1Effects of male and female personality on sexual cannibalism in the Springbok
2mantis
3Pietro Pollo1, *, Nathan W. Burke2 and Gregory I. Holwell2
4
5¹ Evolution and Ecology Research Centre, School of Biological, Earth and
6Environmental Sciences, University of New South Wales, Kensington, Sydney 2052
7NSW, Australia
8² School of Biological Sciences, University of Auckland, New Zealand
9* Corresponding author: [email protected]
10
11Abstract
12Behaviours that are consistent across contexts (also known as behavioural syndromes)
13can have evolutionary implications, but their role in scenarios where the sexes
14conflict, such as sexual cannibalism, is poorly understood. The aggressive spillover
15hypothesis proposes that cannibalistic attacks during adulthood may depend on female
16aggressiveness during earlier developmental stages, but evidence for this hypothesis is
17scarce. Male activity may also influence sexual cannibalism if males approach
18females quickly and carelessly, yet this has not been explored. Here we use the
19Springbok mantis, Miomantis caffra, to explore whether male activity levels and
20female aggressiveness can explain high rates of sexual cannibalism prior to
21copulation. We show that male and female personality traits affect male mating
22decisions, but not sexual cannibalism. Females that were aggressive as juveniles were 23not more likely to cannibalize males when adult, but these females were approached
24by males more frequently. More active males were more likely to approach females,
25but they were neither faster at doing so nor were they more likely to be cannibalized.
26We also found that size and age influenced mating decisions of both sexes: young
27females were more like to cannibalize males while young and large males took longer
28to approach females. Taken together, our results suggest that several traits, including
29personality, play a role in sexual encounters in M. caffra. Our study further highlights
30the importance of examining the traits of both sexes when assessing mating dynamics,
31especially in the context of sexual cannibalism.
32
33Keywords: animal temperament, male mate choice, mating decisions, sexual conflict.
34
35Introduction
36 Animals often exhibit consistent individual differences in behaviour, also
37known as personalities (Réale et al. 2007). Personality presents as varying levels of
38behavioural expression between individuals (behavioural types), such as “more
39aggressive” or “less aggressive” (Sih, Bell, Johnson, et al. 2004; Sih, Bell, and
40Johnson 2004). Behavioural types that correlate across contexts or situations are
41known as behavioural syndromes, and are often characterised by trade-offs where a
42behavioural type is advantageous in certain contexts but detrimental in others (Sih,
43Bell, Johnson, et al. 2004; Sih, Bell, and Johnson 2004). Behavioural syndromes are
44thus predicted to have ecological and evolutionary implications, such as in population 45dynamics and species interactions (Sih et al. 2012; Wolf and Weissing 2012; Sih
462017).
47 One evolutionary context in which behavioural syndromes may play a role is
48sexual conflict (Royle et al. 2010). Sexual cannibalism (i.e. attacking and consuming
49a prospective or realized mate) represents an extreme case of sexual conflict as it
50precludes the victims’ future reproductive success (Elgar 1992). This behaviour
51occurs in several taxa and is usually performed by females (Elgar 1992; Elgar and
52Schneider 2004; but see Aisenberg et al. 2011). Personality traits and behavioural
53syndromes may provide insights into the causes and consequences of sexual
54cannibalism. More specifically, they may help to explain why females attack males
55and whether some males are more likely to become the victims of sexual cannibalism
56than others.
57 The aggressive spillover hypothesis is one of the best-known examples of a
58behavioural syndrome. It proposes that females that are more aggressive towards prey
59as juveniles tend to be more cannibalistic towards prospective mates as adults
60(Arnqvist and Henriksson 1997). This is because aggressiveness that is advantageous
61in a foraging context when females are developing may “spill over” to the mating
62context in the adult stage. Therefore, according to this hypothesis, sexual cannibalism
63prior to mating would be maladaptive if it increases the chance of females dying as
64virgins (Arnqvist and Henriksson 1997). There is some support for the aggressive
65spillover hypothesis (Johnson and Sih 2005; Berning et al. 2012; Rabaneda-Bueno et
66al. 2014; but see Kralj-Fišer et al. 2012; Kralj-Fišer et al. 2013; Kralj-Fišer et al.
672016), indicating that behavioural syndromes play a role in the evolution of sexual 68cannibalism in at least some taxa. However, the extent to which aggressive spillover
69provides a general explanation for sexual cannibalism remains unclear as research on
70this topic has been carried out almost exclusively on spiders, and other hypotheses
71have received more attention (reviewed in Elgar and Schneider 2004).
72 While effects of personality on sexual cannibalism have primarily focused on
73females, male personality traits have been relatively neglected in sexually
74cannibalistic species (but see Kralj-Fišer et al. 2012). Male personality could
75influence sexual cannibalism in the context of female mate choice (Elgar and
76Schneider 2004), and activity is a personality trait that could be especially important.
77Greater male activity may lead to increased attack by females if males approach their
78potential mates too rapidly or incautiously. For example, in the spider Nephila
79fenestrata, males that mate with idle females suffer more injuries than males that mate
80with feeding females (Fromhage and Schneider 2005), suggesting that inopportune
81approaches can be costly. Similarly, males of the praying mantis Pseudomantis
82albofimbriata approach females with great caution to avoid being captured (Barry et
83al. 2009). However, the extent to which male activity levels mediate the likelihood of
84sexual encounter and female attack in cannibalistic taxa remains unclear.
85 The mantis Miomantis caffra is ideal to investigate the causes and
86consequences of pre-copulatory sexual cannibalism because females of this species
87regularly attack approaching males (Walker and Holwell 2016). Moreover, M. caffra
88males avoid engaging with females in approximately 70% of experimental mating
89trials (Fea et al. 2013; Walker and Holwell 2016; Burke and Holwell 2021a),
90suggesting that males are cautious in deciding when to approach females (Burke and 91Holwell 2021b). Here, we investigate whether females and males present personalities
92for aggression and activity, respectively, and whether these individual behavioural
93types form behavioural syndromes. For females, we tested whether juvenile
94aggressiveness towards prey in a foraging context influences the propensity to
95cannibalize males in a mating context. For males, we examined whether higher
96activity in an exploratory context correlates with faster and more frequent approaches
97towards females in a mating context. We also explored whether male activity
98predicted the frequency of mating versus cannibalism.
99
100Material and methods
101Study species, rearing and maintenance
102 Miomantis caffra is a South African mantis introduced to New Zealand more
103than four decades ago, commonly found in urban areas (Ramsay 1984). We collected
104M. caffra juvenile individuals in Auckland, New Zealand, between December 2019
105and February 2020. We maintained them in individual overturned 700 mL plastic
106cups with the bottom cut out and covered with mesh to allow air flow. Following a
107similar feeding regime as Walker and Holwell (2016), we fed juveniles and adult
108males with three CO2-anesthetised flies (Musca domestica) three times per week.
109Juvenile females received the same diet up until they moulted to the last instar before
110adulthood (i.e. subadult stage) at which point their feeding regime changed as part of
111the aggressiveness trials (see below).
112
113Female aggressiveness trials 114 To test whether female aggressiveness is consistent across ontogeny, we
115assessed female aggressiveness (N = 35) every weekday during subadult and adult
116stages, 8 to 17 times per stage (754 trials in total). Trials took place in the same
117containers in which individuals were housed. Females were fed a single fly an hour
118before trials to standardise their hunger state. Females that refused to eat the fly
119(approximately 6% of all trials, N = 49/754) were excluded for that day. For each
120trial, we placed two CO2-anesthetised flies in enclosures and noted female latency to
121first capture (in seconds) over a period of one hour. If neither fly was captured
122(approximately 7% of all trials, 50/754), we recorded latency as 3600 secs (one hour).
123We subtracted each latency score from the trial duration (3600 secs) to obtain a
124measure of female aggressiveness. Larger scores corresponded to more rapid capture
125times which we interpreted as indicative of higher aggressiveness (similar to Johnson
126and Sih 2005). Flies that were not eaten within the observation period were left for
127females to capture later. This allowed females to eat a similar number of flies every
128day regardless of their latency to capture prey. Trials in which one or both flies failed
129to move noticeably during the observation period (approximately 2% of all trials,
13016/754) were excluded from analyses.
131
132Male activity trials
133 To assess male activity, we took adult males (N = 39) and placed them inside
134plastic tubes approximately 40 cm long and four cm in diameter positioned vertically
135and enclosed at the top. All males started at the base of the tube and could freely walk
136its length. In pilot trials in which we observed males for 90 min continuously, 137individuals rarely moved. We thus opted to record distance travelled (mm) after a
138period of eight hours (overnight). We performed five trials per male over a period of
139five consecutive days to assess intra-individual consistency and inter-individual
140differences in activity.
141
142Mating trials
143 To conduct mating trials (N = 35), we used mesh enclosures (30 x 30 x 30 cm)
144each containing brown cardboard flooring and a 20 cm branch of Tecoma capensis
145positioned in a 50 mL jar of damp sand. Females were fed three flies the night before.
146In the morning, we placed individual females on the ends of the Tecoma branches and
147allowed females to acclimate in their enclosures for 1 hour prior to the introduction of
148males. Males were gently placed on the jar facing towards the female. We kept pairs
149together for up to 48 hours, observing them every hour in the first 12 hours, every two
150hours from the 24th hour to the 36th hour, and finally at the 48th hour. We noted the
151occurrence of mating by the presence of a spermatophore plugged in the female’s
152gonopore or ejected onto the bottom of the enclosure, and the occurrence of sexual
153cannibalism by the absence of males.
154
155Statistical analysis
156 We calculated the repeatability of aggressiveness in females and activity in
157males to evaluate whether these behaviours were consistent within and between
158individuals. Repeatability refers to the proportion of behavioural variation attributed
159to among-individual variance (Bell et al. 2009). Greater repeatability therefore 160indicates that within-individual variance is smaller compared with among-individuals
161variance (Bell et al. 2009) and can be interpreted as greater behavioural consistency
162through time. We calculated repeatability of female aggressiveness when females
163were subadult separately from when females were adults. We transformed
164behavioural measures to conduct repeatability analyses because the models from the
165rptR package (Stoffel et al. 2017) assume that data is normally distributed.
166Aggressiveness’ distribution was right-skewed, so we transformed it by square
167rooting the subtraction of its value from 3601 (maximum aggressiveness score plus
168one). Distance travelled in activity trials was left-skewed, so we log transformed it.
169We assessed the distribution of residuals using the package DHARMa (Hartig 2020).
170 To explore the factors that influence the likelihood of sexual cannibalism, we
171built a generalized linear model (GLM) with binomial error structures and a logit link
172function. In this model, the occurrence of sexual cannibalism in our mating
173experiments (binary variable) was the response variable. We included mean female
174aggressiveness in the subadult stage as an explanatory variable to test whether females
175with greater aggressiveness in the subadult stage would be more likely to cannibalize
176males in adulthood (Arnqvist and Henriksson 1997). We also included the mean
177distance travelled by males in activity trials as an explanatory variable to test whether
178females would be more likely to cannibalize males of a certain behavioural type. We
179also included in this model covariates that are known to influence sexual cannibalism
180likelihood (Elgar 1992; Elgar and Schneider 2004), such as age (days since females’
181final moult) and size (pronotum length) for both males and females . 182 We further verified some assumptions made by the aggressive spillover
183hypothesis (Arnqvist and Henriksson 1997) by investigating life-history patterns
184associated with female aggressiveness. Female aggressiveness is expected to be
185consistent across ontogeny, and more aggressive adult females should be larger
186because their high aggressiveness during the juvenile stage should allow them to
187capture more prey and grow more (Arnqvist and Henriksson 1997). Therefore, we
188calculated Spearman’s correlation between mean female aggressiveness in the
189subadult stage and three other variables: mean female aggressiveness in the adult
190stage, absolute growth (difference in pronotum lengths) and relative growth
191(difference in pronotum lengths divided by subadult pronotum length) between
192subadult and adult stages.
193 To explore the factors that influence male mating decisions, we built two
194generalized linear models (GLMs): one with binomial error structure and a logit link
195function and another with a Gaussian error structure (i.e. linear regression). The
196response variables in these models were, respectively, whether males approached
197females (binary variable) and, if they did, how long they took to approach them (in
198hours; continuous). Because it is possible that males modulate their approach
199depending on traits that increase sexual cannibalism likelihood, we included the same
200explanatory variables and covariates as in the female sexual cannibalism likelihood
201model (i.e. subadult female aggressiveness and male activity; age and size of both
202females and males; respectively).
203 All statistical analyses were performed in R (R Core Team 2020). We
204calculated repeatability and its 95% confidence interval using the function rpt from 205the package rptR (Stoffel et al. 2017), with 1000 parametric bootstraps and a Gaussian
206distribution. We used likelihood ratio tests (LRTs) that compared each fixed effect
207and covariate against the full model to assess their coefficients’ significance in GLMs
208using the function lrtest from the lmtest package (Zeileis and Hothorn 2002). All
209continuous explanatory variables were centred and standardized before model fitting.
210
211Results
212Female aggressiveness and male activity
213 Repeatability of female aggressiveness in the subadult stage was small, but
214different from zero (estimate = 0.09, 95% CI = [0.02 – 0.18], P = 0.001). In contrast,
215repeatability of female aggressiveness in the adult stage was not different from zero
216(estimate = 0.1, 95% CI = [0 – 0.08], P = 0.39). Repeatability of distance travelled by
217males in activity trials was different from zero (estimate = 0.32, 95% CI = [0.16 –
2180.44], P < 0.001).
219 Mean female aggressiveness in the subadult stage was not correlated with
220mean female aggressiveness in the adult stage (rho33 = -0.02, P = 0.91), nor with
221absolute (rho33 = 0.02, P = 0.9) or relative growth from the subadult to the adult stage
222(rho33 = 0.19, P = 0.27).
223
224Mating trials: sexual cannibalism and male mating decisions
225 Pairs interacted in 18 out of 35 mating trials (51.4%). Females cannibalized
226males after mating in two trials, but these instances were not tallied as pre-copulatory 227cannibalism as females attacked males several hours after mating. Mating occurred
228only in five out of the 18 trials in which pairs interacted (27.8%), while the remaining
229trials resulted in pre-copulatory sexual cannibalism (72.2%). Males that interacted
230with females took from three to 48 hours to approach them (x̄ = 21.3, SD = 13.9).
231 Our results regarding the influence of several female and male traits on the
232mating decisions of both sexes are summarized in Table 1. Only female age
233influenced the likelihood of pre-copulatory sexual cannibalism: the older the female,
234the lower the likelihood to cannibalize a male. Furthermore, both female
235aggressiveness in the subadult stage and male activity influenced the likelihood of
236male approach. More active males were more likely to approach females, and females
237that were more aggressive in the subadult stage were more likely to be approached by
238a male. Additionally, male age and size influenced the amount of time that males took
239to interact with a female: the younger and the larger the male, the longer it took for
240him to approach a female.
241
242Table 1. Statistical results from three different models exploring the influence of
243different factors in female and male mating decisions.
Sexual cannibalism Male approach likelihood Male approach latency likelihood (N = 35, R2 = 0.324) (N = 18, R2 = 0.457) (N = 18, R2 = 0.431) GLM LRT GLM LRT LM LRT Subadult female -0.182 0.364 0.738 6.044 -5.305 0.104 aggressiveness (0.604) (0.546) (0.396) (0.014) (3.599) (0.747)
0.402 0.009 0.67 4.281 2.296 1.192 Male activity (0.621) (0.924) (0.407) (0.039) (3.351) (0.275)
-0.028 2.329 0.016 0.001 3.822 0.227 Female size (0.456) (0.127) (0.343) (0.97) (2.763) (0.634) -0.191 1.658 0.081 0.122 6.194 6.512 Male size (0.517) (0.198) (0.344) (0.727) (2.858) (0.011)
-1.94 6.139 0.137 0.197 -3.35 1.288 Female age (1.018) (0.013) (0.345) (0.657) (3.479) (0.256)
-1.858 1.243 -0.188 0.011 -13.308 7.211 Male age (1.179) (0.265) (0.348) (0.916) (5.435) (0.007) 244Values given for GLMs and LMs are model coefficients and standard error (in 245brackets). 246Values given for LRTs are chi-square statistics (df = 1) and p-values (in brackets). 247Shaded cells highlight significance effects according to LRTs. 248 249
250Figure 1. Relationship between female and male traits and mating decisions in
251Miomantis caffra. Sexual cannibalism likelihood increases with female age (A). Male
252approach likelihood increases with both female aggressiveness in the subadult stage
253(B) and male activity (C). Latency to reach a female increases with male pronotum
254length (D) and decreases with male age (E). 255
256Discussion
257 In this study, we investigated the existence of personality traits in M. caffra
258and their role in female and male mating tactics. Males showed behavioural
259consistency in activity, while females showed some behavioural consistency in
260aggressiveness in the subadult stage, but not in the adult stage. We found that these
261personality traits influenced males’ decision to approach a female, but not their speed
262in doing so, nor the likelihood of sexual cannibalism. These results suggest that
263personality traits of both sexes can be valuable in understanding the complex
264interactions occurring in M. caffra.
265 Our results do not support the aggressive spillover hypothesis (Arnqvist and
266Henriksson 1997) in M. caffra for multiple reasons. First, this hypothesis proposes
267that aggressiveness is consistent within and between ontogenetic states. We found that
268aggressiveness in M. caffra females was not consistent when they were adults, and
269only somewhat consistent when they were subadult. Second, the aggressive spillover
270hypothesis suggests that, although aggressive females should be penalized in the
271mating context, they can compensate by growing more than other females. We found
272that M. caffra females that were aggressive in the subadult stage did not grow larger
273than less aggressive females, but we note that our experimental design may have
274imposed constraints on this prediction as less aggressive females were not penalized
275for being slow hunters because they received the same amount of prey as other
276females (see Johnson and Sih (2005) for an alternative approach). Third, and most
277importantly, the aggressive spillover hypothesis predicts that aggressiveness in the 278subadult stage increases the likelihood of sexual cannibalism in adulthood, but we did
279not find such a pattern in M. caffra females. Overall, these results support those of a
280previous study of M. caffra (Fisher et al. 2020), despite the differences in the methods
281used (e.g. aggressiveness as attacks on computer generated prey instead of live prey).
282Our results also agree with other studies that found no evidence in favour of the
283aggressive spillover hypothesis (reviewed in Kralj-Fišer et al. 2013). However, M.
284caffra can reproduce parthenogenetically and may therefore avoid costs of mating
285failure due to pre-copulatory sexual cannibalism. Given this reproductive assurance,
286females in this species may have evolved to be highly aggressive on average to
287increase their foraging success without reproductive costs. If this is the case, even the
288least aggressive females would be propense to attack males, explaining the unusual
289high rate of sexual cannibalism seen in this species. Evidence from (Fisher et al.
2902020) may support this argument, as that study found that M. caffra females are, on
291average, more aggressive than a sympatric and ecologically similar mantis species
292(Orthodera novaezealandiae) that is incapable of parthenogenesis.
293 We found that female age influenced the occurrence of pre-copulatory sexual
294cannibalism in M. caffra: older females were less prone to cannibalize males. In that
295regard, our results are similar to those of (Walker and Holwell 2016), even though the
296variation in female age in our trials was much smaller than in their experiments. Such
297an age effect can occur because of the selective pressure on cannibalistic females to
298avoid mating failure (e.g. Uetz and Norton 2007; Wilgers and Hebets 2012; Gavín-
299Centol et al. 2017). However, M. caffra females can potentially escape this constraint
300because of their ability to reproduce parthenogenetically (Walker and Holwell 2016).
301It is possible that females become less mobile as they age due to increasing gravidity 302(e.g. Pollo et al. 2020), which may lead to reduced frequency or success of
303cannibalistic attacks. Alternatively, older females in our experiments may have
304accumulated greater fat reserves making them less compelled to see males as food,
305but this is unlikely given that diet does not influence cannibalism in this system
306(Walker and Holwell 2016). Perhaps there are benefits to females in reproducing
307parthenogenetically early in adult life and switching to sexual reproduction later, but
308this remains to be tested.
309 Surprisingly, males did not approach older females more often or faster
310despite the lower chance of being eaten by these females. Such male mate choice may
311be absent because cues associated with female age (e.g. pheromones; Klein et al.
3122012) may be undetectable by males, or because older females are less desirable for
313other reasons (e.g. decrease in fecundity or egg viability; Lingren et al. 1988; Moore
314and Moore 2001). In contrast, females that were more aggressive as subadults were
315more often approached by males in our experiments, which may be considered a form
316of male mate choice. Mate choice for certain behavioural types can be expected given
317that they are often linked to other traits that confer direct benefits to the chooser
318(reviewed in Munson et al. 2020). As the aggressive spillover hypothesis suggests,
319juvenile aggressiveness could be related to adult size, which is often related to
320fecundity in invertebrates (Honěk 1993; Head 1995). But in our study, females that
321were more aggressive in the subadult stage did not develop into larger adults,
322although such an association is yet to be assessed in wild fed individuals.
323Alternatively, greater female aggressiveness in the subadult stage may represent an
324indirect benefit to males if it is a heritable trait that increases fitness by, for example,
325enhancing offspring survivorship. 326 We found that males that were more active in a non-mating context (i.e. in
327activity trials) were more likely to approach females. More active males are usually
328bolder as well (i.e. more prone to risk; sensu Réale et al. 2007; Biro and Stamps
3292008). However, we did not find any effect of male activity on cannibalism incidence,
330suggesting that active males either do not face greater risk of attack or adjust their
331behaviour to ameliorate such a risk. Given that praying mantises rely on crypsis, and
332movement reduces camouflage effectiveness (Cuthill et al. 2019), overly active males
333may increase their chance of being captured by other predators (Magnhagen 1991;
334Magnhagen et al. 2014), but may benefit by finding females sooner. Increased activity
335could also reflect a “live-fast-die-young” strategy whereby riskier behaviour is
336favoured because it provides greater rewards (Réale et al. 2010). Assessment of
337correlations between behavioural, physiological, and life-history traits could provide
338useful insights on pace-of-life in this system. It is also possible that personality
339underpins cannibalism avoidance strategies more generally. Active and less active
340males may assess risk differently, and therefore use different behaviours in their
341approach, or choose to approach in different contexts. Assessing whether personality
342mediates well-known male tactics such as waiting for females to eat (Fromhage and
343Schneider 2005), playing dead during attacks (Bilde et al. 2006), and offering nuptial
344gifts (Costa-Schmidt et al. 2008; Ghislandi et al. 2014) would be a fruitful next step.
345 Despite the lack of influence of any male trait on sexual cannibalism
346likelihood in our experiments, we found that several such traits influenced male
347mating decisions. Besides more active males being more likely to approach females,
348younger and larger males took longer to approach females. It is possible that we did
349not detect an effect of male activity on sexual cannibalism likelihood because less 350active males infrequently interacted with females, resulting in fewer female
351interactions with less active males in our mating experiments. Using a similar
352rationale, perhaps younger and larger males would have been more often cannibalized
353if not by their slower approach. In other words, males may express distinct decisions
354depending on their own traits as a counter strategy to sexual cannibalism. Indeed,
355sexual conflict can lead to arms races where traits in one sex counter select for traits
356in the other sex (e.g. Friberg et al. 2005; Tatarnic and Cassis 2010). This occurs in M.
357caffra, as males wrestle females to circumvent sexual cannibalism (Burke and
358Holwell 2021a). However, further investigation is required to evaluate whether male
359traits can influence such coercive behaviours and the overall sexual conflict dynamics
360in this species, as our experimental design was not intended to test this.
361 Most studies of animal personality aim to describe behavioural patterns or
362understand the causes of behavioural consistency rather than the ecological or
363evolutionary consequences of such behaviours (Wolf and Weissing 2012; Beekman
364and Jordan 2017). In this study, we not only assessed the consistency of different
365behaviours in males and females but evaluated their effect on mating interactions and
366the likelihood of cannibalism. Our results suggest that female aggressiveness and
367male activity (among other traits) play an important role in mating dynamics in M.
368caffra but have little effect on cannibalism incidence. We highlight the importance of
369investigating both female and male behaviours given that decisions of one sex likely
370influence the decisions of the other under sexual conflict.
371
372Acknowledgements 373 We thank Rose O’Dea for assistance and advice with repeatability analyses.
374We thank Michael Kasumovic for feedback on the manuscript.
375
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