Vision in the mantispid: a sit‐and‐wait and stalking predatory insect

Mantispids (Neuroptera: Mantispidae) are remarkable insects as a result of their close resemblance to the praying mantis (order Mantodea). Although not closely related phylogenetically, as a result of similar selective pressures, both mantispids and mantids have evolved powerful raptorial forelegs for capturing insects. Another striking feature is the hypermetamorphosis in mantispid development, as well as the parasitizing behaviour of the first‐instar larvae. The present review focuses on the role of mantispid vision. First, the morphology and functional significance of the larval eyes (stemmata) are examined. In principle, the stemmata are suitable for spatial vision because of their arrangement and structure. This is then followed by a discussion of how adult mantispids are able to capture fast‐moving insects successfully, although, in contrast to the praying mantis, mantispids rely on superposition eyes rather than on apposition eyes with a frontal region of high acuity. For both larvae and adults, comparisons are made with other insect groups. The present review also addresses the role of mantispid vision as an important cue for triggering mating behaviour; accordingly, sex‐specific differences are considered. Finally, vision in the context of orientation flight is discussed.     

Statilia maculata (Thunberg, 1784) – the first invasive praying mantis (Mantodea,Mantidae) in the fauna of Russia

Summary

Recent years have seen an increasing number of alien praying mantis species being introduced and establishing populations across Europe. Here we add to this count, reporting a stable and spreading population of an Asian species, Statilia maculata (Thunberg, 1784), in Krasnodar Krai of Southern Russia. This is the first introduced praying mantis species in the Russian fauna and the first representative of the genus Statilia Stål, 1877 in Europe. We review what is known about biology of this species and provide diagnostic characters to distinguish nymphs, adults and oothecae from native and introduced European species of praying mantis. Our attempts at captive breeding identify humidity as one of the most important factors limiting possible spread of S. maculata. Invasion to neighbouring territories and other countries with a humid subtropical climate remains a very real possibility with an unknown outcome for local ecosystems.     

Social and maintenance behaviour in two Australian ocypodid crabs (Crustacea: Brachyura)

The feeding, burrowing and fighting behaviour and courtship display of the two ocypodids Heloecius cordiformis (H. Milne Edwards) and Hemiplax latifrons (Haswell) are described from daytime observations in the field over a two-year period and from four months study under laboratory conditions. The behaviour of the two species is compared with that of other ocypodid crabs. Heloecius cordiformis and Hemiplax latifrons live intertidally in estuaries in semi-permanent burrows and feed on detritus sifted from the mud and sand by the mouth-parts. The two species each possess considerably stereotyped patterns of behaviour, particularly in relation to fighting and courtship display in which the two differ most. The courtship display of Heloecius cordiformis closely resembles that of vertically waving, narrow-fronted species of fiddler crabs (genus Uca). The fighting behaviour of Hemiplax latifrons is similar to that of the New Zealand Hemiplax hirtipes (Jacquinot) which apparently lacks courtship display.     

Prey capture in mantids: A non-stereotyped component of lunge

The praying mantis Tenodora aridifolia sinensis strikes at prey with the pincer-like motion of its prothoracic legs. During strike the mantis moves its body forward toward the prey in a lunge which is propelled by its four walking legs. Using a tethered mantis preparation we have studied the lunge produced by the movement of the walking legs. We have found that lunge is correctly oriented toward prey no matter where it moves in three-dimensional space. This demonstrates that the lunge that accompanies the strike is in this species aimed and not invariant in distance and direction as suggested for other mantids.     

It Takes Two to Tango: Relative Influence of Male and Female Identity and Morphology on Complex Courtship Display in a Newt Species

Consistency in behaviour is currently receiving a renewed interest. Although courtship display is generally consistent in terms of behavioural sequence and structure, there is also commonly important variation in the intensity of courtship display between and within males of a given species. Indeed, not all males have the same ability to perform courtship display (variation between males), and each male can potentially adjust his courtship effort in response to the environment (variation within a male). Although the study of male courtship display has received considerable attention in recent years, it is still unclear which part of the variation can be explained by male ability or motivation. We investigated this issue on two phases of the complex courtship display of the palmate newt Lissotriton helveticus. Overall, we found that both male and female identities affected courtship behaviour, but the relative influence of each sex depended on the courtship phase. Male identity explained variation in fan and creep‐quiver display, whereas female identity explained variation in creep‐quiver only. Interestingly, we did not find any link between the expression of courtship display and male or female morphological traits. Our study showed consistency of male courtship display in newts and successfully dissects the different sources of variation that can affect behavioural repeatability/consistency of courtship display.     

Behavioural evidence for a female sex pheromone in Cotesia flavipes (Hymenoptera: Braconidae)

Abstract.  The mating behaviour of Cotesia flavipes Cameron, 1891 (Hymenoptera: Braconidae) was observed under laboratory conditions. The results demonstrate that chemical stimuli play an important role in bringing males and females together for mating. Females from this species release chemical compounds to attract males, which exhibit a distinct courtship behaviour divided into three main steps: (i) antennation, (ii) lateral bouncing and (iii) wing fluttering. Ethological tests using hexane extracts from two different parts of the female's body (head plus thorax and abdomen) demonstrate that males are more attracted and also display a courtship behaviour when a filter paper containing hexane extracts from the abdomen is presented to them, indicating that the abdomen is the source of the sex pheromone of this species. Naive males of this species respond to hexane extracts of this gland by displaying courtship behaviour. Furthermore, additional tests show that C. flavipes females mate only once, whereas virgin males do not show any preference for virgin or mated females, suggesting that mated females of this species continue to produce sex attractants after mating.     

Behavioural interactions between the lizard <Emphasis Type="Italic">Takydromus tachydromoides</Emphasis> and the praying mantis <Emphasis Type="Italic">Tenodera aridifolia</Emphasis> suggest reciprocal predation between them

The Japanese lacertid lizard Takydromus tachydromoides and the praying mantis Tenodera aridifolia are sympatric generalist predators feeding on similar prey. To confirm reciprocal predation between them, we observed the behavioural interactions between the lizards and the mantises of different sizes in a laboratory condition. The lizards caught small mantises (from first to fifth instars), but sometimes escaped from large mantises (from sixth instar to adult). Large mantises occasionally showed catch responses to the lizards. The lizards sometimes caught the mantis without a tongue-flick response (sampling of chemical cues), and they sometimes did not catch the small mantises showing immobile or cryptic responses that prevent visual detection. These results suggested the primary role of vision on recognition of the mantis as a prey. The lizards spent a longer time to approach larger mantises. The time from orienting to catch was longer when the lizards showed tongue-flick responses. The lizard also spent a longer time before deciding to escape from the mantis than to catch it. Biological significance of these differences in timing was discussed.     

Stable and variable parameters in courtship songs of grasshoppers of the subfamily Gomphocerinae (Orthoptera,Acrididae)

The courtship behavior of seven grasshopper species of the subfamily Gomphocerinae from different localities of Russia, Ukraine and Greece was described. Not only the sounds but also the corresponding stridulatory movements of the hind legs and visual display accompanying the courtship song were analyzed. Comparison of the degree of variation in different courtship parameters showed that the most stable traits were the syllable and pulse periods. The potential role of stable and variable traits in the grasshopper courtship songs is discussed.     

Sexual dimorphism of auditory function and structure in praying mantises (Mantodea; Dictyoptera)

Sexual dimorphism of tympanate auditory systems in insects has bees described in only a few taxonomically isolated cases. However, widespread sexual dimorphism occurs in the ultrasound-sensitive, midline ear of the praying mantis.
In dimorphic species, it is always the female mantis that shows a reduction in ultrasonic hearing. The dimorphism may be mild—a difference in tuning and small reduction in sensitivity—or extreme with no evidence of audition in the female. In all but the mildest cases, the reduction in hearing is accompanied by significant anatomical divergence from the male ear structure. Two distinct metathoracic groove ('ear') types are linked to hearing reduction in the females.
Anatomical evidence of auditory sexual dimorphism appears in 34% of the 183 mantis genera examined. The dimorphic genera are widely but non-uniformly distributed within three of the four largest mantis families.
Auditory sexual dimorphism is closely correlated with dimorphism in wing length. In general, mantises with functional wings have sensitive ultrasonic hearing while those with short wings do not. These findings support the hypothesis that ultrasonic hearing in mantises is part of a defensive system against attack by echolocating bats.     

Precocious Courtship and Play in Emydid Turtles

In many mammal species, precocious sexual behaviour is a component of play. A recent model for the evolution of play fighting behaviour in muroid rodents by Pellis (1993) proposes that social play in those species has its origin in precocious sexual behaviour. Captive juvenile emydid turtles perform precocious sexual behaviour. After reviewing its occurrence, we focus on detailed observations of Pseudemys nelsoni. Precocious sexual behaviour in these turtles is characterized by the striking ‘titillation’ display, usually observed only when adult males court females. In this display, the digits of the front feet are rapidly vibrated above the eyes of the female. In our observations, both sexes performed the display, which was energetic, spontaneous, and inconclusive (in that sequences of precocious courtship typically ended when either the displayer or recipient stopped and rested, left, or started some other unrelated non-social behaviour). Displaying individuals clearly preferred conspecific recipients and often appeared to seek out specific individuals. We discuss several plausible explanations for precocious courtship in these turtles but find none entirely satisfying. The characteristics of the precocious courtship behaviour we observed most closely match those commonly listed for social play. Thus, we conclude that the most likely explanation for this behaviour is that it is a component of social play.     

Comparative Studies of Reproductive Behavior in Mantis Shrimps and Fiddler Crabs

Comparative ethological studies of closely related species canplace into clear perspective the functions of behavioral traitsas species diverge and adapt to different environments. In thispaper we contrast the reproductive behavior of species withintwo crustacean groups: the mantis shrimps (stomatopods) andthe fiddler crabs (genus Uca). For the stomatopods, we identifyprobable selective relationships between features of species'ecology and their mating systems. Population density, the kindand availability of shelters in which these shrimps live, theintensity of predation and spawning cycles all play importantroles in molding reproductive behavior in this group. Associationsbetween the ecology and mating systems of fiddler crabs havebeen discussed recently elsewhere. Here we focus on a comparativeanalysis of sexual communication in these crabs. A study ofcompetitive courtship signaling and mate choice in the fiddlercrab Uca beebei has shown that certain male courtship signalsare highly attractive probably because they exploit female sensory-responsesystems that have been molded by selection for escape from predators.Interspecific comparison of male courtship displays and theresponses of females to these displays suggest that sensoryexploitation may play an important role in the evolution ofsexual signals in the genus. Comparative studies have advancedour understanding of how natural and sexual selection affectthe reproductive behavior of both stomatopods and fiddler crabs.     

Similar yet different: differential response of a praying mantis to ant‐mimicking spiders

Batesian mimics typically dupe visual predators by resembling noxious or deadly model species. Ants are unpalatable and dangerous to many arthropod taxa, and are popular invertebrate models in mimicry studies. Ant mimicry by spiders, especially jumping spiders, has been studied and researchers have examined whether visual predators can distinguish between the ant model, spider mimic and spider non‐mimics. Tropical habitats harbour a diverse community of ants, their mimics and predators. In one such tripartite mimicry system, we investigated the response of an invertebrate visual predator, the ant‐mimicking praying mantis (Euantissa pulchra), to two related ant‐mimicking spider prey of the genus Myrmarachne, each closely mimicking its model ant species. We found that weaver ants (Oecophylla smaragdina) were much more aggressive than carpenter ants (Camponotus sericeus) towards the mantis. Additionally, mantids exhibited the same aversive response towards ants and their mimics. More importantly, mantids approached carpenter ant‐mimicking spiders significantly more than often that they approached weaver ant‐mimicking spiders. Thus, in this study, we show that an invertebrate predator, the praying mantis, can indeed discriminate between two closely related mimetic prey. The exact mechanism of the discrimination remains to be tested, but it is likely to depend on the level of mimetic accuracy by the spiders and on the aggressiveness of the ant model organism.     

A new genus of praying mantis,Chlorocalis n. gen., with two new species from the Greater Mekong region (Mantodea: Mantidae)

A new genus of praying mantis is described, Chlorocalis n. gen., with two species, C. maternaschulzei n. sp. and C. prasina n. sp. from the Greater Mekong Region. Chlorocalis is placed in Mantidae and can be immediately recognized by the presence of dark brown (male) or yellow-whitish (female) oblique stripes on the forewings, a large conical bulge on the clypeus, two large conical bulges on either side of the inferior margin of the lower frons, and the male genitalia with two distal processes on the ventral phallomere. Chlorocalis maternaschulzei is a widespread lowland species with records as west as Nakhon Ratchasima Province, Thailand, and as east as mount Núi Chúa in southern Central Vietnam. We consider this species to be of Least Concern. In contrast, C. prasina is a narrow-range mountain endemic of the Vietnamese Central Highlands in the Annamite range and is assessed by us to be Vulnerable. The new findings demonstrate the still very poor knowledge on praying mantis species diversity in South-East Asia and the urgency to resolve this taxonomic deficiency in the face of biodiversity loss.     

Effect of individual chromosomes on sex behavior of Drosophila

Dependence of intensity of Drosophila virilis male sexual behaviour on age-related differences was studied. Most active proved to be those age periods which correlated with the time of S-esterase elevated activity. Courtship displays were compared in strains 101 and 160, interstrain differences correlating with the S-esterase activity level. As S-esterase plays an important role in fertilization, this relationship seems to be non-random. It is quite possible that the level of S-esterase activity regulates male sexual activity in natural populations. Some variations in male courtship display in interspecies hybrids (D. virilis x D. littoralis) were demonstrated which depended on the gene dose of respective species. It may well be that in the bk and dt gene regions those genes are localized which are responsible for courtship behaviour. The analysis of sexual behaviour in interstrain and interspecies hybrids proved to be successful approach to elucidating possible role of separate chromosomes in sexual behaviour determination.     

A new species of Dystacta Saussure, 1871 from Nyungwe National Park,Rwanda (Insecta,Mantodea, Dystactinae)

A recent targeted entomological survey in the Republic of Rwanda has produced two conspecific male and female specimens of an undescribed species of praying mantis (Mantodea). The specimens were collected in Nyungwe National Park in May of 2013. The species is closest morphologically to Dystacta alticeps (Schaum, 1853). Therefore, a new species is described, Dystacta tigrifrutex sp. n., along with the first instar nymphs and ootheca. In addition, the previously monotypic genus Dystacta Saussure, 1871 is re-described to provide a broader definition of the genus group. Habitus images, measurement data, a key to species, natural history information, and locality data are provided.     

Testosterone and its effects on courtship in golden-collared manakins (Manacus vitellinus): seasonal, sex, and age differences

Male golden-collared manakins gather on leks and perform an acrobatic display to attract females. In temperate breeding species, testosterone (T) activation of courtship displays has been well studied. Few studies have examined T activation of displays in tropical species; even fewer have explored the activational role of T in elaborate courtship displays such as in the manakin. In some tropical species, including manakins, territorial aggression or song behavior are uncoupled from T. We have previously shown that T activates display behavior in manakin males when endogenous T levels are low in the non-courtship season. To understand how T functions in breeding birds, we examined T levels in a large group of manakins sampled during the courtship and non-courtship season. In addition, during the courtship season, we gave T implants to adult males, juvenile males, and females. We found that T levels were low during the non-courtship season and comparatively higher on average during the courtship season. However, T levels were low in many adult males during the courtship season, especially when compared to temperate breeding species. Regardless of initial endogenous T levels during the courtship season, T implants did not increase the display frequency of adult males. T-treated females and juvenile males did display under similar conditions. Our data suggest that the effects of T on manakin display vary with season, sex, and age and that high T is not necessary for display.     

Courtship and Female Competition in the Majorcan Midwife Toad,Alytes muletensis

The courtship and mating behaviour of A. muletensis are described based on observations of captive toads. Courtship is prolonged and complex, with both sexes participating actively. Either sex can escape from amplexus and may do so if the partner fails to respond appropriately to courtship manocuvres. Females control the pace and duration of courtship. Females unable to obtain a mate drop their eggs unfertilized. Both sexes produce courtship vocalizations and either sex may initiate courtship, although females were observed to do so only when they were in danger of dropping their eggs. Physical competition in the form of interference and grappling may occur between either males or females, but was observed more frequently between females. The active role played by females during courtship in this species is discussed with regard to sex-role reversal theory and opportunities for mate choice.     

Patterns of praying mantis auditory system evolution based on morphological, molecular, neurophysiological, and behavioural data

Some praying mantids have sensitive ultrasonic hearing arising from a unique 'cyclopean' ear located in the ventral metathorax. The present study explores the evolutionary history of the mantis auditory system by integrating large anatomical, neurophysiological, behavioural, and molecular databases. Using an 'auditory phylogeny' based on 13 morphological characters, we identified a primitively earless form of metathoracic anatomy in several extant taxa. In addition, there are five distinct mantis auditory systems. Three of these can be identified anatomically, and the other two can only be detected neurophysiologically. Superimposing these results onto a phylogenetic tree derived from molecular data from seven genes shows that the cyclopean mantis ear evolved once approximately 120 Mya. All the other auditory system types are either varying degrees of secondary loss, or are recent innovations that each occurred independently multiple times. The neurophysiological response to ultrasound is remarkably consistent across all taxa tested, as is the multicomponent, in-flight behaviour triggered by ultrasound. Thus, mantids have an ancient, highly conserved auditory neural–behavioural system. Although ultrasonic hearing in several insect groups evolved in response to bat predation, mantis hearing predates the appearance of bats (approximately 63 Mya) and must originally have functioned in communication, prey detection, or avoidance of nonbat predators.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 541–568.     

Mating behaviour and reproductive isolation of three species of reptile tick

The mating behaviour of three species of reptile tick, Aponomma hydrosauri, Amblyomma albolimbatum and Amb. limbatum is similar and involves a female sex pheromone which activates males to detach from their hosts and search. After contacting females, a stereotyped six-phase courtship sequence occurs. There are qualitative differences between the species in courtship behaviour at phase 3 (reversal of position by dorsally mounted males) and phase 6 (the copulation position). On-host observations of non-conspecific mating show that females of the three species have species-specific activation pheromones, which is contrary to reports in other species of tick. Such specificity should result in reproductive isolation of the three species; however, under certain circumstances it may not prevent non-conspecific contacts between the sexes. Off-host observations of courtship behaviour show that once males of the three species contact non-conspecific females, they attempt courtship and are persistent with their courtship. Rarely did non-conspecific courtships proceed beyond phase 4 of the courtship sequence, as non-conspecific females did not lift their bodies to allow males venter contact. Differences between the species in leg orientation in the copulation position, together with body size differences, are responsible for a complete barrier to successful non-conspecific copulation. The observations illustrate the role that behavioural mechanisms play in reproductively isolating these three species of tick.     

The unusual inheritance pattern of the courtship songs in closely related grasshopper species of the Chorthippus albomarginatus-group (Orthoptera: Gomphocerinae)

Sibling species of the Chorthippus albomarginatus-group are exceptional among all European grasshopper species because they have an extremely elaborate courtship behaviour. Here, we present a genetic analysis of the courtship song differences between two closely related grasshopper species Ch. albomarginatus and Ch. oschei. Measurements of seven courtship song characters and one parameter of courtship visual display were compared among parentals, F(1) and F(2) hybrids, and backcrosses. Variation in one morphological character, the number of stridulatory pegs, was studied as well. The distributions of song phenotypes were more consistent with a type-III genetic architecture, which involves complementary or duplicate pairs of loci. We suggest that this type of architecture may provide the novel elements of the hybrid songs, even more complex than the parental ones, which may offer a new starting point for sexual selection. One parameter of the visual display appeared to be controlled by a single locus, but differences in other characters of visual display could be explained by a more complex inheritance pattern. Inheritance of the peg number differences was consistent with a simple polygenic additive model. Sex linkage and/or maternal effects were not detected for any trait. We found a disproportionate influence of the Ch. albomarginatus parent on most hybrid song characteristics and the visual display, and this may support a previously suggested idea that Ch. albomarginatus is an ancestral form and Ch. oschei is a derived species.