1Effects of male and female personality on sexual 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 , 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: temperament, male mate choice, mating decisions, .

34

35Introduction

36 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 , 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

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 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. ; 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 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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