaptive population mean. Variation in behavior was
interpreted as plasticity or alternative tactics (Gross 1996). Contrary
to this view, evidence is accumulating that individuals of the same
sex and age within a given population consistently dier from each
other in their behavioral characteristics referred to as personalities
(Zuckerman 1991).
This raises questions about why individual personality dierences
exist, that is, how they are generated and maintained (Sih et al.
2004; Schuett et al. 2010). A number of functional explanations
for the evolutionary maintenance of personalities have been sug-
gested. Between-individual behavioral dierences were proposed to
exist due to life-history trade-os (Stamps 2007; Wolf et al. 2007),
condition-dependent selection (Gross 1996), fluctuating selection,
or negative frequency–dependent selection (Dingemanse and Reale
2005; Smith and Blumstein 2008; Kralj-Fišer and Schneider 2012).
Within-individual behavioral consistency may be explained by
physiological constraints (Sih etal. 2004), benefits of predictabil-
ity (Dall etal. 2004; McNamara etal. 2009), and positive feedback
loops between state and behavior (Dall etal. 2004; Sih etal. 2004).
Recently, Schuett et al. (2010) suggested that personalities might
have evolved due to sexual selection probably acting together with
the other proposed evolutionary processes.
Sexual selection may maintain personalities through nonrandom
mate choice and male–male competition (Schuett et al. 2010).
Choosy individuals may gain benefits if potential mates vary in
fecundity or fertility, parental abilities, resources and status, “good
genes”, and/or genetic compatibility (Bateson 1983; Andersson
1994; Kokko et al. 2003; Ne and Pitcher 2005). A certain
personality type may be associated with quality as a mate while this
quality could be additive or nonadditive. In the former case, certain
personalities would be favored over others, whereas in the latter,
Address correspondence to S.Kralj-Fišer. E-mail:
Received 7 October 2012; revised 5 March 2013; accepted 16 March
Behavioral Ecology
Behavioral Ecology Advance Access published April 18, 2013
by guest on April 19, 2013 from
Behavioral Ecology
the match of the 2 partners matters. Depending on the connection
between personality type, and male and female fitness, the resulting
mating pattern could be assortative or disassortative.
Assortative mating by behavioral type may occur if the same
behavioral type indicates quality as a mate in both sexes. High-
quality individuals will then mate with each other, leaving individu-
als scoring low in this trait to pair up with one another (Johnson
1988; Davies 1989; McNamara and Collins 1990). If both extremes
on a behavioral axis, for example, shy and bold individuals, have
similar fitness, between-individual variability could be maintained
in such a way (Schuett et al. 2010). Assortative mating patterns
could also be produced if mates with compatible behavioral types
are preferred (Tregenza and Wedell 2000; Ne and Pitcher 2005)
although depending on the nature of the best match, compatibility
as a driving force of sexual selection can also produce disassorta-
tive mating patterns by behavioral type. Selecting a mate with the
same behavioral type can be favored by selection if this improves
coordination and cooperation within the pair and increases repro-
ductive success (Budaev etal. 1999; Both et al. 2005; Schuett and
Dall 2009; Royle et al. 2010; Schuett, Dall, etal. 2011; Schuett,
Godin, etal. 2011; Gabriel and Black 2012). On the other hand,
choosing a mate with another behavioral type could be adaptive if
this reduces the probability of genetic or behavioral incompatibili-
ties (Ens etal. 1993; Dingemanse etal. 2004; van Oers etal. 2008).
Studies that consider the role of personalities in sexual selection
have mostly been done with vertebrate systems in which males pro-
vide direct benefits to the female or ospring (reviewed in Schuett
etal. 2010); much less is known about the influence of sexual selec-
tion on personality in species where males provide only sperm (but
see Sinn etal. 2006) and in which benefits of female choice are sup-
posed to be based on good and/or compatible genes that increase
ospring fitness (Zeh and Zeh 1997; Kokko et al. 2003). Among
invertebrates, mate choice and personality have been studied in
the socially polymorphic spider Anelosimus studiosus (Hentz) (Pruitt
etal. 2008, 2011; Pruitt and Riechert 2009a, 2009b). In this spe-
cies, females are larger and the more dangerous sex, and aggres-
sive males preferentially choose to mate with social (docile) females,
whereas docile males more often mate with aggressive ones pro-
ducing a disassortative mating pattern (Pruitt and Riechert 2009a,
However, the reproductive success of a male with a particular
behavioral type may depend on the behavioral type of his mate.
In mating experiments with A.studiosus, aggressive males had lower
reproductive success when mated with aggressive (asocial) females
relative to docile males (Pruitt etal. 2011). It is possible that aggres-
sive females exert some downstream regulatory mechanism that
bias male insemination success; perhaps they do so in accordance
to genetic (in)compatibility, which seems to be associated with
aggressiveness type (Pruitt etal. 2011). Very few studies have dealt
with thisissue.
We aimed to study mating patterns in relation to behavioral
types in the bridge spider Larinioides sclopetarius Clerck (Araneidae),
an orb-weaver inhabiting areas across the Holarctic. Recent
experiments revealed heritable between-individual dierences in
aggressiveness toward a conspecific in both sexes, which were inde-
pendent of spider size (Kralj-Fišer and Schneider 2012). A simu-
lation of high-density conditions suggested that groups consisting
of a mixture of aggressive and nonaggressive types did better than
groups that consisted of nonaggressive types supporting the notion
that negative frequency–dependent selection could be relevant
(Kralj-Fišer and Schneider 2012). A male-biased operational sex
ratio and male–male competition for access to mates may select for
high male aggressiveness. We expected that aggressive males would
outcompete less aggressive rivals in access to preferred female phe-
notypes. Females mate multiply and vary in body mass (Kleinteich
and Schneider 2011), which is a good predictor of fecundity in spi-
ders in general (Bristowe 1958). Females are rarely sexually can-
nibalistic, and body size and aggressiveness of laboratory-reared
females are not correlated, so that we cannot make straightforward
predictions concerning male preference for female behavioraltypes.
Furthermore, we conducted a sperm competition study to test
the prediction that aggressive males may not only win precopu-
latory competition but inseminate more eggs as well. We sequen-
tially mated females with 2 males taken from the extremes of a
distribution of aggression scores. We measured paternity using a
sterile-male method and expected higher numbers of fertilized
eggs by the aggressive male regardless of mating order.
We used subadult spiders, kept in 200-mL plastic cups and fed
with Drosophila sp. until adulthood. We reared Drosophila larva
on a high-quality medium so that the adult flies contain all the
nutrients required for the spiders (Mayntz and Toft 2001). Spiders
grew and survived very well on this diet (Kleinteich and Schneider
2011). When adult, males remained in the 200-mL cups under
the same feeding treatment, whereas females were transferred into
plastic frames (h= 36, d =6, w = 36 cm). Adult female spiders
were fed with larger flies (Calliphora sp.). Spiders (26 males and 30
females) were fed twice per week, watered 5 days per week, and
kept at room temperature under light:dark cycle 14:10 h (early
autumn, breeding) conditions throughout the duration of the
Experimental protocol
Aggressiveness toward same sex conspecific
We tested both male and female aggressiveness by placing 2 indi-
viduals of the same sex close to each other. Generally the female
resident was found in 1 corner of the frame, the intruder was then
carefully placed with the paintbrush approximately 5 cm from the
resident. Females were tested twice, once as residents in their own
web and once as intruders on an unfamiliar web, and the order of
acting as intruder or resident was randomized.
Males give up web-building when adult. To observe males in
a competitive situation, 2 males were placed on a haphazardly
chosen female web. Males were placed in the center of the female
web, approximately 5 cm from each other (ca. 15 cm from a
female). In intrasex contests that lasted 20 min, aggressiveness was
measured as a sum of scores based on the frequency of aggressive
behavior such as approaching (score = 1), web-shaking (score =
1), attacking (score = 2), chasing (score = 3), and biting (score =
3) (Kralj-Fišer, Gregorič, etal. 2011; Kralj-Fišer, Schneider, et al.
2011). “Approach” was defined as a movement by one spider with
the result of shortening the distance to the other individual and
“chasing” as a quick move in the direction of the other individual
resulting in a successful attack or its escape.
According to the mean aggressiveness (average of sum of the
aggression scores from the 2 contests), test individuals were ranked
and divided into 2 groups: aggressive (higher than median) and
nonaggressive (lower than median). Aggressiveness scores showed a
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Kralj-Fišer etal. • Nonrandom mating by personality type
continuous distribution but we chose to simplify the variation into a
dichotomous treatment to facilitate data analysis.
Spiders were allocated into 15 groups of 6 individuals each for the
mating experiments. Each group consisted of 1 aggressive and 1
nonaggressive female (both virgin) and of 2 aggressive and 2 non-
aggressive males. We did not have enough males to use each one
only once (see Appendix 1). To account for a potential correlated
response due to repeated use of the same individuals, male ID was
included in the covariance structure of each linear model used to
analyze data.
Initially, the 2 females were housed in 2 frames separated by a
solid Perspex division (Figure1A). In the field, webs occur in close
proximity and are often not strictly separated from one another
but use common frame threads (Kleinteich 2009). In order to
simulate a realistic choice condition for the males, we simply
removed the division between the 2 frames several days before the
mating experiment so that we had a single large frame (h= 36,
d = 12, w = 36 cm) with 2 females and their webs (Figure 1B).
To minimize the risk of aggressive behavior or even cannibalism
due to hunger, we fed females a day prior to experiment. Prior
to each experiment, we weighed all spiders using an electronic
balance (Mettler Toledo XS105 Dual Range) to an accuracy of
0.01 mg. We marked all spider males and females individually
using nontoxic paint and released the 4 marked males into the
frame (Figure1C), making sure they touched both webs, so they
perceived the presence of the 2 females.
All male–female interactions including aggressiveness, court-
ship, and copulation were recorded for 2 h. Female aggression
toward males was quantified based on the same scoring system as
the aggressiveness test (see above). The exception was biting the
male’s tarsus, which seems to be a part of the courtship or an indi-
cator of the female willingness to mate as it generally occurs during
courtship and never induces termination of courtship or aggressive
behavior (Kralj-Fišer S, personal observation). Therefore, it was not
considered as indicator of mating aggressiveness.
Courtship and copulation were observed and timed with a stop-
watch. To measure the duration of copulation, the stopwatch was
started at genital contact and stopped when genitalia were discon-
nected. Courtship starts with the male entering the web, signaling
his presence to the female. The female either stays passive or comes
out of her retreat (unless she acts aggressively), and the male will
start touching her body with his legs. Courtship ends with copula-
tion or the male leaving the web. We recorded the number (N) of
approaches (meaning the male moved toward the female on the web
without touching her) and touches (the male touched the female
with his legs) from each male toward each female. Copulation was
observed for frequency and order for each male. Very short appar-
ent copulations (less than 1 s) were recorded; however, their validity
as true copulations with sperm transfer were only later confirmed
when we were able to assess whether they resulted in fertilization of
eggs. This was determined by inspecting the incubated egg sacs for
hatching success.
After mating, females were kept individually and fed as before
until they had produced 2 egg sacs. Each egg sac was placed in a
controlled climate chamber at 25°C and light:dark cycle of 14:10
h and was misted with water on 5days per week. One week after
the first spiderlings had hatched, the clutches were frozen at −80°C
and then transferred into 70% ethanol. Numbers of hatched o-
spring were counted under the microscope.
After the experiments had ended, the tibia + patella length of
the first leg from all the tested individuals was measured to an accu-
racy of 0.01 mm using a LEICA MZ 16 stereomicroscope and the
corresponding computer software.
We used unrelated F1 spiders derived from a heritability study,
hence with known heritage of aggressiveness type (Kralj-Fišer and
Schneider 2012). To standardize the contribution of the female
aggression level to paternity, females were daughters of parents
taken from the middle of the distribution of aggressiveness scores,
whereas males were sons of parents with high or low aggressiveness
scores, hence taken from the extremes only. We refer to sons of
aggressive parents as aggressive males (Agg) and to nonaggressive
males (Non) for sons of nonaggressive parents. Half of the males
from each group were randomly picked and irradiated for 50 min
with a dosage of 40 Gray (0.8 Gy/min). The irradiation causes
DNA damage, and although the sperm is still capable of fertilizing
eggs, these eggs will not develop (Schneider and Andrade 2011).
Fifteen females received an irradiated nonaggressive males first
and an untreated aggressive male second (Non(I)/Agg(N)) and
15 females received an irradiated aggressive male first and an
untreated nonaggressive male second (Agg(I)/Non(N)). Hence,
all first males were irradiated. We did not vary the order of the
irradiation as we were interested in the relative paternity; the
absolute fertilization success depending on the irradiation was
irrelevant. We avoided systematic errors due to the irradiation
treatment by treating both groups of males. Two cases had to be
Page 3 of 8
Experimental setup to observe for mating pattern between aggressive and
nonaggressive females and between 2 aggressive and 2 nonaggressive males.
by guest on April 19, 2013 from
Behavioral Ecology
excluded: one because the first male was not prevented from getting
2 insertions and one extreme outlier where the female was very
close to oviposition so that the sperm of the 2nd male could not
be used. The exclusions reduced sample sizes to 14 in the Agg(I)/
Non(N) and 15 in the Non(I)/Agg(N) treatment. The 2 groups of
females did not dier in weight (Anova, F1,27 = 0.02, P=0.88) or
in the clutch size of their 1st (Anova, F1,27 = 1.64, P=0.21) and
2nd egg sacs (Anova, F1,25=0.31, P=0.58). Average clutch size in
the Agg(I)/Non(N) was 115.6 (standard error [SE]= 9.35) in the
1st and 128.5 (SE=8.04) in the 2nd clutch. In the Non(I)/Agg(N)
treatment, 1st clutches contained 99.0 (SE=7.72) and 2nd clutches
contained 109.3 (SE = 12.01) eggs. First and second males did not
dier in body mass or in body size measured as tibia–patella length
(Anovas, all P values >0.68).
The 2 males that mated with the same female were carefully
size matched and a correlation between the weight of the 2 males
and their size was highly significant (weight: Pearson correlation
r=0.95, df=25, P=0.0001; size: r=0.71, df=25, P=0.0001).
However, although the weight match was very good, a dierence
in size was still present as well as a large variation between trials
so that male body size was entered as a covariate into the analysis.
In addition to the above treatments, an irradiation control
treatment was performed in which 5 females mated with 2 irra-
diated males. None of these females produced hatchlings show-
ing that the irradiation was successful. We know from numerous
breeding experiments with this species that natural hatching
success is very high (Kleinteich 2009).
Each male was allowed a single intromission and was then
removed. Copulation duration was measured using a stopwatch.
Females were then kept until they had produced 2 egg sacs. Each
egg sac was left to hatch in a controlled climate chamber (as
above). One week after the first spiderlings had hatched, the egg
sacs were frozen at −80°C and then transferred into 70% ethanol.
Hatched and unhatched eggs were counted under the micro-
scope. Each clutch was counted twice on 2 dierent days without
knowing from which treatment they had derived.
Statistical analyses
The eect of 6 predictors, female patella + tibia I length, male
patella + tibia I length, female age, male age, female behavioral
type, and male behavioral type, on the mating probability, num-
ber of mates, first male choice, copulation frequencies, durations
of copulation, number of approaches, number of touches, dura-
tions of courtship, female aggression scores toward males, egg sac
numbers, egg sac weight, and hatchling numbers was studied using
generalized estimating equations (GEE) or generalized linear mixed
models (GLMM) because we used a block design with repeated use
of the same individuals. GEE were used when only 1 “random”
variable (group ID) was in the model and because we were not
interested in the estimation of variance of the random eects. GEE
models the correlation among subjects, resulting from repeated
use of the same individuals, in the covariance matrix of residuals
and provides correct marginal or population average models even
when the correlation structure is not perfectly specified (Hardin
and Hilbe 2003). GLMM was used in situations when 2 crossed
variables considered “random” (group and male ID) were in the
model, though we were not particularly interested in the estima-
tion of variation for the random eects. The random eects of the
male ID were significant (P< 0.05) in analyses when the follow-
ing response variables were used: copulation frequencies, number
of approaches, number of touches, and female aggression scores
toward males. GLMM models the correlation among subjects in
the covariance matrix of random eects (Zuur etal. 2009). We kept
both “random” variables in the model and simplified only the fixed
part of the linear predictor using stepwise deletion of nonsignifi-
cant eects, which included all predictors in an additive way. The
Poisson error structure (GEE-p, GLMM-p) was chosen when the
response variables were counts; binomial errors were used (GEE-b,
GLMM-b) when the response was of binary character. Alog-nor-
mal model (GEE, GLMM) or gamma errors (GEE-g, GLMM-g)
were assumed when the response variable was time due to increase
of variance with themean.
Generalized linear models with binomial error structure (GLM-
b) were used to study the relationship between several predictors
(female weight, copulation duration, male weight, male aggres-
sion, and male size) and hatching success. Only eggs fathered by
the untreated of the 2 males hatched, whereas the eggs fathered by
the irradiated male died during early development. Hence hatching
success equals paternity success of the untreated male. Data for 2
consecutive egg sacs were pooled and used as a measure of hatch-
ing success. The linear predictor included main eects and their
2-way interactions.
All analyses were performed within R (R Development Core
Team 2010). For the GEE, we used the geepack package (Yan
and Fine 2004) and for the GLMM, the lme4 package (Bates and
Maechler 2009).
Aggressive and nonaggressive males did not dier significantly in
the frequencies of approaches to females in general, regardless
of female aggression type (GLMM-p, X1
P=0.73), in the
number of touches of females (GLMM-p, X1
P= 0.26), or
in courtship durations (GLMM, X1
P=0.29). The approach
frequency (GLMM-p, X1
213 8=
P = 0.0002) and number of
touches (GLMM-p, X1
P = 0.04) decreased with female
age, whereas the eects of all other explanatory variables (female
size, male size, and male age) were not significant (GLMM-p,
P >0.1).
Aggression levels of the male and the female were not important
for the first approach as evidenced by the nonsignificant interaction
(GEE-b, X1
P=1.00). Including all approaches, aggressive
males approached aggressive females almost twice as often as
nonaggressive females, whereas nonaggressive males slightly more
often approached nonaggressive females (GLMM-p, X1
237 1=
P < 0.0001, Figure 2A). Furthermore, aggressive males touched
aggressive females significantly (almost 3 times) more often
(GLMM-p, X1
219 8=
P < 0.0001, Figure2B) and courted them
significantly (3 times) longer (GLMM, X1
P = 0.005,
Figure 2C) than nonaggressive males. The dierence was less
pronounced but reversed in nonaggressive females, which were
more often touched and courted for longer by nonaggressivemales.
Aggressive females were 1.5 times more aggressive toward males
than nonaggressive females. Female aggressiveness decreased with
her size (GLMM-p, X1
P=0.03). Aggressive females were
twice as aggressive toward aggressive males than toward nonaggres-
sive males, whereas nonaggressive females had higher aggression
frequencies toward nonaggressive males (GLMM-p, X1
241 6=
P < 0.0001, Figure 3). Aggressive and nonaggressive males over-
all did not receive significantly dierent levels of aggression from
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Kralj-Fišer etal. • Nonrandom mating by personality type
females (GLMM-p, X1
P = 0.36). Independent of female
and male behavioral types, female aggressiveness toward males
increased with female age (GLMM-p, X1
Aggressive and nonaggressive females did not dier significantly
in the frequencies of mating (GLMM-g, X1
P = 0.92,
Table1), in total number of males they copulated with (GLMM-p,
P = 0.78), or in durations of mating (GLMM-p,
P = 0.4). Aggressive and nonaggressive males did
not dier significantly in the occurrences of mating (GLMM-b,
P=0.54, Table1), in the durations of mating (GLMM,
P = 0.82), in the frequencies of mating (GLMM-b,
P=0.41), and in the number of females they copulate
with (GLMM-p, X1
However, we detected a significant interaction between female
and male aggressiveness type on the probability of mating
(GLMM-b, X1
P = 0.03). Indeed, the mating probability
of aggressive females with aggressive males was more than 3 times
higher than the probability of mating with nonaggressive males
(Figure4A). Accordingly, nonaggressive females significantly more
likely mated (1.45 times) with nonaggressive males (GLMM-p,
219 5=
P < 0.0001, Figure4B). Mating durations were not sig-
nificantly aected by the aggressiveness of either males or females
P = 0.67). Mating frequency decreased with
female age (GLMM-p, X1
219 1=
P < 0.0001).
Reproductive success
Nonaggressive and aggressive females did not dier significantly in
the number of egg sacs (GEE-p, X1
P=0.47), average egg
sac mass (GEE, X1
P=0.29) and average number of hatch-
lings (GEE-p, X1
P = 0.71, Table 1). Number of hatch-
lings decreased with female age (GEE-p, X1
P=0.02) and
increased with female size (GEE-p, X1
227 4=
Paternity success
The proportion of eggs sired by the untreated male (hatched
eggs) was not related to female weight (GLM-b, F1,24 = 0.1,
P = 0.96), copulation duration (GLM-b, F1,24 = 2.7, P = 0.11),
and male weight (GLM-b, F1,24= 3.4, P= 0.08). Nevertheless, it
was positively related to body size of males (GLM-b, F1,24= 7.5,
P = 0.011). Aggressive males, however, sired significantly higher
proportions of eggs than nonaggressive males (GLM-b, F1,25=6.4,
P=0.02, Figure5).
Our study shows that mating between male and female bridge spi-
ders is not random in respect to their behavioral types: aggressive
males more likely mated with aggressive females, whereas non-
aggressive males more likely mated with nonaggressive females
producing a pattern of assortative mating. We propose that non-
random mating by behavioral type perhaps in combination with
negative frequency–dependent selection could maintain between-
individual variability in aggressiveness in L.sclopetarius.
This is the second study to investigate mating patterns by aggres-
siveness type in a spider. In the cooperative breeding spider A. stu-
diosus, docile females more often mate with aggressive males and
vice versa; hence, in contrast to the assortative pattern in the bridge
spiders, these spiders mate disassortatively (Pruitt and Riechert
Page 5 of 8
Aggressive M
Nonaggressive M
Duration [s]
Aggressive M
Nonaggressive M
Aggressive M
Nonaggressive M
Aggressive Nonaggressive
Aggressive Nonaggressive
Comparison of courtship behavior between aggressive and nonaggressive
males toward aggressive and nonaggressive females. Comparison of (A)
the number of approaches, (B) number of touches, and (C) duration of
courtship. Bars are means and whiskers are SEs.
by guest on April 19, 2013 from
Behavioral Ecology
2009a, 2009b; Pruitt etal. 2011). There are several possible expla-
nations for the dierences. Anelosimus studiosus spiders exhibit high
sexual size dimorphism, where larger females readily consume a
male prior to copulation; hence males might be selected to avoid
mating with aggressive females. In this context, docile male spiders
perhaps have better dispositions to perform sneaking behavior or
may invest more in reducing female incentive to attack and are
more likely to succeed in mating with aggressive females. In our
study species, selection to avoid aggressive females may be relatively
weak, as female aggression hardly ever results in sexual cannibalism
and males are not much smaller than females and perfectly able
to resist female attacks. Apositive correlation between sexual size
dimorphism and sexual cannibalism is a general pattern among spi-
ders (Wilder and Rypstra 2008). Furthermore, A.studiosus are social
spiders and males in general prefer to mate with social females
(Pruitt and Riechert 2009a). However, in our study species, it is less
clear which female behavioral type (if only one) is preferred. We
speculated that males prefer aggressive females in the field because
these are likely more fecund (see below).
Although female aggressiveness types did not dier in their
reproductive success when mating with the male of the same
behavioral type, aggressive males sired more ospring than non-
aggressive ones under sperm competition when females were of
medium behavioral type. Because we used sons of parents with
known aggressiveness type, which is heritable (Kralj-Fišer and
Schneider 2012), aggressive males appear to father sons that are
better in sperm competition at least if mated to aggressive females.
If this is a heritable male property, all females should preferentially
copulate with aggressive males. However, nonaggressive females
mated more frequently with nonaggressive males. There are 2
nonmutually exclusive explanations. First, nonaggressive females
do not produce better sperm competitors with aggressive sires,
for example, because the combination of maternal and paternal
behavioral type is relevant for the inheritance of male aggressive-
ness type. In fact, L. sclopetarius and A. studiosus ospring exhibit
aggression levels similar to average parental aggression; hence
high aggressiveness of both parents is likely needed for inheritance
of high aggressiveness (Kralj-Fišer and Schneider 2012). Also, it
would be important to test if aggressive males also win in sperm
competition with nonaggressive males when mated with a docile
Page 6 of 8
Aggressive M
Nonaggressive M
Aggressive Nonaggressive
Comparison of the aggressive scores of aggressive and nonaggressive
females toward aggressive and nonaggressive males. Bars are means and
whiskers are SEs.
Mean and SE of mating and reproductive parameters of
aggressive and nonaggressive females
Aggressive Nonaggressive
No. of matings 2.7 (0.9) 3.7 (1.1)
No. of males mated with 1.0 (0.3) 1.1 (0.2)
Total duration of copulation (s) 22.39 (8.8) 20.69 (4.9)
Egg sac mass 0.11 (0.03) 0.10 (0.03)
No. of hatchlings 46.7 (19.2) 19.7 (17.7)
Mating probability
Aggressive M
Nonaggressive M
Aggressive Nonaggressive
Aggressive M
Nonaggressive M
Aggressive Nonaggressive
Comparison between aggressive and nonaggressive females and between
aggressive and nonaggressive males for (A) the mating probability and (B)
frequency of matings. Bars are means and whiskers are SEs.
by guest on April 19, 2013 from
Kralj-Fišer etal. • Nonrandom mating by personality type
The second possible explanation is that aggressive males
actively avoid mating with nonaggressive females as long as there
are alternatives. The reasons would be the same as above namely
that assortative matings create phenotypes with the highest
quality so that they father better sons only in combination with
aggressive females. Male choosiness could be facilitated if they
were sperm limited and were unable to mate with many females.
We have not specifically studied potential mating frequencies
in bridge spiders but past mating trials provided no evidence
for such a limitation. The presence of male mating preferences
would also explain their increased investment in courtship and
their persistence with aggressive females. These options have to
be further explored.
Aggressive and nonaggressive females did not dier signifi-
cantly in their reproductive success in our study, although there is
a marked (but not significant) dierence in the number of hatch-
lings (Table 1). There was a large variation and small sample
size so that we cannot base any firm conclusion on these data.
Dierences in female reproductive success between aggressive-
ness types may have been masked by the rearing conditions in
the laboratory, where females were kept and fed individually and
with the same quantity and quality of prey. Aggressive females in
predatory species are often reported to be more vigorous hunt-
ers resulting in higher prey capture success, improved growth
rates, and larger body size at a better condition (Bristowe 1958;
Arnqvist and Henriksson 1997). All these parameters are directly
linked to increased fecundity in invertebrates, including spiders
and our study species (this study; Kleinteich 2009). It should be
further explored if female reproductive success between aggres-
sive types remains similar under more natural conditions or
whether aggressiveness is favored through increased hunting suc-
cess. Bridge spiders are nocturnal hunters of small prey insects
that emerge from the water. In urban habitats, they gather around
light sources and compete for the best web sites (Heiling 1999).
Being aggressive may on the one hand improve chances of gain-
ing a rich site but may also be disadvantageous if foraging is
compromised by frequent territorial encounters. Arecent experi-
ment with high densities of females and competition over prey
suggests that groups of only aggressive females have lower mor-
tality than mixed groups (Kralj-Fišer and Schneider 2012). Field
studies are currently under way to address the natural composi-
tion of behavioral types in high-density groups.
The maintenance of alternative phenotypes is often promoted
by frequency-dependent advantages or by trade-os such as high
costs of aggressiveness through injuries or cannibalism. In A. stu-
diosus, docile females have an advantage in high densities and meek
males profit when frequencies of aggressive males are high (Pruitt
and Riechert 2011). In L.sclopetarious, there is evidence that nonag-
gressive behavioral types have advantages if aggressive individuals
are too frequent at profitable web sites (Kralj-Fišer and Schneider
2012) but more studies are required to assess whether such fre-
quency-dependent mechanisms in combination with assortative
mating patterns are sucient to explain the maintenance of consis-
tent dierences in aggressivenesstypes.
In summary, L. sclopetarius spiders mated assortatively by their
aggressive types. Male aggressiveness level was positively related to
the number of sired ospring and likely signaled high quality in
L. sclopetarius males. Female reproductive success was not directly
related to aggressiveness type in this study, but positively correlated
to female size. Our studies suggest that assortative mating pattern
by aggressiveness type might be relevant for maintaining between-
individual dierences in aggressiveness at least in L.sclopetarius (this
study, Kralj-Fišer and Schneider 2012).
S.K.-F. was granted by Humboldt Postdoctoral Fellowship,
Humboldt Return Fellowship, and ARRS Postdoctoral Fellowship
We thank Tomma Dirks and Angelika Tabel-Hellwig for excellent spider
husbandry, and Wiebke Schuett for their comments on the manuscript. We
thank Cene Fišer for drawing Figure 1.
Handling editor: Ben Hatchwell
Page 7 of 8
6.06.5 7.0
0.0 0.2 0.4 0.6 0.8 1.0
Male size [mm]
Hatching success
Aggressive M
Non-aggressive M
Relationship between the male size (patella + tibia length of the first leg)
and the hatching success (i.e., the proportion of eggs hatched) of 2 egg sacs
(pooled) for aggressive and nonaggressive males. Logit models are displayed.
Appendix 1
Males from the 15 mating trials
Group ID
Male ID
Male 1 Male 2 Male 3 Male 4
1 17 23 21 22
2 4 30 28 13
3 6 16 12 26
4 7 10 26 12
5 15 19 29 14
6 15 19 14 29
7 6 16 3 11
8 2 20 25 18
9 23 9 28 13
10 19 15 29 3
11 2 10 22 11
12 2 20 21 22
13 24 4 28 25
14 7 30 26 12
15 9 20 11 3
by guest on April 19, 2013 from
Behavioral Ecology
Andersson M. 1994. Sexual selection. Princeton: Princeton University
Arnqvist G, Henriksson S. 1997. Sexual cannibalism in the fishing spider
and a model for the evolution of sexual cannibalism based on genetic
constraints. Evol Ecol. 11:255–273.
Bates D, Maechler M. 2009. lme4: linear mixed-eects models using S4
classes. R package version 0.999375-31 [Last cited 30 October 2010].
Available from:
Bateson PPG. 1983. Mate choice. Cambridge: Cambridge University
Both C, Dingemanse NJ, Drent PJ, Tinbergen JM. 2005. Pairs of extreme
avian personalities have highest reproductive success. J Anim Ecol.
Bristowe WS. 1958. The world of spiders. London: Collins.
Budaev SV, Zworykin DD, Mochek AD. 1999. Individual dierences in
parental care and behaviour profile in the convict cichlid: a correlation
study. Anim Behav. 58:195–202.
Dall SRX, Houston AI, McNamara JM. 2004. The behavioural ecology of
personality: consistent individual dierences from an adaptive perspec-
tive. Ecol Lett. 7:734–739.
Davies NB. 1989. Sexual conflict and the polygamy threshold. Anim Behav.
Dingemanse NJ, Both C, Drent PJ, Tinbergen JM. 2004. Fitness conse-
quences of avian personalities in a fluctuating environment. Proc R Soc
Lond B. 271:847–852.
Dingemanse NJ, Reale D. 2005. Natural selection and animal personality.
Behaviour. 142:1159–1184.
Ens BJ, Safriel UN, Harris MP. 1993. Divorce in the long lived and monog-
amous oystercatcher, Haematopus ostralegus: incompatibility or choosing the
better option? Anim Behav. 45:1199–1217.
Gabriel PO, Black JM. 2012. Behavioural syndromes, partner com-
patibility and reproductive performance in Steller’s jays. Ethology.
Gross MR. 1996. Alternative reproductive strategies and tactics: diversity
within sexes. Trends Ecol Evol. 11:92–98.
Hardin JW, Hilbe JM. 2003. Generalized estimating equations. Boca Raton:
Chapman & Hall/CRC.
Heiling AM. 1999. Why do nocturnal orb-web spiders (Araneidae) search
for light? Behav Ecol Sociobiol. 46:43–49.
Johnson K. 1988. Sexual selection in pinyon jays I: female choice and male-
male competition. Anim Behav. 36:1038–1047.
Kleinteich A. 2009. Life-history of the bridge spider, Larinioides sclopetarius
(Clerck, 1757) [PhD thesis]. [Germany]: University of Hamburg.
Kleinteich A, Schneider JM. 2011. Developmental strategies in an invasive
spider: constraints and plasticity. Ecol Entomol. 36:82–93.
Kokko H, Brooks R, Jennions MD, Morley J. 2003. The evolution of mate
choice and mating biases. Proc R Soc Lond B. 270:653–664.
Kralj-Fišer S, Gregorič M, Zhang S, Li D, Kuntner M. 2011. Eunuchs are
better fighters. Anim Behav. 81:933–939.
Kralj-Fišer S, Schneider JM. 2012. Individual behavioural consistency and
plasticity in an urban spider. Anim Behav. 84:197–204.
Kralj-Fišer S, Schneider JM, Justinek Ž, Kalin S, Gregorič M, Pekár S,
Kuntner M. 2011. Mate quality, not aggressive spillover, explains sexual
cannibalism in a size dimorphic spider. Behav Ecol Sociobiol. 66:145–151.
Mayntz D, Toft S. 2001. Nutrient composition of the prey’s diet
aects growth and survivorship of a generalist predator. Oecologia.
McNamara JM, Collins EJ. 1990. The job search problem as an employer
candidate game. J Appl Probab. 27:815–827.
McNamara JM, Stephens PA, Dall SRX, Houston AI. 2009. Evolution of
trust and trustworthiness: social awareness favours personality dierences.
Proc R Soc Lond B. 276:605–613.
Ne BD, Pitcher TE. 2005. Genetic quality and sexual selection: an inte-
grated framework for good genes and compatible genes. Mol Ecol.
van Oers K, Drent PJ, Dingemanse NJ, Kempenaers B. 2008. Personality is
associated with extrapair paternity in great tits, Parus major. Anim Behav.
Pruitt JN, Riechert SE. 2009a. Male mating preference is associated with
risk of pre-copulatory cannibalism in a socially polymorphic spider.
Behav Ecol Sociobiol. 63:1573–1580.
Pruitt JN, Riechert SE. 2009b. Sex matters: sexually dimorphic fitness con-
sequences of a behavioural syndrome. Anim Behav. 78:175–181.
Pruitt JN, Riechert SE. 2011. How within-group behavioural variation and
task eciency enhance fitness in a social group. Proc R Soc Lond B.
Pruitt JN, Riechert SE, Harris DJ. 2011. Reproductive consequences of
male body mass and aggressiveness depend on females’ behavioral types.
Behav Ecol Sociobiol. 65:1957–1966.
Pruitt JN, Riechert SE, Jones TC. 2008. Behavioural syndromes and their
fitness consequences in a socially polymorphic spider, Anelosimus studiosus.
Anim Behav. 76:871–879.
R Development Core Team. 2010. R: a language and environment for
statistical computing. Vienna (Austria): R Foundation for Statistical
Computing [cited 10 April 2013]. Available from: http://www.R-project.
Royle NJ, Schuett W, Dall SRX. 2010. Behavioral consistency and the
resolution of sexual conflict over parental investment. Behav Ecol.
Schneider JM, Andrade MCD. 2011. Mating behaviour & sexual selection.
In: Herberstein M, editor. Spider behaviour: variability and versatility.
Cambridge (UK): Cambridge University Press.
Schuett W, Dall SRX. 2009. Sex dierences, social context and personality
in zebra finches, Taeniopygia guttata. Anim Behav. 77:1041–1050.
Schuett W, Dall SRX, Royle NJ. 2011. Pairs of zebra finches with similar
‘personalities’ make better parents. Anim Behav. 81:609–618.
Schuett W, Godin JGJ, Dall SRX. 2011. Do female zebra finches, Taeniopygia
guttata, choose their mates based on their ‘personality’? Ethology.
Schuett W, Tregenza T, Dall SRX. 2010. Sexual selection and animal per-
sonality. Biol Rev. 85:217–246.
Sih A, Bell AM, Johnson JC. 2004. Behavioral syndromes: an ecological
and evolutionary overview. Trends Ecol Evol. 19:372–378.
Sinn DL, Apiolaza LA, Moltschaniwskyj NA. 2006. Heritability and
fitness-related consequences of squid personality traits. J Evol Biol.
Smith BR, Blumstein DT. 2008. Fitness consequences of personality: a
meta-analysis. Behav Ecol. 19:448–455.
Stamps JA. 2007. Growth-mortality tradeos and ‘personality traits’ in ani-
mals. Ecol Lett. 10:355–363.
Tregenza T, Wedell N. 2000. Genetic compatibility, mate choice and pat-
terns of parentage: invited review. Mol Ecol. 9:1013–1027.
Wilder SM, Rypstra AL. 2008. Sexual size dimorphism predicts the fre-
quency of sexual cannibalism within and among species of spiders. Am
Nat. 172:431–440.
Wolf M, van Doorn GS, Leimar O, Weissing FJ. 2007. Life-history
trade-os favour the evolution of animal personalities. Nature.
Yan J, Fine JP. 2004. Estimating equations for association structures. Stat
Med. 23:859–880.
Zeh JA, Zeh DW. 1997. The evolution of polyandry 2. Post-copulatory
defences against genetic incompatibility. Proc R Soc Lond B. 264:69–75.
Zuckerman M. 1991. Psychobiology of personality. Cambridge: Cambridge
University Press.
Zuur A, Ieno EN, Walker N, Saveliev AA, Smith GM. 2009. Mixed eects
models and extensions in ecology with R. Berlin: Springer.
Page 8 of 8
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... Studies on the relationship between behavioural phenotype and reproductive success raise questions regarding the prevalence of assortative (or disassortative) mating (Schuett et al., 2010) on the character or characters that define the phenotype. Assortative mating may be a proximate mechanism for the populationwide maintenance of individual variation in phenotype (Kralj-Fi ser et al., 2013) and itself may be proximately driven by a high probability of encountering a mate of similar phenotype, as may be expected for phenotypes that are based on exploratory behaviour, risk aversion and neophobia. Alternatively, assortative mating may be proximately driven by mating preferences for traits correlated with the phenotype. ...
... In orb-weaving spiders (Larinioides sclopetarius), sons of aggressive parents sire more offspring than sons of nonaggressive parents, a phenomenon that apparently promotes assortative mating (Kralj-Fi ser et al., 2013). In bighorn rams, Ovis canadensis, docile and bold rams show higher survival and higher reproductive success than nondocile and shy rams (R eale et al., 2009). ...
Individually consistent behavioural phenotypes persist in a diversity of populations, despite the expectation that selection would reduce their prevalence. The life-history trade-off between current and future reproduction, combined with individual variation in age or condition and therefore in residual reproductive value, may explain this apparent paradox. Specifically, individuals that are old or of low condition and therefore of low residual reproductive value should take risks and thus elevate current reproduction at the expense of future reproduction. The opposite should be true for individuals that are young or of high condition and therefore of high residual reproductive value, which should be risk averse. Over the course of a long-term study on Lincoln's sparrows, Melospiza lincolnii, we discovered a behavioural phenotype based on trappability that predicts both condition (but not age) and current reproductive success but, surprisingly, in a way that was not expected based on our assumption of how this phenotype is associated with risk taking. We found that, relative to individuals that did not enter traps (the nontrap phenotype, assumed as risk averse), those of the trap phenotype (assumed as risk taking) showed elevated indices of energetic condition and, for females, reduced reproductive success. Assortative mating may be a proximate mechanism for the populationwide maintenance of multiple behavioural phenotypes, and we found that assortative pairings based on trappability phenotype occurred more frequently than random pairing would predict. However, assortative pairing, when compared to dis-assortative pairing, did not affect reproductive success. Nevertheless, the contrasting relationships between current reproductive success and condition that we found for each phenotype are consistent with life-history theory. However, our results are not consistent with the assumption that the trap phenotype is a manifestation of risk-taking behaviour and may, in fact, arise from just the opposite: a risk-averse, shy or neophobic behaviour for the trap phenotype.
... Arthropods are an abundant and taxonomically diverse group of organisms for which mate choice has been observed in multiple taxa, including insects [10,17], arachnids [18,19] and crustaceans [20]. In this taxon, body size is considered to be one of the most prevalent choice criteria [10,21,22]. ...
Full-text available
Background Mate choice is a fundamental element of sexual selection and has the potential to shape the evolution of traits. Mate choice based on body size has been shown to be a common trait in several arthropod species. In hard ticks, a taxon of medical and veterinary importance, engorgement weight is positively correlated with reproductive output but it is unknown whether adult males show mate choice. Here, we experimentally investigated whether males (i) use chemical cues to choose their mating partner, (ii) consistently choose for the same female individual and (iii) prefer females with highest weight after feeding. Methods We used two experimental setups which allowed chemical communication between ticks: (i) a horizontal tube preventing physical contact with the female and (ii) an arena where tactile cues were allowed. In total, we tested 62 different triads in 124 tests (66 tests in the horizontal tube and 58 in the arena) composed of one male that could choose between two engorged females. Specifically, we tested 42 triads in the tube and 46 in the arena; 24 triads were repeatedly tested in the tube while 38 triads were tested in both setups. Results We found no preference for individual or heavier females in either setup. However, in the horizontal tube setup, males significantly preferred females that were not visited by them in the previous test. Conclusions Our results suggest a lack of male mate choice despite heavier females having higher fecundity. However, future studies should take into account that males may recognize the potential mating partners they previously met. Graphical Abstract
... Arthropods are an abundant and taxonomically diverse group of organisms where mate choice has been observed in multiple taxa including insects (Arnqvist et al. 1996;Bonduriansky 2001), arachnids (Bel-Venner et al. 2008;Kralj-Fišer et al. 2013) and crustaceans (Fazhan et al. 2017). In this taxon, body size is considered to be one of the most prevalent choice criteria (Bonduriansky 2001;Crespi 1989;Reading and Backwell 2007 between species. ...
... intriguing role in many taxa. Depending on a species' respective environmental conditions, assortative mate choice of similarly behaving individuals or disassortative mate choice of apparently antagonistic, but complementary behaving individuals may be favored (mammals: e.g., Ihara and Feldman, 2003;Massen and Koski, 2014;Rangassamy et al., 2015;Martin-Wintle et al., 2017; birds: e.g., Both et al., 2005;van Oers et al., 2008;Schuett et al., 2011;Gabriel and Black, 2012;Horton et al., 2012;Fox and Millam, 2014;Pogány et al., 2018;Clermont et al., 2019;Collins et al., 2019; fish: e.g., Ariyomo and Watt, 2013;Laubu et al., 2017;Scherer et al., 2017;Schweitzer et al., 2017;Chen et al., 2018; invertebrates: e.g., Kralj-Fišer et al., 2013;Montiglio et al., 2016;Baur et al., 2019). This could be a decisive competitive advantage both genotypically and phenotypically in the context of bi-parental brood care. ...
Full-text available
The idea of “smart is sexy,” meaning superior cognition provides competitive benefits in mate choice and, therefore, evolutionary advantages in terms of reproductive fitness, is both exciting and captivating. Cognitively flexible individuals perceive and adapt more dynamically to (unpredictable) environmental changes. The sex roles that females and males adopt within their populations can vary greatly in response to the prevalent mating system. Based on how cognition determines these grossly divergent sex roles, different selection pressures could possibly shape the (progressive) evolution of cognitive abilities, suggesting the potential to induce sexual dimorphisms in superior cognitive abilities. Associations between an individual’s mating success, sexual traits and its cognitive abilities have been found consistently across vertebrate species and taxa, providing evidence that sexual selection may well shape the supporting cognitive prerequisites. Yet, while superior cognitive abilities provide benefits such as higher feeding success, improved antipredator behavior, or more favorable mate choice, they also claim costs such as higher energy levels and metabolic rates, which in turn may reduce fecundity, growth, or immune response. There is compelling evidence in a variety of vertebrate taxa that females appear to prefer skilled problem-solver males, i.e., they prefer those that appear to have better cognitive abilities. Consequently, cognition is also likely to have substantial effects on sexual selection processes. How the choosing sex assesses the cognitive abilities of potential mates has not been explored conclusively yet. Do cognitive skills guide an individual’s mate choice and does learning change an individual’s mate choice decisions? How and to which extent do individuals use their own cognitive skills to assess those of their conspecifics when choosing a mate? How does an individual’s role within a mating system influence the choice of the choosing sex in this context? Drawing on several examples from the vertebrate world, this review aims to elucidate various aspects associated with cognitive sex differences, the different roles of males and females in social and sexual interactions, and the potential influence of cognition on mate choice decisions. Finally, future perspectives aim to identify ways to answer the central question of how the triad of sex, cognition, and mate choice interacts.
Assortative mating is non‐random mating by the mutual choice of phenotypes or behavioral types. In polygynandrous species, competition for mating by social rank can lead to assortative mating. However, although not an individual trait, social bonds also influence mating opportunities resembling assortative mating. Stump‐tailed macaques form long‐term close bonds and organize in a linear dominance–subordination hierarchy. Therefore, we studied whether the strength of the social bond and rank closeness influenced mating decisions and increased mating opportunities, particularly for low‐ and middle‐ranking animals. Firstly, we observed whether females directed proceptive behavior to close‐bonded or adjacent rank males. Secondly, we measured whether successful copulations were related to the strength of social bonds and close ranking. Thirdly, to ensure that copulations owed mainly to the aforementioned factors, we also evaluated whether sexual coercion was unrelated to social bonds and rank similarities. Finally, we assessed whether close bonds mediated agonistic support to females. The study subjects were 12 adult female and 11 male captive stump‐tailed macaques. We monitored daily females' reproductive status by vaginal cytology. Sexual behavior was recorded by all occurrences sampling and scan sampling to collect the agonistic and affiliative instances required to calculate social ranks, social bond strength, and agonistic support. The results indicated that the probability of females displaying proceptivity increased during the follicular phase toward close‐bonded and high‐ranking males. Copulation chances increased with male–female social bonds and rank closeness. Forced copulation decreased in close‐bonded individuals, while agonistic support increased in close‐ranking strong‐bonded animals. In conclusion, close social bonds and similar social rank result in non‐random mating in stump‐tailed macaques. In Macaca arctoides, the strength of social bonds favors the probability of copulation in animals sharing a similar social rank (upper panel). This effect of social bonds is particularly evident in the follicular phase (lower panel).
Full-text available
Behavioural patterns often differ consistently across individuals and are linked to fitness. In species with biparental care, the defence behaviour of both parents can affect reproductive success through offspring survival. In addition to the intensity of defence behaviour by both pair members, the similarity in this behaviour among parents may affect offspring survival. However, few studies have investigated the relative impact of both the intensity and similarity of defence behaviour. Here, we examined nest defence behaviour of males and females during the incubation stage in an Arctic population of barnacle geese Branta leucopsis. We calculated the repeatability of defence behaviour to test whether this behaviour is consistent within individuals and investigated how it is associated with age. In addition, we investigated how daily survival rate (DSR) of the nests until hatching is associated with nest defence behaviour and age of the parents, as well as the effect of parent similarity in nest defence behaviour as an emergent trait of the pair bond. Both male and female defence behaviour were highly repeatable. The ages of both partners within breeding pairs were positively related, but age was only significantly associated with defence behaviour in females. Further, we found high similarity in defence behaviour within breeding pairs, but the similarity and intensity of defence behaviour within breeding pairs did not predict DSR. Finally, male defence behaviour positively predicted DSR, but female defence behaviour and male and female age did not. Our results suggest that nest protection is adaptive in males but behavioural similarity of pair members does not enhance nest survival, indicating behavioural similarity itself is not adaptive but rather a by‐product of different effects.
This chapter reviews the main aspects related to the recent trends in animal personality. Newcomers in animal personality raise some questions about the field and its concepts, and discusses terminological and other issues related to personality. The chapter presents a brief history, with an emphasis on behavioral ecology, and reviews the different sources of consistent individual behavioral variation and describes the main methodological advances in the field over the last few decades. Behavioral ecologists investigated the agents of selection acting on behavior and described how long-term, invariant selection pressures could explain the behavioral strategies currently expressed by animals. Physiological processes can translate the diverse genetic or environmental effects into behavioral differences. Behavioral reaction norms provide an interesting framework to study both personality and plasticity within the same context. Positive assortment based on behavior can generate different social conditions for individuals with different behavioral types.
The notion that men are more variable than women has become embedded into scientific thinking. For mental traits like personality, greater male variability has been partly attributed to biology, underpinned by claims that there is generally greater variation among males than females in non-human animals due to stronger sexual selection on males. However, evidence for greater male variability is limited to morphological traits, and there is little information regarding sex differences in personality-like behaviours for non-human animals. Here, we meta-analysed sex differences in means and variances for over 2100 effects (204 studies) from 220 species (covering five broad taxonomic groups) across five personality traits: boldness, aggression, activity, sociality and exploration. We also tested if sexual size dimorphism, a proxy for sex-specific sexual selection, explains variation in the magnitude of sex differences in personality. We found no significant differences in personality between the sexes. In addition, sexual size dimorphism did not explain variation in the magnitude of the observed sex differences in the mean or variance in personality for any taxonomic group. In sum, we find no evidence for widespread sex differences in variability in non-human animal personality.
Within populations, individuals often show repeatable variation in behaviour, called 'animal personality'. In the last few decades, numerous empirical studies have attempted to elucidate the mechanisms maintaining this variation, such as life-history trade-offs. Theory predicts that among-individual variation in behavioural traits could be maintained if traits that are positively associated with reproduction are simultaneously associated with decreased survival, such that different levels of behavioural expression lead to the same net fitness outcome. However, variation in resource acquisition may also be important in mediating the relationship between individual behaviour and fitness components (survival and reproduction). For example, if certain phenotypes (e.g. dominance or aggressiveness) are associated with higher resource acquisition , those individuals may have both higher reproduction and higher survival, relative to others in the population. When individuals differ in their ability to acquire resources, trade-offs are only expected to be observed at the within-individual level (i.e. for a given amount of resource, if an individual increases its allocation to reproduction, it comes at the cost of allocation to survival, and vice versa), while among individuals traits that are associated with increased survival may also be associated with increased reproduction. We performed a systematic review and meta-analysis, asking: (i) do among-individual differences in behaviour reflect among-individual differences in resource acquisition and/or allocation, and (ii) is the relationship between behaviour and fitness affected by the type of behaviour and the testing environment? Our meta-analysis consisted of 759 estimates from 193 studies. Our meta-analysis revealed a positive correlation between pairs of estimates using both survival and reproduction as fitness proxies. That is, for a given study, behaviours that were associated with increased reproduction were also associated with increased survival, suggesting that variation in behaviour at the among-individual level largely reflects differences among individuals in resource acquisition. Furthermore, we found the same positive correlation between pairs of estimates using both survival and reproduction as fitness proxies at the phenotypic level. This is significant because we also demonstrated that these phenotypic correlations primarily reflect within-individual correlations. Thus, even when accounting for among-individual differences in resource acquisition, we did not find evidence of trade-offs at the within-individual level. Overall, the relationship between behaviour and fitness proxies was not statistically different from zero at the among-individual, phenotypic, and within-individual levels; this relationship was not affected by behavioural category nor by the testing condition. Our meta-analysis highlights that variation in resource acquisition may be more important in driving the relationship between behaviour and fitness than previously thought, including at the within-individual level. We suggest that this may come about via heterogeneity in resource availability or age-related effects, with higher resource availability and/or age leading to state-dependent shifts in behaviour that simultaneously increase both survival and reproduction. We emphasize that future studies examining the mechanisms maintaining behavioural variation in populations should test the link between behavioural expression and resource acquisition-both within and among individuals. Such work will allow the field of animal personality to develop specific predictions regarding the mediating effect of resource acquisition on the fitness consequences of individual behaviour.
Lispe (Diptera: Muscidae) is a cosmopolitan genus of predatory flies that inhabit the muddy and sandy surrounds of water bodies. There are more than 163 described species worldwide, many of which are known to exhibit cursorial courtship displays which involve complex visual and vibratory signals. Despite the widespread distribution of these flies and their remarkable courtship displays, the biology and behaviour of most species are entirely unknown. Here, for the first time, we describe the precopulatory mating behaviours of three widespread and common Australian species: Lispe sydneyensis, Lispe albimaculata and Lispe xenochaeta. We demonstrate that all three species exhibit entirely unique courtship displays, consisting of complex behavioural repertoires. Importantly, we highlight intra‐sexual competition in L. sydneyensis, where males engage in competitive dances and combat. We also report female–male aggression in L. albimaculata and L. xenochaeta where females charge and display towards males. These novel mating systems provide unique opportunities to test ecological and evolutionary hypotheses.
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It is well known that secondary chemicals produced at one trophic level may affect organisms at subsequent levels of the food chain. Effects of nutrient supplements may also propagate through trophic levels, but the mechanisms here are less clear. We tested the hypothesis that predators can be affected by the nutrient composition of the prey's food. Wolf spider (Pardosa amentata) hatchlings were raised ad libitum on fruit flies (Drosophila melanogaster) that were cultured in poor basic medium with additions of different nutrients. These additions strongly affected the performance of the spiders. Growth rates increased when additions consisted of 19 different amino acids or fatty acids+cholesterol or commercial dogfood. Survival increased in spiders reared on fruit flies from cultures containing 19 amino acids or methionine or dogfood. The addition of dogfood increased spider growth and survival more significantly than the addition of any single nutrient group alone. Adult female flies from the dogfood culture were significantly heavier than females from the basic culture. The nutrients added to the fruit fly media were thus able to create biological effects at both the second and the third trophic levels. To test whether nutrients passed to the predators through the gut content of the prey, we included a treatment where the spiders were fed flies that had been starved for 48 h in order to empty their guts. Gut emptying of the flies did not reduce the positive effects of the enriched fruit fly media, i.e. the nutritional benefits were not due to nutrients that passed directly through the guts of the flies. Since the nutrients added to the fruit fly media were separated from the spiders that benefited from them by two trophic transformations, this phenomenon was a true tritrophic interaction.
Spider mating behaviour is varied and often surprising. In the past few decades, there has been a shift from descriptive natural history approaches to a more manipulative, theory-based dissection of the behavioural and evolutionary ecology of mating. This approach has yielded evidence in support of important underlying themes of sexual selection. In this chapter, we summarise patterns of mating behaviour in spiders, and the conditions that underlie variation in this behaviour, with an emphasis on how sexual selection theory relates to observed patterns. We end by suggesting spiders may prove particularly tractable models for testing hypotheses regarding mechanisms of sexual selection, sex-specific mating tactics, and reciprocal links between these, and ecology, demography and life history. Introduction. There are a number of traits common to the true spiders (Order Araneae) that lend unusual dimensions to their mating behaviour (e.g. Uhl and Elias, Chapter 5). Almost all spiders are predacious (for an exception see Meehan et al., 2009), and have sensory systems exquisitely tuned to vibrational and pheromonal signals. Males transfer sperm via specialised intromittent organs (males' palps), not directly connected to the gonads, into females' sperm storage organs of variable number (spermathecae), often via independent insemination tubules (Foelix, 1996). In addition, although the mating season holds risks similar to those for all sexual species (e.g. mate rejection, competitive injury, predation), male spiders (and rarely, females; Aisenberg et al., 2009, Cross et al. 2007b, Jackson and Pollard, 1990, Schutz and Taborsky, 2005) also face the additional risk of mortality through their predacious potential mate.
Preface 1. Temperament and personality: trait structure and persistence 2. Psychobiological methods 3. Extraversion/sociability 4. Neuroticism 5. Psychoticism (psychopathy), impulsivity, sensation and/or novelty seeking, conscientiousness 6. Aggression-hostility/agreeableness 7. Consilience References.
The concept of partner compatibility in monogamous animals implies that individuals may reproduce better when paired to a partner with similar traits than to a higher quality, but dissimilar individual. We investigated whether partner similarities in traits that are linked in a behavioural syndrome influence reproductive performance in a wild population of Steller’s jays. In some years, pairs more similar in explorative tendencies and in willingness to take risks initiated nests earlier and were more likely to fledge offspring than dissimilar pairs. Benefits of behavioural similarity differed among breeding seasons, being most pronounced in a year with late breeding onset after a severe winter. Pairing patterns for behavioural traits also varied among years and traits, and assortative pairing of behaviourally similar partners was not only common overall, but was also correlated across the three explorative and risk‐taking tendencies. Pair members with behavioural similarities may yield more compatible and complementary partnerships. Our results indicate that compatibility across a suite of behavioural traits (i.e. a behavioural syndrome) may be beneficial for assortative pairs and support the hypothesis that the combination of traits in behavioural syndromes in itself might be a target for selection.
There is growing evidence that correlated behavioural traits, or behavioural syndromes, influence behavioural evolution in some taxa. Few studies, however, investigate whether the effects of a syndrome are the same for both sexes. We test whether variation in social tendency, inferred from interindividual distance, is correlated with other aspects of behaviour in male comb-footed spiders, Anelosimus studiosus. We compared these results to those from previous studies on female social tendency to determine (1) whether both sexes share the same behavioural syndrome and (2) whether its effects on mating success are the same for both sexes. Trait types in the syndrome analysis include foraging behaviour, antipredator behaviour, exploratory behaviour and activity level. Our results suggest male A. studiosus, like females, can be categorized into two social classes: an aggregative (social) class and an intolerant (asocial) class. Social males (i.e. those with lower interindividual distance scores) were generally less aggressive towards prey and predators, and were less active. Furthermore, we provide evidence from a parent/offspring breeding study for an additive genetic component to male social tendency (heritability = 0.32). To determine the influence of the male syndrome on mating success, we performed staged male–male contests between social and asocial males for access to females. We found that male social tendency was the single best predictor of success in these trials, with asocial males outperforming social. This finding is opposite to the trend observed in female A. studiosus, where social females experience higher mating success. We propose that the diametrically opposed mating outcomes between the sexes could generate evolutionary conflict.
In the standard job search problem a single decision-maker (say an employer) has to choose from a sequence of candidates of varying fitness. We extend this formulation to allow both employers and candidates to make choices. We consider an infinite population of employers and an infinite population of candidates. Each employer interviews a (possibly infinite) sequence of candidates for a post and has the choice of whether or not to offer a candidate the post. Each candidate is interviewed by a (possibly infinite) sequence of employers and can accept or reject each offer. Each employer seeks to maximise the fitness of the candidate appointed and each candidate seeks to maximise the fitness of their eventual employer. We allow both discounting and/or a cost per interview. We find that there is a unique pair of policies (for employers and candidates respectively) which is in Nash equilibrium. Under these policies each population is partitioned into a finite or countable sequence of subpopulations, such that an employer (candidate) in a given subpopulation ends up matched with the first candidate (employer) encountered from the corresponding subpopulation. In some cases the number of non-empty subpopulations in the two populations will differ and some members of one population will never be matched.