bs_bs_banner

Entomological Science (2012)

doi:10.1111/j.1479-8298.2012.00535.x

ORIGINAL ARTICLE

Behavioral response of male mantid Tenodera aridifolia
(Mantodea: Mantidae) to windy conditions as a female
approach strategy
ens_535

1..8

Hiroshi WATANABE and Eizi YANO
Entomological Laboratory, Faculty of Agriculture, Kinki University, Nara, Japan

Abstract
When sexual cannibalism presents a sexual conflict, one expects to find male traits that reduce the risk of
cannibalism. In sexually cannibalistic species, selection is thought to have favored the evolution of male
approach behaviors that reduce the probability that the female will kill the male. We tested the hypothesis
that male mantids change their approach behavior in response to wind to reduce the risk of being noticed
by females. Time between detection of the female by the male and mating was shorter under windy than
windless conditions. Sexual approach behavior was observed more frequently under windy than windless
conditions. Moreover, this behavior was observed more frequently when the female was walking than when
the female was not walking under windy conditions. The detection rate of male mantids by females was
significantly lower on swaying leaves than on fixed leaves. Our results thus indicate that male mantids were
more active in response to wind. Therefore, we suggest that the male’s quick approach strategy toward
females when the wind is blowing at short range is adaptive in reducing the risk of detection by females.
Key words: cryptic, mating behavior, risk avoidance, sexual approach behavior, sexual cannibalism, visual
cues, windy conditions.

INTRODUCTION
Sexual cannibalism in praying mantids is well recognized, with some species displaying sexual cannibalism
at least occasionally (reviewed by Elgar 1992; Maxwell
1999a). Studies over the past two decades have examined a variety of questions surrounding mating behavior
in mantids, with particular attention toward the effect
of sexual cannibalism on female reproductive output
(Parker 1979; Buskirk et al. 1984; Birkhead et al. 1988;
Maxwell 2000; Barry et al. 2008) as well as on male
mating behavior (Liske & Davis 1987; Maxwell 1998,
1999b; Gemeno & Claramunt 2006; Lelito & Brown
2006; Barry et al. 2009, 2010).
Female body condition is positively related to
fecundity in many cannibalistic spiders and mantids
Correspondence: Hiroshi Watanabe, Entomological
Laboratory, Faculty of Agriculture, Kinki University, Nara
631-8505, Japan.
Email: kqhss155@ybb.ne.jp
Received 22 November 2011; accepted 8 April 2012.

© 2012 The Entomological Society of Japan

(Rubenstein 1987; Birkhead et al. 1988; Wise 2006;
Barry et al. 2008), and the high variability of female
body characteristics in natural populations (Eisenberg
et al. 1981; Lawrence 1992; Maxwell 1998) suggests
that males should choose females in relatively good condition to maximize their reproductive fitness (Lawrence
1992; Moya-Laraño et al. 2004; Barry et al. 2008,
2010; Maxwell et al. 2010).
Pheromone emission by females might result in multiple mating for females (Edmunds 1975; Robinson &
Robinson 1979; Lawrence 1992). However, both sexes
are capable of mating several times (Fabre 1897; Bartley
1982; Liske & Davis 1987; Lawrence 1992; Hurd et al.
1994; Holwell et al. 2007). Thus, males may be better
served by surviving copulation and finding further
mating opportunities. Recently, researchers have documented cannibalism of males by females in four mantid
species in nature, reporting that cannibalism comprises
17–31% of these intersexual encounters (Lawrence
1992; Hurd et al. 1994; Maxwell 1998). This suggests
that, at least for some mantid species, selection should
favor the evolution of male anti-cannibalistic behaviors.

The male was upwind. the male generally moves his body away from the female without moving his legs. Birkhead et al. 1988. Nymphs hatched in September 2011 were used to examine the effect of wind on time elapsed for the male’s searching for prey and on the male’s searching for the prey. Watanabe and Yano (2009) reported that Hierodula patellifera (Serville) shows walking and body swaying more under windy than windless conditions. Entomological Science (2012) © 2012 The Entomological Society of Japan . Sphero Aqua Company. We focused on the behavioral response of male Tenodera aridifolia (Stoll) to wind as an approach strategy toward females. Kynaston et al. Yano Males should choose females that are less likely to cannibalize them. Figure 1 Experimental set-up to evaluate the effect of wind on male approach behavior toward females. Japan in October 2010. It was kept in a plastic cage (12. Nara.) Ohwi facing a fan (Fig.H. Sixth instar nymphs were moved individually to acrylic fiber cages (10 cm diameter. Kinki University. 20 cm height) whose top 2 and side openings were covered with a screen to enable the nymph to walk and molt.) Ohwi facing a fan. 2002. Nymphs were fed on fruit flies (Drosophila sp. Thus. All experiments were conducted between 13:00 and 17:00 hours in July 2009 at 25–26°C. and may use female body size or feeding states (Liske & Davis 1987. possibly imitating the movement of the vegetation in the wind. Experimental setup No mantids had mating experience at the beginning of any experiment.) (Sphero Aqua Company). aridifolia adults readily copulate within 8–10 days of the final molt (Liske & Davis 1987). or turns and disappears into the vegetation (Lawrence 1991). One male and one female satiated adult mantid were selected and placed 15 cm apart on a leaf of the kudzu vine Pueraria lobata (Willd. aridifolia was collected from the field at the Faculty of Agriculture. Japan in October 2008. 10 cm height) until the sixth instar. Herberstein et al.. An ootheca of the mantid T. If the female moves toward a male. The cage was moved to a constant-temperature room (30°C) in May 2009 to promote mantid hatching from the ootheca. male mantids behave in a manner clearly indicative of risk avoidance.5°C.5 cm top diameter.). Japan) until the third instar.5 cm top diameter ¥ 10 cm bottom diameter ¥ 10 cm height) in a temperature-controlled cabinet at 10 ! 0. Larvae that hatched in June 2009 were used for all experiments other than the experiment to examine the effect of wind on time elapsed for the male’s searching for prey and on the male’s searching behavior for prey. Eighty-two nymphs were reared in plastic cages (12. Males move toward females most often and most rapidly when females move away or when a breeze moves the surrounding vegetation (Lawrence 1992). 1). Prey items were supplied to each mantid every other day. other than the experiment to examine the effect of wind on time elapsed for the male’s searching for prey and on the male’s searching behavior for prey. or female mating status (Newman & Elgar 1991. An ootheca was moved to a constant temperature room (30°C) in July 2011 to promote mantid hatching from the ootheca. Kinki University. Fourth and fifth instar nymphs were fed on cockroach Blattella germanica (L. 10 cm bottom diameter. Shizuoka. as an indication of a female’s propensity to cannibalize. Males often rock as they approach females. 1994. Sixth and seventh instar nymphs and adults were fed house crickets Acheta domesticus (L. Schneider & Elgar 2002). We hypothesized that male mantids change their approach behavior with wind velocity and use wind to reduce the risk of being noticed by females. Drinking water was sprayed on the cages several times every other day. each adult was 14–23 days old after eclosion because T. Watanabe and E. Moreover. Eighty nymphs were reared by the same rearing method as the other experiment. Andrade 1998). One male and one female satiated adult mantid were selected and placed 15 cm apart on a leaf of the kudzu vine Pueraria lobata (Willd. ensuring adequate nutrition. 50% relative humidity (RH) under conditions of 12 h light : 12 h dark (LD 12:12). MATERIALS AND METHODS Tenodera aridifolia An ootheca of the mantid T. We conducted experiments to study male mantid responses to wind during their approach and the efficacy of an approach strategy toward females under windy conditions. Nara. aridifolia was collected from the field at the Faculty of Agriculture.

We analyzed male behavioral components (i. A Fisher’s exact probability test was performed to test the relationship between the approach behavior of males and the walking behavior of females under windy conditions (n = 24). The time was measured using a stopwatch.0001. The mantid executes two types of body swaying: swaying from side to side and swaying backward and forward. A Wilcoxon rank-sum test was used to compare the time elapsed for approaching for females and that for searching for prey (n = 24). and a rhythmic bending motion of the abdomen at an angle of 0–90° (Liske & Davis 1987). Courtship by male T. 1.001. 1. n = 24) (Fig. Effect of wind on time elapsed for the male’s searching for prey We examined the effect of wind on time elapsed for the male’s searching for prey to compare that for the male’s approaching females. courtship display and resting) and female behavioral components (i. the leaf was fixed with a wire so that it could not sway in the wind. body swaying. 3). we defined the head-turning behavior as a sign of a male’s detection of a moving female.e. or a constant wind velocity of 1. 2). U = 0. Wind velocity was measured when the fan was on by using an anemometer (Model 6533-01. walking.0 for Windows (SPSS Inc.34 m/s. USA). We did not discriminate theses two types of swaying behavior. aridifolia involves an upward thrusting of the forewing and hindwings. Chicago.34 m/s) every 10 s. Effect of wind on mating behaviors of males and females We video-recorded the female behavior as well as the approach behavior of the male after the female had been detected for 100 s when alternating between windless and windy conditions (average wind velocity. U = 45. Moreover.00. n = 24) (Fig.34 m/s) every 10 s to compare the male behavior when approaching females and searching for prey. Responses of females to approaching males We recorded whether the female detected the approaching male under windy conditions. RESULTS Effect of wind on time from detection of the female to mating The time elapsed between the male’s detection of the female and mating behavior was significantly shorter under windy conditions than under windless conditions (Wilcoxon rank-sum test.Sexual approach strategy of male mantid Moreover. A one-way analysis of variance (anova) was used to compare the percentage of the total duration of each Entomological Science (2012) © 2012 The Entomological Society of Japan behavioral component under windless and windy conditions after arcsine transformation. IL. The experiment was started when the male mantid detected the female except for the experiment carried out without females. Effect of wind on time elapsed for the male’s searching for prey The time spent for approaching females was significantly longer than the searching for prey under constant windy condition (Wilcoxon rank-sum test. All statistical analyses were performed using SPSS v14. body swaying.. A Fisher’s exact probability test was performed to test the difference in detection or attacking rates between treatments (n = 24). Effect of wind on time between the male’s detection of the female and mating The time between the male’s detection of the female and mating was examined under two conditions: either windless. Osaka. P < 0. We recorded the time elapsed to arrive at the end portion of a leaf 15 cm from the starting position. P < 0. The head-turning behavior was used as a sign of a male’s detection of a moving female. Mantids react to moving objects first by turning their heads.00. Kanomax. Effect of wind on male’s searching behavior for prey We video-recorded the male behavior under the condition without a female for 100 s when alternating between windless and windy conditions (average wind velocity. we conducted the experiment between 13:00 and 17:00 hours in December 2011 at 25–26°C to examine the effect of wind on time elapsed for the male’s searching for prey and on the male’s searching behavior for prey.34 m/s). A one-way anova was used to compare the percentage of the total duration of each behavioral component under windless and windy conditions after arcsine transformation. Japan). Effect of wind on male and female mating behaviors Males spent significantly longer time walking under windy conditions than under windless conditions 3 . grooming. Therefore.e. we recorded the occurrence of the aggressive strike by the female after the female detected the approaching males under windy conditions. The male mantid was released from the starting point on the end portion of a leaf and it began searching under constant windy conditions (1. resting and walking). For the control condition.

= 1.001.001. d.342. Watanabe and E. 5A). d. = 1. or (ii) a constant wind velocity of 1. d.660. d. n = 24.702. P < 0. Values are means + SE.720.0001. = 1. 4 Figure 4 Percentage of the total duration of (A) walking behavior. n = 24) (Fig. 4C). We recorded the time elapsed to arrive at the end portion of a leaf 15 cm from the starting position.780. We recorded the approach behavior of males after the male had detected the female mantid. Males spent significantly longer time resting when Entomological Science (2012) © 2012 The Entomological Society of Japan .001. (B) resting behavior and (C) courtship display of males during observations for 100 s when the wind condition was alternated between windless and windy at a constant wind velocity of 1. Yano Figure 2 Time between the detection of the female by the male and mating was examined under two conditions: (i) windless.f.f. 5B). Effect of wind on the male’s searching behavior for prey Males spent significantly less time walking when approaching females than when searching prey (anova: F = 173.f.001. Males spent significantly less time resting under windy conditions than under windless conditions (anova: F = 46.34 m/s. Males spent significantly longer time in courtship display under windy conditions than under windless conditions (anova: F = 18. Females spent significantly less time resting under windy conditions than under windless conditions (anova: F = 39. n = 24) (Fig. = 1. 4B). (anova: F = 89. P < 0. n = 24) (Fig. 4A). 6).34 m/s). n = 24. P < 0. The male mantid was released from the starting point on the end portion of a leaf and it began searching under constant windy conditions (1. n = 24) (Table 1). P < 0.34 m/s every 10 s. n = 24) (Fig. = 1. n = 24) (Fig.f.001. Males approached walking females more frequently than non-walking females under windy conditions (Fisher’s exact probability test: P < 0. P < 0. P < 0.f.001. Females spent significantly more time walking under windy conditions than under windless conditions (anova: F = 79.f. n = 24) (Fig.H. d. Figure 3 Time spent approaching females or searching for prey under windy conditions. = 1. d. n = 24.783.

DISCUSSION The time between a male’s detection of a female and mating was significantly shorter under windy conditions Entomological Science (2012) © 2012 The Entomological Society of Japan No. Experimental conditions are as for Figure 4. Table 2 Detection rate of approaching male mantids by conspecific females under swaying and fixed-leaf conditions Figure 5 Percentage of the total duration of (A) walking behavior and (B) resting behavior of females under alternating windy and windless conditions.001. These behavioral responses to wind are also a known antipredator strategy of mantids (Watanabe & Yano 2009).001. n = 24) (Table 3). of observations) Approaching Not approaching 40 1 9 70 approaching females than when searching for prey (anova: F = 24. aridifolia are honest indicators of female mate availability and a lower risk of sexual cannibalism (Lelito & Brown 2008).34 m/s) when approaching females and searching for prey. Thus. 6). Responses of females to approaching males Females detected males significantly less frequently under the swaying leaf condition than the fixed leaf condition (Fisher’s exact probability test: P < 0. n = 24) (Fig. = 1. n = 24. Female-emitted pheromones are definitely impor- 5 . Table 1 Relationship between approach behavior of males and walking behavior of females under windy conditions Behavior of females Walking Not walking Behavior of males (No.001. Mate attraction signals of female T. Females attacked males significantly less frequently under the swaying leaf condition than the fixed leaf condition (Fisher’s exact probability test: P < 0. of observations Leaf condition Detected Not detected Detection rate (%) Fixed Swaying 9 2 15 22 38 8 Table 3 Attacking rate of approaching male mantids by conspecific females under swaying and fixed leaf conditions No. n = 24) (Table 2). When the wind stopped.f. mantids stopped moving to rest (Fig. d. this was unlikely given that we placed the male upwind from the female.Sexual approach strategy of male mantid Figure 6 Percentage of the total duration of walking behavior and resting of males during observations for 50 s when wind is blowing (1. 2).737. Males walked for a significantly longer time under windy conditions than under windless conditions (Fig. P < 0. 3B). 3A). However. An alternative hypothesis was that the males quickly approach females when the wind is blowing because they are better able to detect and locate the females by pheromones. a male mantid’s behavioral change in response to wind could be thought of as a strategy for approaching females. of observations Leaf condition Attacked Not attacked Attacking rate (%) Fixed Swaying 7 1 17 23 29 4 than under windless conditions (Fig.

Lelito & Brown 2006. Therefore. Maxwell et al. Males showed courtship displays frequently even when females did not look at the males under windy conditions. Moreover. Maxwell et al. and almost always on the horizontal plane (Liske & Davis 1987. approaching a female responding to wind is more common than the above conditions in nature. their pre-mating movements are induced by both their movements and the wind. Barry 2004. without any further stalking (Roeder 1935) or abdominal flexing (Liske & Davis 1987). females may approach males by walking. we demonstrated that the males’ cryptic or quick approach to females when the wind blows reduces the risk of detection by females. in part. Because both females and males do not move frequently under windless conditions. mantids are normally found on plants (Barrows 1984. This switch in orientation could be interpreted as either risk avoidance or simply facilitation of proper alignment for mounting. Entomological Science (2012) © 2012 The Entomological Society of Japan . Most inferences come from purely descriptive accounts of male mating behavior (Roeder 1935. We documented male approach behaviors toward females under more natural conditions for the first time by simulating wind blowing against mantids on a leaf in the laboratory. religiosa mantids approach females much faster (¥6. religiosa starts to walk.6). When this happens. Watanabe and E. Females were more active under windy conditions than under windless conditions (Fig. this male approach behavior is quite different from the searching behavior because this male behavior is to approach to the female as one of the processes of mating behavior after the male recognized the female (Fig. no strong experimental evidence exists to demonstrate either the complicity of males or active male risk avoidance. 3C). Our work is the first laboratory experiment to confirm the results of observation in the field by Lawrence (1992). 6). Kynaston et al. physiological condition and predatory posturing (Lelito & Brown 2008. 2009. Maxwell 1999a). However. males approach the female more rapidly (Lawrence 1992). The male approaches the female more slowly and cautiously than when it searches for the prey (Figs 3. If a female M. which allows for more complex male approach strategies than laboratory arenas. a courtship display is induced by female behavior as well as by wind. 4A). However. in the field. Vision likely guides a male’s detection and location of a female. Similar results reported by Watanabe and Yano (2009) suggested an antipredator strategy of mantids. Hurd 1999). For example. courtship displays by males were more frequently observed under windy conditions than under windless conditions (Fig. Thus.4) when females are catching and eating prey than when they are just sitting and waiting for prey. 1994. for leaves to shake under windless conditions in the field would be rare. 1994. We suggest that the males’ cryptic or quick approach to females when the wind blows is adaptive and reduces the risk of detection by females. 2010). 1999b. Pheromones might be distributed by the wind. 2010). to reduce the risk of cannibalism. Moreover. The frequency of approach behavior by males significantly increased when females walked than when females did not walk under windy conditions (Table 1). Moreover. whereas visual cues and signals become important at shorter ranges (Maxwell 1999a. Note that most laboratory observations of mantid approach behavior (including the present study) have been performed on flat surfaces. Lawrence 1992. Based on our results and studies of other sexual cannibalistic species. 6 The detection rate of male mantids by conspecific females was significantly lower under the swaying leaf condition than under the fixed leaf condition (Table 1). the male’s quick approach strategy toward females at short range when the wind is blowing may be adaptive in reducing the risk of detection by females. as well as helps with species identification and provides information regarding the female’s phenotype. Maxwell 1998. Barry et al. Although a female’s movement in response to wind stimulates male movement.H. aridifolia has evolved. The female’s behavior may contribute to quick approaches by males. We think that the male approach behavior has cryptic effects because the wind undoubtedly stimulates the approach behavior by males. Perhaps this is an indication of male arousal following detection of a female. Yano tant for long-distance attraction of mantids. Mating with feeding females may not be uncommon under natural conditions (Gemeno & Claramunt 2006). the males may mount them quickly. Distinguishing the effects of wind from those of a shaking substratum is necessary. Future studies of male approach behavior toward females in the field are necessary. Kynaston et al. we hypothesize that the adjustment of male speed to the female behavior of T. However. Despite the notoriety of sexual cannibalism in mantids. Liske & Davis 1987. who observed that males moved towards the female most often and most rapidly when females moved away or when a breeze moved the surrounding vegetation. 2010). However. male M. the male’s approach behavior is also stimulated by the wind. Gemeno & Claramunt 2006. but the adaptive significance of the courtship display when females did not look at males under windy conditions is unclear. Females that are eating may be safer to approach than non-eating females because well-fed females are less likely to attack a male than starved ones (Gemeno & Claramunt 2006).

Entomologist’s Monthly Magazine 111. 5. 193–198. 287–355. Animal Behaviour 35. 377– 383. 1108–1111. and possible sex pheromones in three species of Mantodea. 710–715. Herberstein ME. Behavioral Ecological Sociobiology 63. Bartley JA (1981) Ecological consequences of food limitation for adult mantids (Tenodera aridifolia sinensis Saussure). Paris. Gemeno C.). Newman JA. 33–42. We also thank Dr Ikuo Kandori of Kinki University for technical advice. 739–741. Hurd LE. Stagmomantis limbata and Tenodera aridifolia sinensis (Mantodea: Mantidae). Lawrence SE (1992) Sexual cannibalism in the praying mantid. 612– 625. (1994) Cannibalism reverses male-biased sex ratio in adult mantids: female strategy against food limitation? Oikos 69. 7 . Fagan WF et al. Mantis religiosa: a field study. Schneider JM. mating. Crespi BJ (eds) Cannibalism: Ecology and Evolution among Diverse Taxa. Hurd LE (eds) The Praying Mantid. Lee KE. The Johns Hopkins University Press. Fabre JH (1897) Souvenirs entomologiques Nr. Annals of the Entomological Society of America 103. In: Elgar MA. 205–219. 307–313. MD. Baltimore. Wells H. 1372–1395. PhD Thesis. Davis WJ (1987) Courtship and mating behaviour of the Chinese praying mantis. Environmental Entomology 11. 313–320. Hurd LE (eds) The Praying Mantid. pp 69–89. Holwell GL. Behavioral Ecology 9. Universityof Sheffield. Kynaston SE. Eisenberg RM. Hurd LE. Maxwell MR (1999a) The risk of cannibalism and male mating behaviour in the Mediterranean praying mantid Iris oratoria. Pascual J. 1524–1537. Oxford University Press. Animal Behaviour 79.Sexual approach strategy of male mantid ACKNOWLEDGMENTS We thank Dr Taku Iwasaki for reviewing earlier drafts of the manuscript. In: Prete FR. Animal Behaviour 43. The Johns Hopkins University Press. Barry KL. Macquarie University (Department of Biological Sciences). Holwell GI. Moya-Laraño J. Mill PJ (1994) Courtship. Sphodromantis lineola. Young P (1988) Sexual cannibalism in the praying mantis Hierodula membranacea. Elgar MA (1992) Sexual cannibalism in spiders and other invertebrates. Journal of Insect Behavior 19. Wise DH (2004) Approach strategy by which male Mediterranean tarantulas adjust to the cannibalistic behaviour of females. Barry KL. Herberstein ME (2007) Mate location. McErlain-Ward P. Edmunds M (1975) Courtship. Ethology 110. Barrows EM (1984) Perch sites and food of adult Chinese mantis (Dictyoptera: Mantidae). Herberstein ME (2008) Female praying mantids use sexual cannibalism as a foraging strategy to increase fecundity. 209–218. Maxwell MR (1999b) Mating behavior. Behavioral Ecology 19. Proceedings of the Entomological Society of Washington 86. Wells PH. Honours thesis. Journal of Ethology 27. 120–127. England. American Midland Naturalist 106. Animal Behaviour 55. American Naturalist 168. Herberstein ME (2010) Multimodal mate assessment by male praying mantids in a sexually cannibalistic mating system. Herberstein ME (2009) Male mating behaviour reduces the risk of sexual cannibalism in an Australian praying mantid. Maxwell MR. Behaviour 106. Wells PH. Buskirk RE. Baltimore. Holwell GI. Maxwell MR (1998) Lifetime mating opportunities and male mating behaviour in sexually cannibalistic praying mantids. In: Prete FR. Wells H. 53–57. antennal morphology and ecology in two praying mantids (Insecta: Mantodea). Maxwell MR (2000) Does a single meal affect female reproductive output in the sexually cannibalistic praying mantid Iris oratoria? Ecological Entomology 25. Barry KL. Animal Behaviour 47. pp 128–155. Claramunt J (2006) Sexual approach in the praying mantid Mantis religiosa (L. Elgar MA (2002) Costs of courtship and mating in a sexually cannibalistic orb-web Entomological Science (2012) © 2012 The Entomological Society of Japan spider: female mating strategies and their consequences for males. 717– 724. Barry KL (2004) Mating behavior of a sexually cannibalistic praying mantid Pseudomantis albofimbriata. American Naturalist 123. Lelito JP. Holwell GI. Behavioral Ecology and Sociobiology 51. 569– 583. Lelito JP. MD. Eisenberg RM. Elgar MA (1991) Sexual cannibalism in orbweaving spiders: an economic model. Barry KL. Ross KG (1984) The natural selection of sexual cannibalism. (Mantodea). Hurd LE (1999) Ecology of preying mantids. 263– 269. mating behaviour and sexual cannibalism in the praying mantis. Brown WD (2006) Complicity or conflict over sexual cannibalism? Male risk taking in the praying mantis Tenodera aridifolia sinensis. Brown WD (2008) Mate attraction by females in a sexually cannibalistic praying mantis. 112–118. Biological Journal of the Linnean Society 91. Tenodera aridifolia sinensis. Frohlich C. Liske E. REFERENCES Andrade MCB (1998) Female hunger can explain variation in cannibalistic behavior despite male sacrifice in redback spiders. 440– 446. 1165– 1172. 1011–1028. American Naturalist 138. 898–901. pp 43–60. Librairie Ch Delagrave 5. 8 fig. Behaviour 136. Barry KL. 731–740. Sydney. Johns PM (2010) Examinations of female pheromone use in two praying mantids. Bartley JA (1982) Movement patterns in adult male and female mantids (Tenodera aridifolia sinensis Saussure). Lawrence SE (1991) Sexual cannibalism in praying mantids. 54–62. Birkhead TR. Oxford.

Academic Press. Science 205.). Annual Review of Entomology 51. intraspecific competition and the regulation of spider populations. Roeder KD (1935) An experimental analysis of the sexual behavior of the praying mantis (Mantis religiosa. 517–522. Elgar MA (2002) Sexual cannibalism in Nephila plumipes as a consequence of female life history strategies. 203–220. Watanabe and E. 825–827. In: Blum MS. Journal of Evolutionary Biology 15. food limitation. Entomological Science (2012) © 2012 The Entomological Society of Japan . Robinson MH. Annals of the Entomological Society of America 102. Watanabe H. Behavioral Ecology and Sociobiology 20. Robinson B (1979) By dawn’s early light: matutinal mating and sex attractants in a Neotropical mantid.H. Blum NA (eds) Sexual Selection and Reproductive Competition in Insects. Biological Bulletin 69. 8 Schneider JM. pp 123–166. 229–237. New York. Yano Parker GA (1979) Sexual selection and sexual conflict. Yano E (2009) Behavioral response of mantid Hierodula patellifera to wind as an antipredator strategy. Rubenstein DI (1987) Alternative reproductive tactics in the spider Meta segmentata. Wise DH (2006) Cannibalism. 441–465. 84–91. L.