Eunuchs as better fighters?

Abstract

Male-male competition for females can significantly affect a male's reproductive success and hence his fitness. Game theory predicts that an individual should avoid fighting when its future reproductive potential is high, but should fight forcefully when its future reproductive potential is insignificant. When mates are scarce, extreme competition and fatal fighting is expected. We recently showed that Nephilengys malabarensis eunuchs, i.e. sterile spider males that lost their genitals during copulation, become more aggressive during male-male contests. Here, we add crucial comparative data by exploring eunuch fighting behaviour in Nephilengys livida from Madagascar, specifically by testing the 'better fighter hypotheses' in a laboratory setting. Similar to N. malabarensis, N. livida copulations resulted in total male castration with the severed palp plugging the female genitals in 70.83% cases, which mostly (63.63%) prevented subsequent copulations. Unexpectedly, however, N. livida eunuchs exhibited lower aggressiveness than virgin males. We interpret these results in the light of different mating biology between the so far studied species known for the eunuch phenomenon, which might reflect differing plug effectiveness due to variation in genital anatomy in N. livida, N. malabarensis and Herennia multipuncta. However, detected differences in aggressive behaviour of N. livida versus N. malabarensis eunuchs might also be explained by the species' ecology, with lower population densities resulting in a relaxed male-male competition making excessive aggression and mate guarding redundant. This study thus questions the generality of overt aggressiveness in mated males with no reproductive value, and highlights the importance of understanding the natural history of species in the question.

Full text

ORIGINAL PAPER
Eunuchs as better fighters?
Simona Kralj-Fišer &MatjažKuntner
Received: 26 October 2011 /Revised: 27 November 2011 /Accepted: 30 November 2011 / Published online: 14 December 2011
#Springer-Verlag 2011
Abstract Male–male competition for females can signifi-
cantly affect a male’s reproductive success and hence his
fitness. Game theory predicts that an individual should
avoid fighting when its future reproductive potential is high,
but should fight forcefully when its future reproductive
potential is insignificant. When mates are scarce, extreme
competition and fatal fighting is expected. We recently
showed that Nephilengys malabarensis eunuchs, i.e. sterile
spider males that lost their genitals during copulation, be-
come more aggressive during male–male contests. Here, we
add crucial comparative data by exploring eunuch fighting
behaviour in Nephilengys livida from Madagascar, specifi-
cally by testing the ‘better fighter hypotheses’in a labora-
tory setting. Similar to N. malabarensis,N. livida
copulations resulted in total male castration with the severed
palp plugging the female genitals in 70.83% cases, which
mostly (63.63%) prevented subsequent copulations. Unex-
pectedly, however, N. livida eunuchs exhibited lower ag-
gressiveness than virgin males. We interpret these results in
the light of different mating biology between the so far
studied species known for the eunuch phenomenon, which
might reflect differing plug effectiveness due to variation in
genital anatomy in N. livida,N. malabarensis and Herennia
multipuncta. However, detected differences in aggressive
behaviour of N. livida versus N. malabarensis eunuchs
might also be explained by the species’ecology, with lower
population densities resulting in a relaxed male–male com-
petition making excessive aggression and mate guarding
redundant. This study thus questions the generality of overt
aggressiveness in mated males with no reproductive value,
and highlights the importance of understanding the natural
history of species in the question.
Keywords Aggression .Cannibalism .Emasculation .
Guarding .Plugging .Nephilidae .Nephilengys livida
Introduction
Male–male competition over females and related resources
can significantly affect a male’s reproductive success and
hence his fitness (Huntingford and Turner 1987). Yet, costs
associated with competition favour the assessment of a
resource value relative to the contestant’s fighting ability
(also termed resource-holding value, e.g. weapons, size)
over confronting all rivals indiscriminately (Maynard Smith
and Parker 1976; Parker 1970; but see Moore et al. 2008).
An important factor is an individual’s expected future re-
productive value (Enquist and Leimar 1990). When its
future reproductive potential is high, an animal should avoid
Communicated by: Sven Thatje
Electronic supplementary material The online version of this article
(doi:10.1007/s00114-011-0873-1) contains supplementary material,
which is available to authorized users.
S. Kralj-Fišer :M. Kuntner
Institute of Biology, Scientific Research Centre,
Slovenian Academy of Sciences and Arts,
Ljubljana, Slovenia
S. Kralj-Fišer
Biozentrum Grindel, University of Hamburg,
Martin-Luther-King Platz 3,
20146, Hamburg, Germany
M. Kuntner
National Museum of Natural History, Smithsonian Institution,
Washington, DC, USA
M. Kuntner (*)
College of Life Sciences, Hubei University,
Wuhan 430062 Hubei, China
e-mail: kuntner@gmail.com
Naturwissenschaften (2012) 99:95–101
DOI 10.1007/s00114-011-0873-1

fighting and hence injury, but should fight forcefully when its
future reproductive potential is insignificant (e.g. Fromhage
and Schneider 2005a; Innocent et al. 2007). When females are
limited, extreme competition and fatal fighting is expected
(Bean and Cook 2001; Enquist and Leimar 1990; Maynard
Smith and Price 1973).
Male competitive behaviour for females often varies,
being dependent on the density of rivals, the availability
and value of the contestant resource, as well as the related-
ness between competitors (Elias et al. 2010; Enquist and
Leimar 1987; Innocent et al. 2011; Kasumovic et al. 2008;
Kokko and Rankin 2006; Moore et al. 2008; Murray and
Gerrard 1985; Reinhold 2003; West et al. 2001,2002). Male
encounter rates with females depend on female density
within a patch, on patch size and on the distance between
patches (Schneider and Lubin 1998). When males’chances
to encounter more females are high, they are expected to
avoid the risk of intensive fighting. On the other hand, if
males’chances of encountering more than one female are
slim, they are predicted to maximize their reproductive
success by investing all of their resources in the first female
they encounter (Buskirk et al. 1984). However, the situation
also depends on the density of the males, and in the cases
where the density of males is as low as that of rare females,
there might not be a need for the male to defend his paternity
(Fromhage et al. 2005,2008).
In many sexually dimorphic spider species, males break
their copulatory organs, the palps, during copulation to plug
up a female (Fromhage and Schneider 2006; Kuntner 2005,
2007; Kuntner et al. 2008; Kuntner et al. 2009a; Uhl et al.
2010). A ‘mating plug’hypothesis postulates that these
palpal leftovers function as physical barriers to future cop-
ulations (Kralj-Fišer et al. 2011a). In most species, the males
with damaged or missing palps are functionally sterile and
have no further reproductive value. Despite high costs re-
lated to mating plugs and sterility, these do not always
effectively prevent female remating, and this may depend
on both male and/or female genital morphology (Kuntner et
al. 2009b; Uhl et al. 2010). Due to incomplete plug effec-
tiveness, those emasculated males that survive copulation
commonly practice post-copulatory mate guarding (Fromhage
and Schneider 2005b). In accordance with game theory
(Enquist and Leimar 1990), Nephilengys malabarensis males,
which entirely emasculate their palps during copulation (such
sterile males are termed ‘eunuchs’), commonly escalate fierce
fighting and win in contests against rivals—a‘better fighter’
phenomenon (Kralj-Fišer et al. 2011a). To elucidate the gen-
erality of detected patterns in eunuch behaviour, we here
expand our research to include the second Nephilengys
species.
Nephilengys livida is a highly sexually size dimorphic
and synanthropic species (Kuntner 2007;Kuntnerand
Agnarsson 2011), which exhibits a combination of inviting
sexual traits such as complex genitalia, male genital damage
resulting in plugging and eunuchs, as well as sexual canni-
balism and post-copulatory mate guarding (Kralj-Fišer et al.
2011b; Kuntner 2007; Kuntner et al. 2009c). The interaction
between current resource value, future resource value, in-
vestment in sperm plugs and male–male competition for a
female, make this species an interesting system for testing
the predictions of game theory models. We explored male
fighting behaviour in N. livida, by conducting a series of
laboratory male–male contests on female webs. We com-
pared contests between two virgin rivals with those between
a virgin male and a eunuch. Since plug efficiency likely
affects male fighting behaviour, e.g. effective plugs reduce
sperm competition, we staged laboratory mating trials to test
if plugged female genitalia could be reused by another male.
In accordance with the better fighter hypothesis (Kralj-Fišer
et al. 2011a), we predicted that eunuchs would escalate
fighting intensity to outcompete virgin rivals and that
eunuchs would be more aggressive in a species with lower
plug efficiency.
Materials and methods
Study animals
We collected N. livida (formerly known as Nephilengys bor-
bonica, Kuntner 2007) spiders in Andasibe-Mantadia (Toa-
masina Province) and Ranomafana (Fianarantsoa Province) in
Madagascar, between 24 February 2010 and 4 April 2010. To
examine remating, we collected subadults and reared them to
adulthood in the laboratory (females019, males033). We
housed females in glass frames (50×50×10 cm), and males
in smaller plastic cups (250 ml). We provided water to spiders
daily and fed them fruit flies, crickets and mealworms twice a
week. As previously, we define eunuchs as those adult males
lacking bothpalps, and half-eunuchs as those lacking one palp
(Kralj-Fišer et al. 2011a). We compared N. livida with N.
malabarensis from Southeast Asia (for details see Kralj-
Fišer et al. 2011a).
Experimental protocol
We tested the ‘better fighter’and the ‘mating plug’
hypotheses in series of laboratory tests. The spiders
reached maturation at various times, thus we had to
continuously adapt our ongoing experiments to the
available spiders (see work flow table in ESM). Due
to the small number of wild-caught spiders, several
spiders were reused (Nreused males024; Nreused
females019; see also ESM), however, we considered
this in our analyses.
96 Naturwissenschaften (2012) 99:95–101

Better fighter hypothesis
To establish if palpal severance during copulation triggers
mate guarding, particularly fighting as known in N. mala-
barensis (Kralj-Fišer et al. 2011a), eunuchs were placed on a
random female web, immediately followed by the introduc-
tion of the second, virgin male (N012 trials). The control
group staged contests between two virgin males (N012
trials). During male–male contests, we noted the frequencies
of guarding behaviours, i.e. walking towards female, touch-
ing female and exploring (walking back and forth), and
estimated the distance between males and a female every
five minutes. Male–male fighting behaviour was scored as
frequencies of walking towards another male (score 01),
web shaking (score 02), chasing (score03) and attacking
(score04). Aggressiveness intensity levels were estimated
as sums of scores (Kralj-Fišer et al. 2011a,b). We also noted
frequencies of escaping and web plucking (defined as male
tapping on web threads). We noted whether copulations
took place. Trials lasted 60 min. After a trial the spiders
were given at least a 24-h rest.
Mating plug hypothesis
To test for genital reuse, our aim was to document at least
two successful insertions in the same copulatory opening
(CO). We took 11 females that received plugs and males that
could only insert their palp(s) in the plugged CO (insertions
were always ipsilateral; N013, see ESM). If mating into the
used CO was attempted but was unsuccessful in two subse-
quent mating trials, we assumed that plugging prevented
successful access to the previously used CO (Kralj-Fišer et
al. 2011a). Observations lasted for 60 min. After a trial the
spiders were given at least a 24-h rest. In the analysis, we
considered also rematings into the used CO that occurred
during male–male contests.
Morphological examination
At the end of all trials, the females were euthanized and
preserved in 70% ethanol. The epigyna of 17 females (two
females escaped) were excised and macerated in concentrated
KOH for 24 h, cleaned in distilled water, then further treated
with methyl salicilate (Kuntner et al. 2009b). Examinations of
epigynal microscopic preparations were done under a Leica
MZ16 stereomicroscope.
Statistical analyses
We compared plugging efficiency between N. livida and
N. malabarensis using a chi square test. To detect the
effect of emasculation on fighting and guarding behav-
iours of N. livida eunuchs, we employed generalized
linear mixed models (GLMMs). The fixed factors were
male’sandrival’s reproductive status (10eunuch, 20
virgin). We introduced male identities as random fac-
tors, because some males were reused. Response varia-
bles were behavioural scores. We sequentially deleted
fixed terms in order of decreasing significance; only
terms with P≤0.1 remained in the final model. Excluded
terms were re-entered one by one into the final model to
confirm that they did not explain a significant part of the
variation (Poesel et al. 2006). We present Wald statistics for
final models including fixed terms with P≤0.1 only. The
differences of behavioural measures—where P≤0.1—were
interpreted using the graphs. The analyses were done in
PASW version 18.
Results
Better fighter hypothesis
Virgin males were generally more aggressive than
eunuchs in male–male contests: virgin males shook the
web more often, walked more frequently towards a rival
and had higher general aggressiveness intensity than eu-
nuch males (for probabilities see Table 1; Fig. 1a). Virgin
males were more aggressive when confronted with an-
other virgin male than when opposed with a eunuch
(frequency of walking towards a rival, aggressiveness
intensity; Table 1). Interestingly, virgin males exhibited
escape behaviour more frequently than eunuchs (Table 1).
Virgin males signalled (web plucking) more frequently
than eunuchs, again more often so when confronted with
Table 1 Statistical results of the final GLMM model
State of ♂1 State of ♂2
Signal (pluck web) 0.037 0.006
Touch female >0.1 >0.1
Explore web >0.1 >0.1
Walk towards female 0.064 >0.1
Distance to female 0.003 0.004
Shake 0.034 >0.1
Walk towards rival 0.002 0.01
Attack >0.1 >0.1
Chase >0.1 >0.1
Intensity of aggressiveness 0.009 0.015
Escape 0.1 >0.1
Probabilities that fixed factors, e.g. number of palps in male 1, number of
palps in the rival male (male 2) affect the occurrences of behaviours in
male 1. Statistical probabilities P≤0.05 bolded, 0.05<P<0.1 italicized.
N024 contests, Nof trials virgin vs virgin012; Nof trials virgin vs
eunuch012
Naturwissenschaften (2012) 99:95–101 97

another virgin rival (Table 1). Eunuchs did not employ
guarding behaviour: eunuchs and virgin males did not
differ in frequencies of touching a female and exploring
a web; however, virgin males more often walked towards
a female and stayed closer to her than eunuchs (Table 1;
Fig. 1b). Again, virgin males stayed closer to a female
when opposed with a virgin rival than when opposed
with a eunuch male (Table 1, Fig. 1b).
Mating plug hypothesis
In total we observed 31 copulations (Nfemales019; N
males021). A palp insertion always ended with total emas-
culation, resulting in whole-palpal mating plug in the used
CO. Mating plugs remained in the used female CO after
copulation termination, but were usually absent (externally)
several hours thereafter. In two cases, a plug was observed
Fig. 1 Male behaviours
during male–male contests.
aaggressiveness intensity;
baverage distance to the
female (cm) of the male 1
in the contest with male 2
98 Naturwissenschaften (2012) 99:95–101

externally even a day after copulation. We observed four
females removing their plugs using the third pair of legs.
Plugging mostly prevented subsequent copulation into the
used female CO: in seven out of 11 females with a plugged
CO (63.63%) no further copulations occurred, whereas in
36.36% (four out of eleven) of cases a subsequent male used
the already plugged CO and produced the second plug seen
externally. Plugging efficiency by means of unsuccessful in-
sertion attempts in N. livida did not significantly differ from
plugging efficiency in N. malabarensis (χ
2
00.21, df01, p0
0.647, N022).
Our morphological examination revealed broken parts of
embolic conductor (EC) within the internal female genital
tract (within copulatory duct and/or spermathecae) in 17 out
of 24 cases (70.83%, Nfemales017). Eleven females were
subjected to males that could only copulate in the used CO.
We found a double plug in one such female, i.e. two parts of
EC in the remated organ. The other remated female had no
internal plug despite the previous occurrence of two external
palpal plugs. In seven females that did not remate, we found
parts of EC in the used genitals.
Discussion
Our study found significantly different results to those pre-
dicted by theory, where mated (and functionally sterile)
males with no residual reproductive value (eunuchs) are
predicted to fight with maximal force when facing an
intruder (e.g. Fromhage and Schneider 2005a; Kralj-Fišer
et al. 2011a). Unexpectedly, eunuch behaviour in N. livida
differed strikingly from that in N. malabarensis, despite
these species being congeneric and exhibiting a similar
sexual biology (Kralj-Fišer et al. 2011a). In both species,
copulation leads to total emasculation of the used palp and
plugging of the female copulatory opening; about 70% of
males fall victim to sexual cannibalism, and about 30% of
females can remate using the same CO (this study; Kralj-
Fišer et al. 2011a). Contrary to the theory predicting that
surviving males with no reproductive value will escalate
fatal fighting (Enquist and Leimar 1990; Maynard Smith
and Price 1973), this was never observed in N. livida and
only rarely in N. malabarensis (this study; Kralj-Fišer et al.
2011a). Nevertheless, N. malabarensis eunuchs exhibited
higher aggressiveness levels than their virgin rivals in the
male–male contests on the female webs as predicted by
game theory (Kralj-Fišer et al. 2011a), while N. livida
eunuchs behaved rather meekly in comparison with virgin
males. A study on parasitoid wasps Mellittobia similarly found
mated and virgin males exhibiting similar levels of aggression
(Innocent et al. 2011). The contrasting results between the
studies suggest that a male fighting behaviour depends on
additional factors, not only future reproductive value. Potential
benefits of fighting may vary with plug efficiency, sperm
precedence, ability to assess resource value, and/or resource
density (e.g. Innocent et al. 2011; Reinhold 2003).
Perhaps unexpectedly, this study invalidates the general-
ity of eunuch-enhanced fighting abilities in spiders, but it
may provide some new insights into the N. livida mating
system in particular and into the eunuch biology in general.
In both studied Nephilengys species and in Herennia, eu-
nuch contests were observed at least 1 day after copulation:
after the male had lost a palp(s), it was separated from the
female for a day, then placed on a random female web for a
male–male contest (Kralj-Fišer et al. 2011a; Kuntner et al.
2009b). Despite a time lag and an unfamiliar female, N.
malabarensis or Herennia eunuchs aggressively fought a
virgin rival off, while the N. livida eunuchs in this study
did not. A possible explanation is that in nature, N. livida
eunuchs guard their mates only shortly after copulation, but
leave the female web, or even offer themselves as prey to the
female thereafter. Such strategy is known in other orbweav-
ing spiders, e.g. Argiope keyserlingi (Herberstein et al.
2005), and hints at a pronounced first sperm priority, where
mate guarding is only adaptive until the female fertilizes the
eggs. That N. malabarensis and Herennia eunuchs are more
persistent in mate-guarding might suggest a fiercer sperm
competition and perhaps longer sperm storage.
A possible reason for discrepancies between both Nephi-
lengys studies may also be a difference in efficiency of sperm
plugs to prevent paternity of subsequent suitors. This is be-
cause a mere reuse of a plugged CO does not necessarily
Fig. 2 Morphological comparison in palpal structure between N. livida
(short, wide and broad tip), N. malabarensis (long and thin tip) and H.
multipuncta (long and thin tip). Redrawn from Kuntner (2005,2007).
Scale lines00.1 mm. CB cymbium, EC embolic conductor, ST subtegu-
lum, Ttegulum
Naturwissenschaften (2012) 99:95–101 99

imply paternity. Although the ultimate test of this would be
paternity analysis, which was beyond our scope here, genital
morphology suggests plug efficiency by means of protecting
paternity (Kuntner et al. 2009b). N. malabarensis and Here-
nnia multipuncta males have longer, thinner and more hooked
embolic conductors compared with short, wide and broad-
tipped ones in N. livida (Kuntner 2005;2007; Fig. 2). We
speculate that broader embolic plugs better shield the female
CO and thus prevent subsequent sperm transfer into sperma-
thecae than thinner plugs. Although the shift to more complex
and wider palps (N. livida) may have coevolved with
corresponding counter adaptations in females (Kuntner et al.
2009b), we find it nevertheless plausible that plug efficiency is
higher in N. livida with more complex palps than in N.
malabarensis and H. multipuncta with relatively simpler ones.
If true, more aggressive behaviour of N. malabarensis and H.
multipuncta compared to N. livida eunuchs might be means of
compensating for plug inefficiency.
Our results might also imply that in contrast to N. mala-
barensis and Herennia, N. livida males might be able to assess
whether a given female has in fact been their mate. Further-
more, N. livida males might also be able to assess their rivals’
status, i.e. virgin versus sterile male, in the female web. Virgin
males signalled, tried to approach the female and engaged in
agonistic interactions more often when confronted with an-
other virgin male than when opposed by a eunuch. Such
behaviour of virgin males might be enhanced by females,
which significantly more often cannibalized non-aggressive
than aggressive males (Kralj-Fišer et al. 2011b;e.g.
Stoltz et al. 2008). Such female choice strategy might
explain the differences in aggressiveness in N. livida and N.
malabarensis virgin males. Interestingly, despite their higher
aggressiveness, virgin males nevertheless retreated more often
than eunuchs. Such higher risk aversion in virgin males com-
pared with mated ones is in accordance with game theory
(Fromhage and Schneider 2005a; Kralj-Fišer et al. 2011a).
Finally, competitive and mating behavioursmay depend on
ecological factors. Males may adapt their mating strategies
according to female density, levels of male competition, patch
size, etc. (Innocent et al. 2011; Kasumovic et al. 2008; Kokko
and Rankin 2006; Reece et al. 2007). Population density, in
particular, is important for mate guarding behaviour; if popu-
lation densities intensify male–male competition through
male-biased operational sex ratio, the intensity of mate guard-
ing is expected to increase (Davis and Brown 1999; Hardling
2004; Jormalainen 1998). According to our observations in
nature, N. livida occur at lower local abundances compared to
N. malabarensis. In populations with lower densities, male
survival during mate searching might be lower than in pop-
ulations with high densities, resulting in further reduced
numbers of rival males. If so, sperm competition in N. livida
is reduced, which makes paternity protection additional to
sperm plugs unnecessary (Fromhage et al. 2005,2008).
In conclusion, our study found no support for the better
fighter hypothesis in N. livida eunuchs. Differing mating
biology between the so-far-studied species known for the
eunuch phenomenon may account for this result. However,
non-aggressive behaviour of N. livida eunuchs might also be
explained by population ecology, with lower population
densities perhaps resulting in relaxed male–male competi-
tion, which makes excessive aggression and mate guarding
redundant.
Acknowledgements We thank three anonymous reviewers and Jutta
Schneider for comments on the manuscript, MatjažGregoričfor the
spider collection and laboratory help; Živa Justinek for help with
experiments and lab support; and Ingi Agnarsson, Sahondra Lalao
Rahanitriniaina and Honore Rabarison for their help in field. This work
was funded by the Slovenian Research Agency (grant J1-2063 to MK)
and the National Geographic Society (grant 8655-09 to I. Agnarsson,
M. Kuntner and T. Blackledge). SKF was supported by Humboldt
fellowship for postdoctoral researchers and Humboldt return
fellowship.
References
Bean D, Cook JM (2001) Male mating tactics and lethal combat in the
nonpollinating fig wasp Sycoscapter australis.AnimBehav
62:535–542
Buskirk RE, Frohlich C, Ross KE (1984) The natural selection of
sexual cannibalism. Am Nat 123:612–625
Elias DO, Botero CA, Andrade MCB, Mason AC, Kasumovic MM
(2010) High resource valuation fuels “desperado”fighting tactics
in female jumping spiders. Behav Ecol 21:868–875
Davis JA, Brown CR (1999) Costs of coloniality and the effect of
colony size on reproductive success in purple martins. Condor
101:737–745
Enquist M, Leimar O (1987) Evolution of fighting behavior—the
effect of variation in resource value. J Theor Biol 127:187–205
Enquist M, Leimar O (1990) The evolution of fatal fighting. Anim
Behav 39:1–9
Fromhage L, Schneider JM (2005a) Virgin doves and mated hawks:
contest behaviour in a spider. Anim Behav 70:1099–1104
Fromhage L, Schneider JM (2005b) Safer sex with feeding females:
sexual conflict in a cannibalistic spider. Behav Ecol 16:377–382
Fromhage L, Schneider JM (2006) Emasculation to plug up females:
the significance of pedipalp damage in Nephila fenestrata. Behav
Ecol 17:353–357
Fromhage L, Elgar MA, Schneider JM (2005) Faithful without care:
the evolution of monogyny. Evolution 59:1400–1405
Fromhage L, Houston AI, McNamara JM (2008) A model for the
evolutionary maintenance of monogyny in spiders. J Theor Biol
250:524–531
Hardling KA (2004) Male brood care without paternity increases
mating success. Behav Ecol 15:715–772
Herberstein ME, Barry KL, Turoczy MA, Wills E, Youssef C, Elgar
MA (2005) Post-copulation mate guarding in the sexually canni-
balistic St Andrew's Cross spider (Araneae Araneidae). Ethol Ecol
Evol 17:17–26
Huntingford FA, Turner AK (1987) Animal Conflict. Chapman and
Hall, London
Innocent TM, Savage J, West SA, Reece SE (2007) Lethal combat and
sex ratio evolution in a parasitoid wasp. Behav Ecol 8:709–715
100 Naturwissenschaften (2012) 99:95–101

Innocent TM, West SA, Sanderson JL, Hyrkkanen N, Reece SE (2011)
Lethal combat over limited resources: testing the importance of
competitors and kin. Behav Ecol 22:923–931
Jormalainen V (1998) Precopulatory mate guarding in crustaceans:
male competitive strategy and intersexual conflict. Q Rev Biol
73:275–304
Kasumovic MM, Bruce MB, Andrade MCB, Herberstein ME (2008)
Spatial and temporal demographic variation drives within-season
fluctuations in sexual selection. Evolution 62:2316–2325
Kokko H, Rankin DJ (2006) Lonely hearts or sex in the city? Density-
dependent effects in mating systems. Phil Trans R Soc B
1466:319–334
Kralj-Fišer S, GregoričM, Zhang SC, Li D, Kuntner M (2011a)
Eunuchs are better fighters. Anim Behav 81:933–939
Kralj-Fišer S, Schneider JM, Justinek Ž, Kalin S, GregoričM, Kuntner
M (2011b) Mate quality, not aggressive spillover, explains sexual
cannibalism in a size dimorphic spider. Behav Ecol Sociobiol
doi:10.1007/s00265-011-1262-7
Kuntner M (2005) A revision of Herennia (Araneae: Nephilidae:
Nephilinae), the Australasian 'coin spiders'. Invertebr Syst
19:391–436
Kuntner M (2007) A monograph of Nephilengys, the pantropical
'hermit spiders' (Araneae, Nephilidae, Nephilinae). Syst Entomol
32:95–135
Kuntner M, Coddington JA, Hormiga G (2008) Phylogeny of extant
nephilid orb-weaving spiders (Araneae, Nephilidae): testing mor-
phological and ethological homologies. Cladistics 24:147–217
Kuntner M, Kralj-Fišer S, Schneider JM, Li D (2009a) Mate plugging
via genital mutilation in nephilid spiders: an evolutionary hypoth-
esis. J Zool 277:257–266
Kuntner M, Coddington JA, Schneider JM (2009b) Intersexual arms
race? Genital coevolution in nephilid spiders (Araneae, Nephilidae).
Evolution 63:1451–1463
Kuntner M, Agnarsson I, GregoričM (2009c) Nephilid spider eunuch
phenomenon induced by female or rival male aggressiveness. J
Arachnol 37:266–271
Kuntner M, Agnarsson I (2011) Biogeography and diversification of
hermit spiders on Indian Ocean islands (Nephilidae: Nephi-
lengys). Mol Phylogenet Evol 59:477–488
Maynard Smith J, Price GR (1973) Logic of animal conflict. Nature
246:15–18
Maynard Smith J, Parker GA (1976) The logic of asymmetric contests.
Anim Behav 24:159–175
Moore JC, Obbard DJ, Reuter C, West SA, Cook JM (2008) Fighting
strategies in two species of fig wasp. Anim Behav 76:315–322
Murray MG, Gerrard R (1985) Putting the challenge into resource
exploitation—a model of contest competition. J Theor Biol
115:367–389
Parker GA (1970) Sperm competition and its evolutionary consequen-
ces in insects. Biol Rev Camb Philos Soc 45:525
Poesel A, Kunc HP, Foerster K, Johnsen A, Kempenaers B (2006)
Early birds are sexy: male age, dawn song and extrapair paternity
in blue tits, Cyanistes (formerly Parus)caeruleus. Anim Behav
72:531–538
Reece SE, Innocent TM, West SA (2007) Lethal male–male combat in
the parasitoid Melittobia acasta: are size and competitive envi-
ronment important? Anim Behav 74:1163–1169
Reinhold K (2003) Influence of male relatedness on lethal combat in
fig wasps: a theoretical analysis. Proc R Soc Lond Ser B Biol Sci
270:1171–1175
Schneider JM, Lubin Y (1998) Intersexual conflict in spider. Oikos
83:496–506
Stoltz JA, Elias DO, Andrade MCB (2008) Females reward courtship by
competing males in a cannibalistic spider. Behav Ecol 62:689–697
Uhl G, Nessler S, Schneider J (2010) Securing paternity in spiders? A
review on occurrence and effects of mating plugs and male genital
mutilation. Genetica 138:75–104
West SA, Murray MG, Machado CA, Griffin AS, Herre EA (2001)
Testing Hamilton’s rule with competition between relatives. Nature
409:510–513
West SA, Pen I, Griffin AS (2002) Conflict and cooperation—cooper-
ation and competition between relatives. Science 296:72–75
Naturwissenschaften (2012) 99:95–101 101