Species delimitation of the praying mantis emHierodula/em empatellifera/em (Audinet-Serville) based on morphological and molecular characters (Mantodea: Mantidae).

An integrative taxonomic analysis of Hierodula patellifera (Audinet-Serville) is presented based on morphological and molecular characters (COI, 28S rDNA). During repeated trips to the Korean peninsula, we collected unusual specimens from Wanju-gun. They were similar to H. patellifera, but can be distinguished by a larger body size, the number and shape of spines on foreleg, and the shape of male genitalia. To examine the phenotypes and delimit H. patellifera from fourteen sampled populations, we used forecoxal spines and male genitalia as key morphological characters, as well as molecular data including gene tree monophyly and genetic divergence data. The molecular analyses (p-distance, neighbor-joining, and parsimony analyses) did not separate the specimens as two distinct species. The diagnostic characters of H. patellifera are illustrated with habitus images.

Evolutionary diversification in the raptorial forelegs of Mantodea: Relations to body size and depth perception.

Limb proportions have evolved among animals to meet functional demands among diverse environments. Studies from terrestrial, vertebrate locomotion have demonstrated that variation in limb proportions have adaptively evolved so animals can perform in a given environment. Most of the research on limb proportion evolution is among vertebrates and terrestrial locomotion, with little information on limb segment evolution in invertebrates or for other functional roles. For example, among invertebrates, multisegmented raptorial forelimbs have evolved multiple times independently to capture prey, but there is little information on the adaptive evolution and diversity of these limbs. Furthermore, as feeding performance is influenced by the sensory system, few studies have examined the coevolution of sensory-motor systems. Using mantises (Mantodea) I examined forelimb diversification among 97 species with a combination of methods, including ternary plots for morphospace visualization, phylogenetically informed allometric relationships, and comparison of evolutionary rates of diversification. Furthermore, using head width as a proxy for depth perception, I examined the correlated evolution of foreleg diversity with depth perception. The results show that among the three segments of the foreleg, the tibia is the smallest, most diverse, and has the highest rate of evolution after body size corrections. Furthermore, while all foreleg segments were related to head width, head width explained the most variation in tibial length compared with other foreleg segments. The results suggest a potential adaptive functional role of tibia length related to the displacement or force produced in this mechanical lever. Furthermore, results from this study support distinct ecomorphs of mantises, as several independent evolutions to grass mimicry evolve similar morphologies. RESEARCH HIGHLIGHT: This study demonstrates interspecific variation among segments of an invertebrate raptorial foreleg. Among Mantodea species the tibia is the most diverse and is related to a proxy for depth perception, while the other segments had strong relationships with body size. This suggests an adaptive, functional role of the tibia during prey capture.

A New Species of Praying Mantis from Peru Reveals Impaling as a Novel Hunting Strategy in Mantodea (Thespidae: Thespini).

A new species of lichen-mimicking praying mantis, Carrikerella simpira n. sp., is described from Tingo María region in Peru. The new species differs from its congeners in having reduced tergal lobes, a relatively sinuous pronotum, and it is found in the highland tropical rainforest of the Central Andes. Behavioral observations conducted on captive individuals revealed that juveniles and adults hunt by impaling prey using modified foretibial structures. Anatomical examinations of the incumbent trophic structures revealed functional adaptations for prey impaling in the foretibiae, primarily consisting of prominent, forwardly oriented, barbed spines. We provide an overall description of this novel hunting behavior in Mantodea and hypothesize on its evolutionary origin and adaptive significance for the Thespidae.

Three-dimensional atlas of thoracic ganglia in the praying mantis, Tenodera aridifolia.

The praying mantis is a good model for the study of motor control, especially for investigating the transformation from sensory signals into motor commands. In insects, thoracic ganglia (TG) play an important role in motor control. To understand the functional organization of TG, an atlas is useful. However, except for the fruitfly, no three-dimensional atlas of TG has not been reported for insects. In this study, we generated a three-dimensional atlas of prothoracic, mesothoracic, and metathoracic ganglia in the praying mantis (Tenodera aridifolia). First, we observed serial sections of the prothoracic ganglion stained with hematoxylin and eosin to identify longitudinal tracts and transverse commissures. We then visualized neuropil areas by immunostaining whole-mount TG with an anti-synapsin antibody. Before labeling each neuropil area, standardization using the iterative shape averaging method was applied to images to make neuropil contours distinct. Neuropil areas in TG were defined based on their shape and relative position to tracts and commissures. Finally, a three-dimensional atlas was reconstructed from standardized images of the TG. The standard TG are available at the Comparative Neuroscience Platform website (cns.neuroinf.jp/modules/xoonips/detail.php?item_id=11946) and can be used as a common reference map to combine the anatomical data obtained from different individuals.

Unraveling the functional organization of lobula complex in the mantis brain by identification of visual interneurons.

The praying mantis shows broad repertories of visually guided behaviors such as prey recognition and defense against collision. It is likely that neurons in the lobula complex (LOX), the third visual neuropil in the optic lobe, play significant roles in these behaviors. The LOX in the mantis brain consists of five neuropils: outer lobes 1 and 2 (OLO1 and OLO2); anterior lobe (ALO); dorsal lobe (DLO); and stalk lobe (SLO), and ALO comprise ventral and dorsal subunits, ALO-V and ALO-D. To understand the functional organization of LOX, intracellular electrodes were used for recording from and staining neurons in these neuropils of the mantis (Tenodera aridifolia). The neurons belonged to three categories based on their response properties and morphologies. First, tangential ALO-V neurons projecting to ventromedial neuropils (VMNP) (TAproM1 and 2), tangential DLO (or ALO-D) neurons projecting to VMNP (TDproM1 and 2), and tangential ALO-V centrifugal neurons (TAcen) all showed directional sensitivity and sustained excitation to gratings drifting in preferred direction (outward-downward, inward-upward, outward-upward, inward-downward, and inward, respectively). Second, tangential OLO neurons projecting to VMNP or ventrolateral neuropils (VLNP) (TOproM or TOproL), columnar OLO commissural neurons (COcom), and SLO commissural neurons (Scom) all showed strong excitation to 2°-8° moving squares but little excitations to drifting gratings. COcom and SLO neurons ramified in both left and right LOX. Last, the class of tangential ALO-V neurons projecting to VLNP (TAproL1, 2, and 3) responded best to looming circles and showed little excitation to receding, darkening, and lightening circles.

Epaphroditidae sensu novo, an Endemic Caribbean Family of Morphologically Divergent Praying Mantises (Insecta, Mantodea).

Three endemic Caribbean praying mantis genera with a complex taxonomic history were recently discovered to be part of a lineage that colonized the Caribbean region during the Cretaceous period (Svenson & Rodrigues, Proc R Soc B Biol Sci 284, 2017). In all classification systems proposed up to now, the three genera, Callimantis, Epaphrodita, and Gonatista, were never considered as close relatives, a reflection of their divergent morphology. More recently, the genus Brancsikia was placed with Epaphrodita in a family based on the similarity of camouflage-related morphology. To address recent phylogenetic results that do not track current classification, we compared the morphology of the three Caribbean genera with each other and representative members of traditional or current family groups. Our morphological analysis of external and male genital characters provides strong support for the Caribbean lineage despite the divergent morphological evolution present in the three genera. We raise this Caribbean lineage to family status by employing a precedent family-group name, Epaphroditidae Brunner de Wattenwyl, 1893 sensu novo. We remove Brancsikia from our new concept of Epaphroditidae, rendering the genus incertae sedis.

Template for the neural control of directed stepping generalized to all legs of MantisBot.

We previously developed a neural controller for one leg of our six-legged robot, MantisBot, that could direct locomotion toward a goal by modulating leg-local reflexes with simple descending commands from a head sensor. In this work, we successfully apply an automated method to tune the control network for all three pairs of legs of our hexapod robot MantisBot in only 90 s with a desktop computer. Each foot's motion changes appropriately as the body's intended direction of travel changes. In addition, several results from studies of walking insects are captured by this model. This paper both demonstrates the broad applicability of this control method for robots, and suggests neural mechanisms underlying observations from walking insects.

Organization of the antennal lobes in the praying mantis (Tenodera aridifolia).

Olfaction in insects plays pivotal roles in searching for food and/or for sexual partners. Although many studies have focused on the olfactory processes of nonpredatory insect species, little is known about those in predatory insects. Here, we investigated the anatomical features of the primary olfactory center (antennal lobes) in an insect predator whose visual system is well developed, the praying mantis Tenodera aridifolia. Both sexes of T. aridifolia were found to possess 54 glomeruli, and each glomerulus was identified based on its location and size. Moreover, we found a sexual dimorphism in three glomeruli (macroglomeruli) located at the entrance of the antennal nerves, which are 15 times bigger in males than their homologs in females. We additionally deduced the target glomeruli of olfactory sensory neurons housed in cognate types of sensilla by degenerating the sensory afferents. The macroglomeruli received sensory inputs from grooved peg sensilla, which are present in a large number at the proximal part of the males' antennae. Furthermore, our findings suggest that glomeruli at the posteriodorsal part of the antennal lobes receive sensory information from putative hygro- and thermosensitive sensilla. The origins of projections connected to the protocerebrum are also discussed. J. Comp. Neurol. 525:1685-1706, 2017. © 2016 Wiley Periodicals, Inc.

Taxonomic revision of Stagmatoptera Burmeister, 1838 (Mantodea: Mantidae, Stagmatopterinae).

Stagmatoptera Burmeister, 1838 includes medium to large-sized (49.8-98.7mm) praying mantises distributed in the Neotropical region. They are characterized for having a central, circular spot on each forewing, and 2-4 carinae in the head's frontal shield. This study is an updated revision of the genus on the basis of type and non-type material deposited in various scientific collections from Europe and the Americas. The following nomenclatural procedures were conducted: Stagmatoptera nova is a new junior synonym of Stagmatoptera pia, Stagmatoptera ignota as the junior synonym of Stagmatoptera femoralis, and Stagmatoptera flavipennis is reinstated as the junior synonym of Stagmatoptera supplicaria. Two new species are described: Stagmatoptera diana Rodrigues sp. n. from Colombia and Venezuela, and Stagmatoptera cerdai Rodrigues sp. n. from Trinidad & Tobago and Venezuela. Stagmatoptera now includes 14 species, two of which (Stagmatoptera abdominalis and Stagmatoptera indicator) are considered as species inquirenda. We provide detailed morphological descriptions of all species, a species-level identification key, in addition to abundant visual material to assist identification. Female genitalia, a structure seldom used for taxonomic purposes, was a useful character system to distinguish among species of the genus. Distribution maps for all species are also provided.

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).

Take-off speed in jumping mantises depends on body size and a power-limited mechanism.

Many insects such as fleas, froghoppers and grasshoppers use a catapult mechanism to jump, and a direct consequence of this is that their take-off velocities are independent of their mass. In contrast, insects such as mantises, caddis flies and bush crickets propel their jumps by direct muscle contractions. What constrains the jumping performance of insects that use this second mechanism? To answer this question, the jumping performance of the mantis Stagmomantis theophila was measured through all its developmental stages, from 5 mg first instar nymphs to 1200 mg adults. Older and heavier mantises have longer hind and middle legs and higher take-off velocities than younger and lighter mantises. The length of the propulsive hind and middle legs scaled approximately isometrically with body mass (exponent=0.29 and 0.32, respectively). The front legs, which do not contribute to propulsion, scaled with an exponent of 0.37. Take-off velocity increased with increasing body mass (exponent=0.12). Time to accelerate increased and maximum acceleration decreased, but the measured power that a given mass of jumping muscle produced remained constant throughout all stages. Mathematical models were used to distinguish between three possible limitations to the scaling relationships: first, an energy-limited model (which explains catapult jumpers); second, a power-limited model; and third, an acceleration -: limited model. Only the model limited by muscle power explained the experimental data. Therefore, the two biomechanical mechanisms impose different limitations on jumping: those involving direct muscle contractions (mantises) are constrained by muscle power, whereas those involving catapult mechanisms are constrained by muscle energy.

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.

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.

Phylogeny of Dictyoptera: Dating the Origin of Cockroaches, Praying Mantises and Termites with Molecular Data and Controlled Fossil Evidence.

Understanding the origin and diversification of organisms requires a good phylogenetic estimate of their age and diversification rates. This estimate can be difficult to obtain when samples are limited and fossil records are disputed, as in Dictyoptera. To choose among competing hypotheses of origin for dictyopteran suborders, we root a phylogenetic analysis (~800 taxa, 10 kbp) within a large selection of outgroups and calibrate datings with fossils attributed to lineages with clear synapomorphies. We find the following topology: (mantises, (other cockroaches, (Cryptocercidae, termites)). Our datings suggest that crown-Dictyoptera-and stem-mantises-would date back to the Late Carboniferous (~ 300 Mya), a result compatible with the oldest putative fossil of stem-dictyoptera. Crown-mantises, however, would be much more recent (~ 200 Mya; Triassic/Jurassic boundary). This pattern (i.e., old origin and more recent diversification) suggests a scenario of replacement in carnivory among polyneopterous insects. The most recent common ancestor of (cockroaches + termites) would date back to the Permian (~275 Mya), which contradicts the hypothesis of a Devonian origin of cockroaches. Stem-termites would date back to the Triassic/Jurassic boundary, which refutes a Triassic origin. We suggest directions in extant and extinct species sampling to sharpen this chronological framework and dictyopteran evolutionary studies.

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.

Four new species of Ciulfina Giglio-Tos, 1915 (Mantodea: Liturgusidae, Liturgusinae) from the Northern Territory, Australia .

The praying mantid genus Ciulfina Giglio-Tos includes many small, gracile tree-trunk dwelling species found throughout northern Australia. Four new species of Ciulfina: C. annecharlotteae, C. herbersteinae, C. ianrichardi, and C. terrymariceae are formally described on the basis of male genital morphology.

A revision of Rhodomantis Giglio-Tos, 1917 (Mantodea: Mantidae: Mantinae).

The genus Rhodomantis Giglio-Tos is revised. A total of nine species are recognised. Seven new species, R. helenae, R. kimberley, R. macula, R. microptera, R. mitchell, R. napier, and R. rentzi are described. Rhodomantis carinicollis Werner and R. gracilis Tindale are new synonyms of R. queenslandica Sjöstedt.

A revision of Apteromantis (Mantodea: Mantidae, Amelinae): a comprehensive approach to manage old taxonomic and conservation problems .

The genus Apteromantis Werner, 1931 comprises two species of wingless mantids, the Iberian A. aptera (Fuente, 1894) and the North African A. bolivari (Werner, 1929). Although A. aptera and A. bolivari have been traditionally considered as separate and valid species, their external appearance is quite similar and no comprehensive taxonomic study has analyzed their morphological and genetic characteristics. This taxonomic uncertainty has important implications for conservation because A. aptera is considered an Iberian endemic and the only praying mantis protected by international laws. In this study, we apply a comprehensive approach, including quantitative morphological and molecular analyses, to shed new light on the taxonomic and conservation status of the genus Apteromantis and the putative species. We have found that the Iberian and North African specimens analyzed herein significantly differ in female head shape, male genitalia morphology and several other traits related to body size. Molecular data suggest the presence of two main lineages, with sequence divergence rates of approximately 4 %, which are within the range reported for other well defined insect species. Overall, this study supports that A. aptera and A. bolivari are valid species despite their ecological and morphological similarity and highlights the importance of comprehensive approaches to resolve old taxonomic and conservation problems.