Morphological allometry and intersexuality in horsehair-worm-infected mantids, Hierodula formosana (Mantodea: Mantidae).

Parasitic castration is a strategy used by parasites to minimize damage to the host by consuming its reproductive system, which results in the morphological alteration of the host. We determined that the forewing shape and density of the antennal sensilla of field-collected adult male mantids (Hierodula formosana), infected by horsehair worms (Chordodes formosanus) was partially feminized (intersexuality), and both male and female mantids infected by horsehair worms exhibited allometric changes in their wings and walking legs. In addition, the testes of most infected male adults disappeared or reduced in size, whereas the number of ovarioles in infected female adults was unaffected. The infection mainly influenced the structures related to host reproduction and locomotion, suggesting unbalanced energy exploitation and the reduction of parasitic virulence. In addition, the intersexuality of infected male adults indicated that sexual differentiation in insects, which researchers have considered to be an autonomous process, was influenced by the infection. The similarity of the antennae of infected male adults with those of last-instar female nymphs suggested that parasitic juvenilization may cause such feminization, but the mechanism of parasitic influence on insect sex characteristics should be studied further.

Rediscovery of Mantellias pubicornis Westwood, 1889, a rare praying mantis from the Amazon (Mantodea, Thespidae, Oligonicinae).

Many praying mantis species remain known from type specimens only. The majority of these taxa have vague taxonomic limits, as original descriptions are often very short, making strong emphasis on superficial characters (Rivera 2010). One clear example of this is the Amazonian Mantellias Westwood, 1889, a monotypical genus represented by Mantellias pubicornis Westwood, 1889.

Cohabiting With the Enemy: Comparative Population Ecology of Two Mantid Species in a Successional Old Field.

Two praying mantids, Tenodera aridifolia sinensis Saussure and Tenodera angustipennis Saussure, are commonly found in the same old-field habitats in the eastern United States and in much of temperate zone Asia. Naturally established populations of these two species were studied intensively over two consecutive years (2010 and 2011) in an old field in southeastern Virginia, to compare life history features relevant to how they coexist, or whether one or the other of them is likely to be more successful in the same habitat. Populations of both species declined about 50% from 2010 to 2011 (adults from 47 to 21 for T. a. sinensis; 37 to 20 for T. angustipennis), but T. a. sinensis oviposited 10 oothecae and T. angustipennis only one in 2011. Tenodera a. sinensis was more abundant in the study site in both years, hatched earlier, and matured and oviposited earlier than T. angustipennis. Fewer females of both species survived to maturity in 2011 than in 2010, possibly indicating a reduction in prey or habitat suitability in 2011. We suggest that T. angustipennis will always be at a disadvantage as a result of its smaller body size, because of interspecific predation (and potentially competition) from its congener, lower clutch size, and susceptibility to egg parasitism. Further, environmental variability across field habitats and years profoundly affects populations of both species in successional old fields.

Chirality of male genitalia in Otomantis casaica Giglio-Tos, 1915 (Mantodea: Hymenopodidae, Acromantinae).

Reversal of male genitalia are known in various insect orders, such as in Odonata, Orthoptera, Dermaptera, Hemiptera and Trichoptera (Schilthuizen 2007) and, within the Dictyoptera, in several species of Ectobiinae (Blattodea) (Bohn 1987), and Mantodea. Balderson (1978) first described reversal of the phallic complex in Stenomantis Saussure and Ciulfina Giglio-Tos, reporting this condition in eleven of 17 specimens representing two species of the latter-informally named as "Ciulfina sp.2" and "Ciulfina sp.7" (see Balderson 1978: 238). Subsequently, Anisyutkin & Gorochov (2004) reported the same condition at the time of describing Haania doroshenkoi from Cambodia. The male external genitalia within the Mantodea ("praying mantises") are markedly asymmetrical and generally develop in a single orientation (Klass 1997; Huber et al. 2007). Typically, the phallic complex consists of three phallic lobes surrounding the gonopore, all contained in a genital chamber between the ninth sternite and the paraprocts. Two of the three phallic lobes (phallomeres of La Greca 1955) are situated above the gonopore-one to the left and one to the right-while the third lies ventral to the genital opening. The right phallomere (RP) (Fig.1) ("right epiphallus" of Beier 1964) is usually dorsally positioned and its base extends almost completely across the wall of the genital chamber. The left phallomere (LP) (Fig.1) ("left epiphallus" of Beier 1964) is the most complex of the three lobes and it lies above the ventral phallomere (VP) (Fig.1) (hypophallus of Beier 1964).

The postabsorptive and postprandial metabolic rates of praying mantises: Comparisons across species, body masses, and meal sizes.

The metabolic rate of an animal affects the amount of energy available for its growth, activity and reproduction and, ultimately, shapes how energy and nutrients flow through ecosystems. Standard metabolic rate (SMR; when animals are post-absorptive and at rest) and specific dynamic action (SDA; the cost of digesting and processing food) are two major components of animal metabolism. SMR has been studied in hundreds of species of insects, but very little is known about the SMR of praying mantises. We measured the rates of CO2 production as a proxy for metabolic rate and tested the prediction that the SMR of mantises more closely resembles the low SMR of spiders - a characteristic generally believed to be related to their sit-and-wait foraging strategy. Although few studies have examined SDA in insects we also tested the prediction that mantises would exhibit comparatively large SDA responses characteristic of other types of predators (e.g., snakes) known to consume enormous, protein-rich meals. The SMR of the mantises was positively correlated with body mass and did not differ among the four species we examined. Their SMR was best described by the equation µW=1526*g0.745 and was not significantly different from that predicted by the standard 'insect-curve'; but it was significantly higher than that of spiders to which mantises are ecologically more similar than other insects. Mantises consumed meals as large as 138% of their body mass and within 6-12h of feeding and their metabolic rates doubled before gradually returning to prefeeding rates over the subsequent four days. We found that the SDA responses were isometrically correlated with meal size and the relative cost of digestion was 38% of the energy in each meal. We conclude that mantises provide a promising model to investigate nutritional physiology of insect predators as well as nutrient cycling within their ecological communities.

Habitat Preference of German Mantis religiosa Populations (Mantodea: Mantidae) and Implications for Conservation.

In Germany, the thermophilic European mantid (Mantis religiosa L.) is endangered. Here, we study habitat requirements during its life, and discuss the applicability of the two conservation measures grazing and mowing to this insect species. At two study sites in south-western Germany, which were each subject to one of these measures, we recorded structural and climatic conditions within different microhabitats. We also conducted capture-mark-recapture studies for adult M. religiosa, and mapped adult roosting, oothecae deposition, egg hatching, and imaginal molting in microhabitats over two vegetation periods. In order to assess microhabitat preference of M. religiosa during its life, and identify climatic conditions driving preferences, we applied the Lille habitat preference index and conducted logistic regression analysis for life phases. Our results suggest that temperature is important for egg and nymph development. For egg deposition, females preferred solid substrates with high heat-storing capacities, as those attenuate the negative influence of cold weather periods on egg development. Being ambush predators, males and females preferred roosting sites with sufficient shelter and high prey abundance. Contrary to our expectation, the conservation measures reduced adult population sizes, and presumably reproduction rates. We thus suggest that mowing during the adult phase should reduce vegetation height to a moderate level to keep prey abundance high. Mowing with a clearing saw or grazing over a short period in small fenced areas should be preferred over prolonged grazing, as grazers collaterally stamp down the vegetation. Grazers indirectly reduce prey availability by deteriorating prevailing microclimatic conditions.

Mantises exchange angular momentum between three rotating body parts to jump precisely to targets.

Flightless animals have evolved diverse mechanisms to control their movements in air, whether falling with gravity or propelling against it. Many insects jump as a primary mode of locomotion and must therefore precisely control the large torques generated during takeoff. For example, to minimize spin (angular momentum of the body) at takeoff, plant-sucking bugs apply large equal and opposite torques from two propulsive legs [1]. Interacting gear wheels have evolved in some to give precise synchronization of these legs [2, 3]. Once airborne, as a result of either jumping or falling, further adjustments may be needed to control trajectory and orient the body for landing. Tails are used by geckos to control pitch [4, 5] and by Anolis lizards to alter direction [6, 7]. When falling, cats rotate their body [8], while aphids [9] and ants [10, 11] manipulate wind resistance against their legs and thorax. Falling is always downward, but targeted jumping must achieve many possible desired trajectories. We show that when making targeted jumps, juvenile wingless mantises first rotated their abdomen about the thorax to adjust the center of mass and thus regulate spin at takeoff. Once airborne, they then smoothly and sequentially transferred angular momentum in four stages between the jointed abdomen, the two raptorial front legs, and the two propulsive hind legs to produce a controlled jump with a precise landing. Experimentally impairing abdominal movements reduced the overall rotation so that the mantis either failed to grasp the target or crashed into it head first.

A survey of the praying mantises of Rwanda, including new records (Insecta, Mantodea).

We report the results of two surveys targeting praying mantises in four localities in Rwanda, specifically Akagera National Park, Nyungwe National Park, Volcanoes National Park, and the Arboretum de Ruhande at the National University of Rwanda. Using an assortment of collecting techniques, including metal halide light traps, sweep netting vegetation and general searching, we obtained 387 adult and 352 juvenile specimens, representing 41 species. A total of 28 novel species records for Rwanda are added to the 18 previously recorded species for the country, in addition to 20 novel species records for the broader region, including neighbouring Uganda and Burundi. This study provides high resolution images of the dorsal habitus of both sexes of representative species, both pinned and living. Species distribution records are presented and discussed. With a 155% increase in species recorded from Rwanda, this survey illustrates the need for further taxonomic work in the region.

The Ontogenetically Variable Trophic Niche of a Praying Mantid Revealed by Stable Isotope Analysis.

Praying mantids have been shown to exert strong influences on arthropod community composition. However, they may not occupy the same trophic level throughout their lives. Trophic shifting over a life cycle could explain the documented variation in results from field studies, but specific interactions of predators within food webs have been difficult to determine simply by comparing control and treatment assemblages in field experiments. We examined the trophic position of the Chinese praying mantid, Tenodera aridifolia sinensis (Saussure), using stable isotope analysis (SIA). We measured the δ(13)C and δ(15)N of field-collected arthropods, and of laboratory groups of mantids fed known diets of these arthropods chosen from the most abundant trophic guilds: herbivores (sap feeders and plant chewers), and carnivores. We also collected mantids from the field over a growing season and compared their SIA values to those of the laboratory groups. Both δ(13)C and δ(15)N of mantids fed carnivorous prey (spiders or other mantids) were higher than those fed herbivores (grasshoppers). SIA values from field-collected mantids were highly variable, and indicated that they did not take prey from trophic guilds in proportion to their abundances, i.e., were not frequency-dependent predators. Further, δ(15)N decreased from a high at egg hatch to a low at the third instar as early nymphs fed mainly on lower trophic levels, and increased steadily thereafter as they shifted to feeding on higher levels. We suggest that the community impact of generalist predators can be strongly influenced by ontogenetic shifts in diet.

Extreme convergence in egg-laying strategy across insect orders.

The eggs of stick and leaf insects (Phasmatodea) bear strong resemblance to plant seeds and are commonly dispersed by females dropping them to the litter. Here we report a novel egg-deposition mode for Phasmatodea performed by an undescribed Vietnamese species of the enigmatic subfamily Korinninae that produces a complex egg case (ootheca), containing numerous eggs in a highly ordered arrangement. This novel egg-deposition mode is most reminiscent of egg cases produced by members of unrelated insect orders, e.g. by praying mantises (Mantodea) and tortoise beetles (Coleoptera: Cassidinae). Ootheca production constitutes a striking convergence and major transition in reproductive strategy among stick insects, viz. a shift from dispersal of individual eggs to elaborate egg concentration. Adaptive advantages of ootheca formation on arboreal substrate are likely related to protection against parasitoids and desiccation and to allocation of specific host plants. Our phylogenetic analysis of nuclear (28S, H3) and mitochondrial (COI, COII) genes recovered Korinninae as a subordinate taxon among the species-rich Necrosciinae with Asceles as sister taxon, thus suggesting that placement of single eggs on leaves by host plant specialists might be the evolutionary precursor of ootheca formation within stick insects.

Different paths to sexual size dimorphism in two praying mantids, Pseudomantis albofimbriata and Hierodula majuscula.

Sexual size dimorphism (SSD) is widespread among diverse animal taxa and has attracted the attention of evolutionary biologists for over a century. SSD is likely to be adaptive and the result of divergent selection on different size optima for males and females, given their different roles in reproduction. The developmental trajectory leading to SSD may help us to understand how selection acts on male and female size. Here, we describe the growth and development of two Australian praying mantids, Pseudomantis albofimbriata and Hierodula majuscula including the number of moults, time to adulthood, size at each moult, and the degree of SSD. While both species exhibit the common pattern of female-biased SSD, the number of moults required for individuals to reach adulthood differed between males and females and between species. Despite their larger adult size, P. albofimbriata females require fewer moults and less time than males to reach adulthood, but are significantly larger than males from the second instar onwards. In contrast, H. majuscula males reached adulthood in fewer moults, and less time than females, however males and females did not differ in size until females went through their final moult into adulthood. H. majuscula also required more time and more moults to reach adulthood than P. albofimbriata. We discuss these different developmental pathways in light of the existing knowledge of reproductive biology for each species. We also suggest that these differences may relate to the different phenologies that occur in strongly seasonal temperate environments compared with those in the tropics. This study provides evidence that SSD can result from two different patterns of growth and development in closely related species.

Antennal development in the praying mantis (Tenodera aridifolia) highlights multitudinous processes in hemimetabolous insect species.

Insects possess antennae equipped with a large number of segments (flagellomeres) on which sensory organs (sensilla) are located. Hemimetabolous insects grow by molting until they reach adulthood. In these species, the sensory structures develop and mature during each stage of development; new flagellomeres are generated at each molt elongating the antennae, and new sensilla appear. The praying mantis (Tenodera aridifolia) is a hemimetabolous insect with 7 different instars before it reaches adulthood. Because their antennae are provided with an atypical sensillar distribution, we previously suggested that their antennae develop with a different mechanism to other hemimetaboulous insect species. In the present study, we measured the number, length and width of flagellomeres along the antennae in nymph and adult mantis Tenodera aridifolia. For this study, we developed a new and innovative methodology to reconstruct the antennal development based on the length of flagellomeres. We observed and confirmed that the antennae of mantises develop with the addition of new segments at two distinct sites. In addition, we constructed a complete database of the features of the flagellum for each stage of development. From our data, we found that sexual dimorphism appears from the 6 instar (larger number and wider flagellomeres in males) in accordance with the appearance of their genital apparatus. The antennal sexual dimorphism completes at adulthood with longer flagellomeres and the emergence of a huge number of grooved peg sensilla in males during the last molting, which suggests once again their function as sex-pheromone receptive sensilla.

Allometry and ontogeny in Callibia diana Stål (Mantodea: Acanthopidae).

The life-cycle of Callibia diana Stål is described and linear and geometric morphometrics are used for studying allometrics and shape changes throughout this neotropical mantid species' life-cycle. Significant changes were expected in the allometry and shape of the raptorial leg and abdomen, given the importance of hunting and reproduction. The allometric slopes were obtained by using total length as the independent variable. Geometric morphometrics of landmarks were used for frontal femur and tibia. Hunting and reproduction-related structures had the steepest slopes and positive allometries. Negative growth of both disc width and head width found in the last moulting event may be a consequence of prothoracic muscle growth which is responsible for predatory strike strength. The tibial claw and femur of the raptorial leg become larger, while their spines become more orthogonal to the longitudinal axes which may facilitate prey retention. These changes in mantid shape throughout ontogeny were consistent and suggested the resource allocation and development programming of the body that improved reaching distance and prey retention.

Allometry and ontogeny in Callibia diana Stål (Mantodea: Acanthopidae)

The life-cycle of Callibia diana Stål is described and linear and geometric morphometrics are used for studying allometrics and shape changes throughout this neotropical mantid species' life-cycle. Significant changes were expected in the allometry and shape of the raptorial leg and abdomen, given the importance of hunting and reproduction. The allometric slopes were obtained by using total length as the independent variable. Geometric morphometrics of landmarks were used for frontal femur and tibia. Hunting and reproduction-related structures had the steepest slopes and positive allometries. Negative growth of both disc width and head width found in the last moulting event may be a consequence of prothoracic muscle growth which is responsible for predatory strike strength. The tibial claw and femur of the raptorial leg become larger, while their spines become more orthogonal to the longitudinal axes which may facilitate prey retention. These changes in mantid shape throughout ontogeny were consistent and suggested the resource allocation and development programming of the body that improved reaching distance and prey retention.