ANIMAL BEHAVIOUR, 2006, 71, 481–490
doi:10.1016/j.anbehav.2005.05.011
REVIEWS
Sexual cannibalism and mate choice
JOHN PRENTER, CALUM MacNEIL & ROBERT W. ELWOOD
School of Biology & Biochemistry, Queen’s University Belfast
(Received 17 September 2004; initial acceptance 29 October 2004;
final acceptance 7 May 2005; published online 13 February 2006; MS. number: RV-53R)
Sexual cannibalism, where a female kills and consumes a courting male, represents an extreme form of sexual conflict and has been proposed as a mechanism of mate choice. We evaluate the evidence for mate
choice through premating sexual cannibalism via mate rejection, other indirect mechanisms of mate
‘choice’ and choice in postmating sexual cannibalism. We highlight a paucity of investigations, particularly
of field studies, and note gaps in our knowledge. There is empirical support for the size-dependent sexual
cannibalism predicted by mate choice through premating sexual cannibalism. This may represent mate
choice operating on absolute male size but it could be a by-product of female foraging behaviour and greater
vulnerability of relatively smaller males. Thus, indirect mate choice is as plausible an explanation of sizedependent sexual cannibalism as is direct mate choice based on discrimination of male traits. Direct female
choice, mediated through premating sexual cannibalism, has yet to be demonstrated. We suggest a framework for distinguishing direct and indirect choice and note an absence of information on which to test it.
There is evidence for sequential mate choice in postmating sexual cannibalism, but the nature or basis of
the female’s discriminatory behaviour remains unclear. Costs and long-term fitness benefits of the putative
mate choice have been largely ignored. Reversed sexual cannibalism, in which the male eats the female,
presumably occurs when the gain from food is high and potential gain from mating low and probably
has little to do with mate choice.
Ó 2005 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.
Mr. Blackwall has sometimes seen two or more males on the same web with a single female, but their courtship is too tedious and prolonged an affair to be easily observed. The male is extremely cautious in making
his advances, as the female carries her coyness to a dangerous pitch. De Geer saw a male that ‘‘in the midst
of his preparatory caresses was seized by her in a web and then devoured’’, a sight as he adds filled him
with horror and ‘‘indignation’’.
C. Darwin (1871).
Sexual cannibalism, where females kill and consume
potential or actual mates before, during or after mating,
is common in spiders and occurs in some other invertebrates (Elgar 1992; Johns & Maxwell 1997; Schneider &
Elgar 2002; Elgar & Schneider 2005; Huber 2005). It may
represent the pinnacle of sexual conflict (reviewed in
Chapman & Partridge 1996; Brown et al. 1997; Schneider
& Lubin 1998; Elgar & Schneider 2005; Schneider et al.
2006; but see Andrade 1998). Males use various strategies
to avoid being cannibalized (reviewed in Robinson 1982;
Elgar 1992; Moya-Laraño et al. 2004), including waiting
Correspondence: J. Prenter, School of Biology & Biochemistry, Queen’s
University Belfast, Medical Biology Centre, 97 Lisburn Road, Belfast
BT9 7BL, U.K. (email: j.prenter@qub.ac.uk). C. MacNeil is now at
the Government Laboratory, Ballakermeen Road, Douglas, Isle of
Man IM1 4BR, U.K.
0003–3472/05/$30.00/0
for the female to catch a fly before attempting to mate
(e.g. Austin & Anderson 1978; Prenter et al. 1994a, b;
Schneider et al. 2001; Fromhage & Schneider 2005) and
mating with vulnerable, newly moulted females (e.g. Elgar
1992). The caution with which males court females and
the tenaciousness of their postmating escape attempts
suggest that sexual cannibalism is a strong selection force,
even if the incidence is low (Morse 2004) and, from the
male’s point of view, maladaptive (Foellmer & Fairbairn
2004); however, it has long been viewed as a potential
mechanism for female mate choice (Darwin 1871; Elgar
& Nash 1988; Eberhard 1996). We review the evidence
for sexual cannibalism as a mate choice mechanism in
premating sexual cannibalism (PRE-SC) and postcopulatory or postmating sexual cannibalism (POST-SC). We also
briefly examine reversed sexual cannibalism, where courting males kill females (e.g. Dick 1995; Tsai & Dai 2003;
Schütz & Taborsky 2005), and the possibility of male
481
Ó 2005 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.
482
ANIMAL BEHAVIOUR, 71, 3
mate choice in sexual cannibalism. While there is a paucity
of studies that address the central issue of mate choice
through sexual cannibalism, there is a need to examine
support for the various hypotheses critically and also
highlight gaps in knowledge that may serve as a strategic
focus for further study. Most of our discussion will relate
to spiders because of a bias in the literature.
DIRECT AND INDIRECT MATE CHOICE
Direct mate choice by sexual cannibalism is an outcome of
a specific discrimination by females, often by size, since size
is strongly correlated with many fitness characteristics (e.g.
Blanckenhorn 2000), and may be inferred from decreased
aggression towards certain males and increased aggression
directed at others. Hebets (2003) showed that adult female
wolf spiders, Schizocosa uetzi, preferred to mate with males
of a familiar phenotype (subadult exposure). Unfamiliar
males were more likely to be cannibalized by females. However, we have excluded this study from our survey of PRE-SC
(see below, Table 1) because of the unnatural nature of the
manipulation of male phenotypes undertaken.
Whenever variation in a given male trait, for example
size, is associated with variation in female behaviour, for
example sexual cannibalism, and in male reproductive
success, selection on the particular trait is expected.
Differential sexual cannibalism, however, may not always
involve direct female choice. We are thus faced with the
dichotomy of ideas that represent direct and indirect mate
choice (e.g. Darwin 1871; Kirkpatrick 1987; Andersson
1994; Wiley & Poston 1996). Indirect mate choice embraces all other behaviour or attributes that restrict the mates
of females but without discrimination or active choice on
the part of the female.
MATE REJECTION
Established criteria guide researchers investigating direct
mate choice (e.g. Halliday 1983; Searcy & Andersson 1986;
Andersson 1994; Eberhard 2000). Eberhard (1996) suggested that (1) females should respond differently to
some conspecifics (i.e. discriminate between them), (2)
the discrimination should occur under natural conditions,
(3) discrimination by the female between different males
Table 1. Summary of evidence for size-related precopulatory sexual cannibalism (PRE-SC) and mate choice
Criteria for
mate choice
Species
Differential sexual
cannibalism and nature
Other female
discrimination
1
2
3
4
5
Source
Y
N
Y(?)
Y
?
Elgar & Nash 1988
Y
N
Y(?)
Y
?
Arnqvist 1992; Arnqvist
& Henriksson 1997
N
N
N
N
?
Johnson 2001, 2005b
Y
Y
Y(?)
Y
?
Rubenstein 1987
Y
N
?
Y(?)
?
Persons & Uetz 2005
N
N
N
N
Y
N
?
?
?
?
?
N
?
?
?
Uhl & Vollrath 1998
Uhl & Vollrath 1998
Elgar & Fahey 1996
?
Y
?
?
?
Araneus
diadematus
Smaller males
cannibalized
Dolomedes
fimbriatus
Larger males avoided
PRE-SC
Dolomedes triton
ND
Metellina
segmentata
Schizocosa
ocreata
Smaller males
cannibalized*
Smaller males more
likely to be attacked
and cannibalized
Nephila edulis
Nephila clavipes
Nephila plumipes
ND
ND
ND
Micrathena
gracilis
Gasteracantha
minax
Gasteracantha
cancriformis
Misumenia vatia
N/A
Males with higher
pseudocopulation rates
less likely to suffer PRE-SC
ND; larger males better at
gaining copulations
(once mounted)
ND; larger males escape
attack more often but not
translated into differential
reproductive success
Only larger males mate
multiply
Attack smaller males
(with smaller leg tufts),
probability of PRE-SC varied
with relative size of pair;
males with larger leg tufts
more likely to mate
N/A
N/A
Females responded to larger
males and ignored smaller
males but no data on attack
behaviour
N/A
ND
N/A
N
N
?
N
?
Bukowski &
Christenson 2000y
Elgar & Bathgate 1996
N/A
N/A
?
Y
?
?
?
Bukowski et al. 2001y
ND
ND
N
?
?
?
?
Morse 2004z
See text for explanation of criteria for mate choice (based on Eberhard 1996). Y ¼ yes, criteria met; N ¼ no, criteria not met; ? ¼ uncertain/no
available data; Y(?) ¼ probable; ND ¼ none detected; N/A ¼ not applicable/data not available.
*Cannibalism by females of adult males during male–male fights.
yStudy focused on male behaviour and sperm competition.
zDespite a lack of empirical evidence, Morse (2004) suggested that a size bias in cannibalized males is likely in M. vatia.
REVIEW
should result in differential reproductive success for males,
(4) the female bias should be associated with specific male
morphological or behavioural trait(s), and (5) the variation among males in characteristics used by females to discriminate should be heritable.
Evidence that, in spiders, females respond differently to
certain conspecific males and that discrimination is based
on some measurable morphological or behavioural trait
is available for only four species. Elgar (1992) regarded
reported sexual cannibalism of smaller males in Metellina
segmentata (Rubenstein 1987) as anecdotal, presumably
because of the lack of formal statistical analysis. However,
using data from the original paper (Figure 9 in Rubenstein
1987), we confirmed that smaller males were cannibalized
at a significantly greater frequency than larger males (number cannibalized: small 11/25; large 0/15; Fisher’s exact
probability test: P ¼ 0.003). In Araneus diadematus, females
were more likely to cannibalize small males (Elgar & Nash
1988) and those that engaged in lower premating pseudocopulation rates (mock copulation without sperm transfer).
However, pseudocopulation rate was not correlated with
male size and without this association it is difficult to see
how selecting for high rates would lead to small males being
cannibalized preferentially. The basis of the differential cannibalism in these two species, therefore, remains unclear
and we cannot determine direct or indirect mate choice. Female Schizocosa ocreata preferred to mate with males with
enlarged secondary sexual characters (leg tufts) but sexual
cannibalism was not related to this feature (Persons &
Uetz 2005). Female fishing spiders, Dolomedes fimbriatus,
did not differ in their attack behaviour towards males
(Arnqvist 1992; Andrade 1998); the attack success of females, however, was negatively related to male size because
larger males escaped or avoided capture. This seems to be
a clear case of indirect mate choice. Premating sexual cannibalism has been noted in various other species but without
evidence of either direct or indirect mate choice (Table 1).
Returning to the remainder of Eberhard’s (1996) criteria,
confirmation of PRE-SC resulting in differential mating
success under natural conditions is restricted, reflecting
a continued (Elgar 1992) paucity of field studies on sexual
cannibalism (Table 1). Obtaining evidence that fulfils the
criterion of female discrimination resulting in differential
reproductive success for males is more troublesome. In so
far as PRE-SC limits the lifetime reproductive success of
cannibalized males the criterion is fulfilled; however, the
actual reproductive success of surviving and mated males
is often not available. Furthermore, information on the
heritability and long-term fitness benefits (e.g. Elgar &
Nash 1988) of the outcomes of female mate choices (preferences) is also absent. The mate rejection, female mate
choice hypothesis for PRE-SC remains to be rigorously or
specifically tested experimentally; most of the evidence
for or against it comes from wider studies of courtship,
mating or sexual cannibalism.
ABSOLUTE AND RELATIVE SIZE
If an apparent discrimination of smaller males is simply
a by-product of the relative physical attributes of males
and females (Fox 1975; Polis 1981), then the idea of sexual
selection resulting from female preferences must be rejected. Indirect mate choice may then be the explanation
for the size-dependent nature of PRE-SC. Both mechanisms would result in selection on male size since larger
males survive the selection event better than smaller
males. The two models, however, make very different assumptions about the ability of females to discriminate between males (Johnson 2001), and the behaviour of males
and females is crucial in distinguishing between them.
Females attacking courting males and males attempting
to escape are two distinct facets of PRE-SC. Information
about the aggressive tendencies of females towards courting males is essential and will be a major factor in
unravelling the causes of sexual cannibalism and the
role of mate choice. If females attack all males, for
example as in D. fimbriatus (Arnqvist 1992; Arnqvist &
Henriksson 1997), then size-dependent sexual cannibalism is probably the by-product of indirect mate choice.
That female attack success, although not attack behaviour,
varies with male size in D. fimbriatus suggests that larger
males are better able to avoid/escape the female’s attacks.
We should ask whether females are consistent in the size
class of males rejected (although variation in choice
criteria might be expected under variation in environmental circumstances, for example Elgar & Nash 1988;
Newman & Elgar 1991) or whether the rejection criteria
vary with the difference in size between the interacting
pair. If sexual cannibalism varies with the relative size of
the interacting males and females, then we propose that
direct mate choice may be ruled out as an explanation
of PRE-SC. Thus, the outcome of courtship interactions
(cannibalism or mating) would be determined by the relative vulnerability of the male to the predatory advances of
the larger female. However, if females are consistent in
their choice of males for cannibalistic attack and mate
with the others (at least under similar environmental conditions), and the absolute size of males determines the
outcome of courtship interactions, then direct mate
choice is a more likely explanation. The distinction between the effects of absolute size and relative size may,
therefore, be crucial in the determination of direct or indirect mate choice via PRE-SC. The data necessary to distinguish between these hypothesis, however, are rarely
available. When sexual size dimorphism (ratio of female
to male size) was low in D. triton, males were more likely
to escape attacks from females and go on to mate successfully (Johnson 2005b). Relative size of the pair (along with
female body condition) determined the probability of successful PRE-SC in S. ocreata (Persons & Uetz 2005),
although PRE-SC did not represent a mechanism of mate
choice. The paucity of information about the effects of relative size on PRE-SC is in stark contrast to the plethora of
studies on aggressive behaviour and assessment spawned
by game theory, where relative size and its effects are commonly investigated (Taylor & Elwood 2003).
Relatively small females might be incapable of making
and imposing a choice because they do not possess the
ability to reject all unpreferred males. Males may be able to
repel attacks and engage in a war of attrition, ultimately
resulting in the female abandoning her attacks. As
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ANIMAL BEHAVIOUR, 71, 3
a consequence, a female attempting to select males
actively might show cannibalism based on relative size,
even though the choice is based on absolute size. Effectively, small females may not be capable of rejecting many
males, but larger females may have a greater opportunity
to make a choice through sexual cannibalism. Even in this
case, our prediction about different female behaviour,
with regard to absolute and relative size of males and
females, should still hold. A more formal theoretical
approach might be needed, however, to consider the
outcome of the hypothesized war of attrition.
Examination of female motivation to attack and capture
males, rather than actual ability to do so (which may be
constrained by various factors including size-related male
escape and female capture abilities) might prove a profitable approach to the investigation of the tendency of
females to cannibalize different courting males. An interruption technique has been successfully used to probe
the motivation of invertebrates in aggressive encounters
(Elwood et al. 1998) and could be adapted to the problem
of sexual cannibalism.
Owing to the lack of information about the effects of
relative or absolute size of the interacting pair on the
occurrence of sexual cannibalism, the theory that PRE-SC
forms a mechanism of direct mate choice must be treated
with caution. Mate choice in the classical/direct sense,
where females actively select males on the basis of some
morphological or behavioural characteristic, remains to be
demonstrated clearly with regard to PRE-SC. There can be
little doubt, however, that indirect mate choice may arise
via sexual cannibalism on males that are small relative to
females, resulting in size-dependent patterns in PRE-SC, as
a consequence of female foraging strategies, and that both
scenarios will produce selection for increased size in males
to avoid sexual cannibalism.
POSTCOPULATORY SEXUAL CANNIBALISM
Males are generally expected to be under stronger selection to control fertilization than are females, because their
potential loss from failing to fertilize any eggs is greater
than that for females restricted to mating with males of
inferior quality (Parker 1984; Stockley 1997; Simmons &
Siva-Jothy 1998). However, the polygamous nature of
many mating systems (but see Fromhage et al. 2005),
and the resulting possibility of sperm competition, offers
an opportunity for females to manipulate and control paternity. POST-SC offers a similar potential for females to
influence copulation duration, sperm transfer and
paternity.
Eberhard (1996) proposed a modification to the mate rejection hypothesis for cannibalism occurring during or after copulation, that the termination of copulation by
initiating sexual cannibalism allows females to control fertilization of their eggs. Here, sexual cannibalism forms
a postcopulatory mate choice mechanism, by which females can influence which males father their offspring.
The specific conditions that must be met before cryptic
choice can be determined are controversial (Birkhead
1998, 2000; Birkhead & Møller 1998; Eberhard 2000;
Pitnick & Brown 2000). Paramount among the prerequisites, in terms of POST-SC, are the requirements that (1)
variation in paternity is attributable to a male characteristic (e.g. size or specific behaviour) and (2) females are able
to control this variation in paternity by varying the onset
of cannibalism (Andrade 1998). In simple terms, preferred
males may be allowed to copulate for longer before cannibalism is initiated or if cannibalism occurs and these males
will transfer more sperm and should, therefore, sire more
offspring. Crucially, this argument depends on the mechanism of sperm transfer used and hinges on a positive
relation between copulation duration and sperm transferred. This positive relation is known for some species
(e.g. Schneider et al., 2006) but not for others (Bukowski
et al. 2001; Snow & Andrade 2004; see below).
Satisfying the criterion that establishes female control of
paternity, that some variation in paternity is attributable
to the female, is often problematic, because it is difficult to
separate the contributions of the two sexes to variation in
paternity. Females often favour the same traits (e.g. large
size) that provide males with advantages over their rivals.
Certainly, if the female is to exert control of paternity by
sexual cannibalism, cannibalism, copulation duration and
paternity should be related (Schneider & Elgar 2001).
In the past decade or so there have been significant
advances in the study of POST-SC, especially from the
standpoint of male adaptive (suicidal or sacrificial) behaviour in spiders (e.g. Forster 1992; Andrade 1996, 1998; Andrade et al. 2005). There has been little investigation of
mate choice via POST-SC; however, studies on mating behaviour and sperm competition in sexually cannibalistic
spiders (see Elgar 1998 for a review of sperm competition
in arachnids) provide data that can be evaluated in this
light (Table 2). Information is lacking, however, that
would allow examination of female mate choice in mantids other than that there is potential for cryptic choice
(Lawrence 1992).
Where POST-SC has been studied, there is a lack of
consistent evidence that differential POST-SC occurs.
Where differential female behaviour has been identified,
this may not translate into differential POST-SC or increased paternity for cannibalized males. POST-SC can
either prolong or shorten the duration of copulation for
cannibalized males compared to survivors (Table 2) and
copulation duration may be an important determinant
of fertilization success for males. While POST-SC was not
discriminatory on the basis of any male trait or behaviour
measured in the orb-weaving spiders Argiope aurantia (in
which males invariably spontaneously expire after their
second palpal intromission), Argiope bruennichi (Foellmer
& Fairbairn 2003, 2004; Fromhage et al. 2003; but see
Schneider et al. 2006) and Nephila fenestrata (Fromhage
& Schneider 2005), it still resulted in increased copulation
duration. Female mate choice may, therefore, be an unlikely explanation for POST-SC in these species.
We must be cautious, however, in regarding increased
copulation duration as a benefit of POST-SC. In species
where males attempt to make two pedipal intromissions
with a female (e.g. A. aurantia: Foellmer & Fairbairn 2003,
2004; A. bruennichi: Schneider et al. 2006), POST-SC during or after the first insertion may drastically shorten
REVIEW
Table 2. Summary of evidence for cryptic female choice through postcopulatory sexual cannibalism (POST-SC)
Criteria for mate choice
Species
Argiope
keyserlingi
Differential
sexual
cannibalism
and comment
Females delayed onset
of POST-SC on smaller
second mates*; no evidence
that small males escape
POST-SC better
Argiope
ND; POST-SC in up to
bruennichi 80% of matings
Argiope
Larger males with longer
aurantia
legs more likely to be
attacked during first palpal
insertion (but not translated
into differential POST-SC)y
Nephila
Smaller males cannibalized
plumipes
less than larger males when
mating with virgin
(but not mated) females
(but not translated to
increased
fertilization success)
Nephila
ND (but females attacked
edulis
larger males more often in
first matings, while NS trend
for cannibalized males to
weigh less); mating
advantage for
smaller malesx
Nephila
ND, no correlation between
fenestrata
male mass, size or age with
occurrence of cannibalism
Latrodectus ND, no size, mass or
hasselti
condition differences
detected between
cannibalized and
noncannibalized males,
but males cannibalized on
their second palpal insertion
obtain longer copulations
and higher paternity
Latrodectus Small males cannibalized
revivensis
more when attempting
second insemination into
second spermatheca
Schizocosa Smaller males (poorer condition
ocreata
with relatively small leg tufts)
more likely to be cannibalized
by larger (older) females
Paternity
attributable
to male
character?
Female ability
to control
variation in
male paternity?
Effect of sexual
cannibalism on
copulation duration
Source
Y
Y
[ by delaying
POST-SC*
Elgar et al. 2000
N
?
[ with POST-SC
?
?
[ with POST-SC(?)
(palpal insertion
duration
[ with cannibalism)
Fromhage et al. 2003;
Schneider et al. 2006
Foellmer & Fairbairn
2003, 2004
?
(But males
cannibalized by
nonvirgin females
increased paternity
compared to
surviving males)
?
Y(?)z
[ with POST-SC
Elgar & Fahey 1996;
Schneider & Elgar 2001;
Schneider et al. 2001
?
None
Uhl & Vollrath 1998;
Schneider et al. 2000;
Elgar et al. 2003
N
?
Fromhage &
Schneider 2005
?
But increased
paternity for
cannibalized
males
Y(?)
May control
copulation
duration with
onset
of POST-SC
[ with POST-SC
(but with no
effect on paternity)
[ with POST-SC
?
?
?
B. Berendonck, personal
communication
?
?
?
Persons & Uetz 2005
Forster 1992; Andrade
1996, 2003; Andrade &
Banta 2001, 2002;
Snow & Andrade 2004
[ ¼ increased. See Table 1 for other abbreviations.
*Applies only to second mates where two males were cannibalized (not to all males cannibalized).
ySubject to statistical provisos with regard to multiple tests (effect not significant when corrected for multiple tests) males spontaneously die at
the second palpal insertion in this species (Foellmer & Fairbairn 2004).
zCannibalized males (mating with nonvirgin females) achieved longer copulations and increased paternity while not transferring more sperm;
therefore females may make some form of sperm choice under these circumstances.
xAdvantage in fertilization success for smaller males unrelated to POST-SC (small males copulated for longer and fertilized more eggs than large
males).
copulation duration (and also leave an unfilled spermatheca available for rival males). Furthermore, even in species that insert only one pedipalp, escaping affords the
opportunity for further mating, which would considerably
increase a male’s fitness over a cannibalized male.
Elgar et al. (2000) succeeded in demonstrating female
control of paternity separate from the confounding effects
of intrasexual selection in the St Andrew’s cross spider
Argiope keyserlingi. In this species, females terminate copulation by detaching males from their genital openings,
485
486
ANIMAL BEHAVIOUR, 71, 3
wrapping and then killing them. Females copulating with
relatively smaller males delayed sexual cannibalism, allowing prolonged copulations that resulted in more eggs
being fertilized (Table 2). The apparent lack of evidence
of an ‘inherent’ advantage to small males of escaping cannibalism and, thereby, prolonging copulation duration,
supports direct mate choice explanations of sexual cannibalism in A. keyserlingi, according to Elgar et al. (2000). Additional evidence of a mating advantage for smaller males
is also present in the cannibalistic spider Nephila edulis
(Schneider et al. 2000). Why females of these species
should favour smaller, rather than larger males, however,
is unclear. Sexual selection may favour small size under
conditions of scramble competition, when mate searching
entails climbing (Moya-Laraño et al. 2002) or if these
males mature earlier and have a higher probability of
locating a female than later maturing males (Andersson
1994; see also Vollrath & Parker 1992). Although whether
or not and even why sexual cannibalism by females influences this is unclear (Table 2).
POST-SC in Latrodectus revivensis has been proposed as
a female strategy to counter monopolization of both
(paired) sperm storage organs by males (B. Berendonck,
personal communication). In this desert widow spider, females ensure they obtain sufficient quantities of sperm to
fertilize their eggs by allowing males to inseminate one
spermatheca. Thereafter, females practise size-dependent
sexual cannibalism on small males, denying them the opportunity to copulate more than once. Thus, females can
practise sequential choice via POST-SC, allowing only larger
or better-quality males to fill the second spermatheca, or
even replace the first ‘inferior’ male’s sperm from the already filled sperm storage organ. Specific details of female
discriminatory behaviour, other than the gross effects of
cannibalism, however, remain unclear.
In N. plumipes, females cannibalize large males
(Schneider & Elgar 2001). Although they did not deliver
significantly more sperm to the female’s spermathecae,
males cannibalized by nonvirgin females secured a higher
proportion of paternity than males that survived mating
(Schneider & Elgar 2001). We interpret these results as being suggestive of sequential (cryptic) mate choice, although the basis of any discrimination and choice by
the female is unclear. However, Schneider & Elgar (2002)
suggested that sexual cannibalism in N. plumipes is a consequence of female life history strategies and attributed it to
an increased foraging voracity of females that matured at
a smaller size and in poorer condition. Mate choice may
represent only a contributing factor, therefore, to the evolution and maintenance of POST-SC in N. plumipes.
Returning to the criteria for female choice, there is
evidence of POST-SC leading to increased paternity for
cannibalized males (Table 2). In only one case (Elgar et al.
2000), however, has variation in POST-SC and paternity
been associated with or attributable to an identifiable
male trait. Whether females can control the variation in
paternity that differential POST-SC produces, by varying
the onset of sexual cannibalism, is debatable. In three
cases (A. keyserlingi, L. hasselti and N. plumipes), females
may be able to manipulate or otherwise influence the paternity of their broods through postcopulatory control of
sperm uptake or transfer. Nevertheless, females of some
sexually cannibalistic spiders do appear to practise cryptic
female choice but the exact mechanism of cryptic choice
is debatable, requires further investigation and may not
be a simple reflection of the increased copulation time
that POST-SC seems to provide some males (Table 2).
Only further detailed examination of the mechanism of
transfer of sperm and other extragametic substances in
the ejaculate by males (e.g. Snow & Andrade 2004), and
their usage by females, will determine whether females
are using sperm from ‘preferred’ males differentially and
whether sexual cannibalism can influence this process.
Certainly, in the redback spider L. hasselti, there would appear to be a linkage between sperm uptake and POST-SC.
However, while no clear mechanism for female control
of paternity has been shown, possible male influences
(e.g. sperm manipulation) cannot be ruled out. Snow &
Andrade (2004) attributed reduced postmating receptivity
in cannibalistic L. hasselti females to manipulation by
males through substances transferred in the ejaculate.
Examination of possible copulatory courtship (Eberhard
1996) by males may enable traits/activities to be identified
that distinguish between cannibalized and surviving
males and can be related to differences in paternity. Courtship duration itself may constitute a signal used by female
L. hasselti to discriminate against males. Andrade & Banta
(2001) experimentally manipulated courtship duration
(short or long) and found that males in the short courtship group had lower mating success and experienced
more female rejection behaviour. They argued that this
represented female discrimination, via a form of endurance test. In addition, as indicated earlier, POST-SC in
this species increases male paternity (Andrade 1996) and
this occurred more often with males in the long courtship
group.
MATE CHOICE AND SEXUAL CANNIBALISM
Sexual cannibalism is thought to drive the evolution of
various morphological and behavioural traits, particularly
in spiders (reviewed in Robinson 1982; Elgar 1992) and
mantids (e.g. Maxwell 1998, 1999). The power of sexual
cannibalism as a selective agent derives from the drastic
and instantaneous effect it can have on male survival
and prospective mating opportunities (Persons & Uetz
2005) and is manifest in the cautious nature of courtship
in cannibalistic species (Morse 2004). Sexual cannibalism
may be a strong and important selective force even
when it occurs infrequently (Morse 2004).
There may be more than one cause for the evolution
and maintenance of sexual cannibalism for any species
(e.g. Johnson 2001; Morse 2004; Elgar & Schneider 2005);
various hypotheses are not mutually exclusive and may affect life history stages differently (Johnson 2001). Thus,
mate choice, either overt or cryptic, clearly may not be
the only explanation for sexual cannibalism. Mistaken
identity (e.g. Robinson 1982; Gould 1984; Elgar 1992;
Arnqvist & Henriksson 1997), hunger levels (Newman &
Elgar 1991; Andrade 1998; Samu et al. 1999; Herberstein
et al. 2002) and complicit cannibalism (male sacrifice) to
REVIEW
ensure paternity (Andrade 1996) contribute to the evolution and maintenance of sexual cannibalism. Even
though male L. hasselti derived a paternity advantage (Andrade 1996) from sexual cannibalism, their fate was still
determined by the females’ nutritional state (Andrade
1998). However, feeding history was not important in determining the likelihood of PRE-SC in fishing spiders
(Johnson 2005a) or the jumping spider Phidippus johnsoni
(Jackson 1980). In praying mantids, hungrier females were
more likely to cannibalize males than better-fed females
(Liske & Davis 1984, 1987; Birkhead et al. 1988; Kynaston
et al. 1994). Previous juvenile experience may also affect
PRE-SC in adult females. Johnson (2005a) showed that
in D. triton, juveniles’ experience of cohabitation with
adult males (an indicator of the availability of males/future mating opportunities) increased the likelihood of
PRE-SC (see also Hebets 2003).
Notwithstanding the possibility that cannibalism may
act in mate choice, it seems unlikely that mate choice has
been the most important driving force behind the evolution of sexual cannibalism. There are numerous opportunities for females to influence the paternity of their young
without resorting to killing and consuming males. However, even if the initial function of sexual cannibalism was
to obtain food it might then have become a mechanism of
mate choice; this may have added to the benefits of
cannibalism (see Persons & Uetz 2005) and may remain
an important influence on its maintenance in cannibalistic species, imposing directional selection on male morphology and behaviour, particularly in polygamous
mating systems. Indeed, this may be why we find more evidence for indirect than direct mate choice though sexual
cannibalism.
REVERSED SEXUAL CANNIBALISM
Reversed sexual cannibalism may occur particularly in
species in which males are larger than females. Male
amphipods (Gammarus pulex and G. duebeni celticus) can
cannibalize smaller females, especially at the moult
when they are most vulnerable (Dick et al. 1993; Dick
1995). Similarly, female brachyurid paddle crabs, Ovalipes
catharus, are also vulnerable at their moult when their
shell is soft and may be subject to cannibalism during or
after mating (Haddon 1995). Male Ichthyoxenus fushanensis (an isopod body cavity parasite of fish) may cannibalize
females during or after mating (Tsai & Dai 2003). In the
case of Gammarus species, sexual cannibalism would
seem to be opportunistic in nature and not have any
mate choice implications. Not all females produce eggs
at each moult, however, and males may preferentially cannibalize females without eggs, but this has not been tested.
Cannibalism of the female may be similarly opportunistic
in O. catharus; however, males seem to avoid cannibalism
of females they have recently inseminated. Resource limitation may explain reversed sexual cannibalism in I. fushanensis (Tsai & Dai 2003), but requires further empirical
investigation.
There is anecdotal evidence of a male eating a female in
the mantid Tarachodes afzeli (Edmunds 1975). Unusually
for a spider, males of the water spider Argyroneta aquatica
are larger than females, prefer to approach larger females
and may cannibalize females smaller than themselves
(Schütz & Taborsky 2005). Females prefer to mate with
larger males, but are more wary of them, fleeing from
them more often than from males closer to their own
size. The vulnerability to sexual cannibalism of the victim
appears to be key in A. aquatica and would suggest that an
indirect mechanism, rather than direct mate choice,
produces the size-dependent reversed sexual cannibalism
observed. With the male-biased size dimorphism present
and the relative size of the female determining her vulnerability, more smaller females will tend to be cannibalized.
Presumably in all cases of reversed sexual cannibalism
there is little prospect of the female producing offspring
for the male or the need for food exceeds the fitness
gain from mating. Since males tend towards promiscuity
to maximize their reproductive output, it is difficult to
envisage mate choice through sexual cannibalism for
males, unless the sex roles are reversed or the sex ratio is
extremely female biased, offering males the opportunity
to be choosy. Costly courtship and mating may promote
male mate choice (Bonduriansky 2001), but there are no
clear examples of male mate choice via reversed sexual
cannibalism.
CONCLUSIONS AND RECOMMENDATIONS
There are clear gaps in our knowledge with regard to mate
choice and sexual cannibalism that need to be filled.
Research on mantids has concentrated on male behaviour
and avoidance of sexual cannibalism and sheds little light
on mate choice (e.g. Maxwell 1998, 1999), although the
escape behaviour and copulation success of attacked males
argues against mate choice (Liske & Davis 1987; Maxwell
1999). Sexual cannibalism also occurs in scorpions but it
is not as prevalent as previously presumed and appears
to play no role in mate choice in this group (Polis & Sissom 1990; Peretti et al. 1999; Benton 2001). While there
is no similar doubt over the prevalence of sexual cannibalism in spiders, its function remains the subject of debate.
The occurrence of mate choice through sexual cannibalism in arachnids is supported by some empirical
evidence, manifested by size dependency in PRE-SC
(Rubenstein 1987; Elgar & Nash 1988; Arnqvist 1992)
and by cryptic choice in POST-SC (Elgar et al. 2000). There
is little evidence, however, for direct mate choice by female spiders or scorpions (Peretti et al. 1999) in PRE-SC.
Rather, sexual cannibalism may be economic in nature,
and may reflect opportunism and voracity in female predatory behaviour, the effects of relative size and the reduced
ability of smaller males to fend off larger females. However,
whichever mechanism is in operation, the same directional
selection on male size will still be exercised. We have presented a framework by which research may be advanced
and direct and indirect choice distinguished that requires
empirical and theoretical examination.
There remains a paucity of evidence and rigorous
investigation of the mate rejection hypothesis and PRESC. This represents a poor state of affairs in such an
487
488
ANIMAL BEHAVIOUR, 71, 3
important predatory animal group as arachnids, particularly when the historical notoriety of sexual cannibalism
in spiders is taken into account (Darwin 1871). Furthermore, greater emphasis must be placed on field rather
than laboratory study, and on determining the longterm fitness benefits of the apparent mate choices of females, since a direct benefit of cannibalism is known
only for a few species (e.g. Elgar 1992; Fromhage et al.
2003; Elgar & Schneider 2005). Further investigation of
environmental (e.g. feeding rate, male encounter rate, effects of population density) and especially genetic (e.g.
population differences) influences on female behaviour
towards courting males is also necessary.
More detailed examinations of females’ behaviour towards males, particularly their motivation to attack different males, and investigations of the effects of relative size
and the ability of males to withstand the advances of
females, are required if we are to understand sexual
cannibalism in spiders better and to assess the role of
mate choice. The direct causes of size dependency in sexual
cannibalism are unclear and the examination of the costs
and benefits of this behaviour has been inadequate. Future
investigations should focus on ameliorating these inadequacies, for example, by comparing size at emergence,
foraging ability, growth rate and ability of the offspring of
preferred males to avoid cannibalism with those of the
offspring of males that would ordinarily be cannibalized.
Genetic benefits from mate choice may be context
dependent and, therefore, selection pressures on female
mate choice will vary across environments. Qvarnström
(2001) proposed that a profitable approach to the evolution of mate preferences would be to examine them in
terms of optimality, as in other reproductive decisions.
The optimal mate choice might be dynamic, rather than
constant, and be determined by a variety of factors. Fluctuations in internal factors (e.g. reproductive status of females and their sperm store) and external factors (e.g.
food supply and male encounter rate) should all influence
the female’s decision to mate or cannibalize a male.
Evidence suggests female choice by POST-SC in several
spiders, with cannibalized males achieving longer copulations and potentially greater paternity than noncannibalized males (but see above). It has been assumed that
longer copulation allows increased quantities of sperm to
be transferred to the female, with a resultant increase in
paternity. However, this has been called into question
(Bukowski et al. 2001; Snow & Andrade 2004). While cryptic choice would seem likely, the mechanism by which it is
achieved is unclear. This highlights the need for detailed
investigation of the mechanics of sperm transfer and of
mechanisms by which females may control paternity of
their offspring. In general, research should be focused on
examining the various criteria for demonstrating mate
choice in both PRE-SC and POST-SC. In the latter, it is important to examine both variation in paternity with POSTSC and the mechanism by which it may be achieved.
Where POST-SC leads to variation in male paternity, but
no obvious mechanism for female control presents itself,
further detailed examination of male and female copulatory
behaviour is advocated. Little is known about reversed
sexual cannibalism, although it appears to be associated
with opportunism and vulnerability in the victim and its
occurrence may be unrelated to mate choice.
There is a growing consensus that sexual cannibalism
may have multiple explanations and could have developed in response to a variety of lifestyle and phylogenetic
constraints (e.g. Morse 2004; Elgar & Schneider 2005).
Mate choice may be just one of these explanations. We
caution that alternative explanations of the evolution of
sexual cannibalism, especially economic cannibalism
and life history implications, be considered before explanations based on mate choice, especially direct mate choice,
are adopted.
Acknowledgments
Mark Elgar, Jutta Schneider, Michael Taborsky, Chad
Johnson, Maydianne Andrade, Bernhard Huber and Tina
Berendonck kindly provided information on unpublished
research. We are grateful to Leigh Simmons, Mark Elgar,
Tina Berendonck, Mark Briffa, Gillian Riddell and Liz
Pothanikat for their comments on the manuscript. J.P.
acknowledges support from the UK Biotechnology and
Biological Sciences Research Council (S20306) and a Research Fellowship from the Institute of Irish Studies
(QUB). C.M. thanks the UK Natural Environment Research Council (GR3/12871) and the Isle of Man Government for support.
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