Anatomical organization of the cerebrum of the praying mantis Hierodula membranacea.

Many predatory animals, such as the praying mantis, use vision for prey detection and capture. Mantises are known in particular for their capability to estimate distances to prey by stereoscopic vision. While the initial visual processing centers have been extensively documented, we lack knowledge on the architecture of central brain regions, pivotal for sensory motor transformation and higher brain functions. To close this gap, we provide a three-dimensional (3D) reconstruction of the central brain of the Asian mantis, Hierodula membranacea. The atlas facilitates in-depth analysis of neuron ramification regions and aides in elucidating potential neuronal pathways. We integrated seven 3D-reconstructed visual interneurons into the atlas. In total, 42 distinct neuropils of the cerebrum were reconstructed based on synapsin-immunolabeled whole-mount brains. Backfills from the antenna and maxillary palps, as well as immunolabeling of γ-aminobutyric acid (GABA) and tyrosine hydroxylase (TH), further substantiate the identification and boundaries of brain areas. The composition and internal organization of the neuropils were compared to the anatomical organization of the brain of the fruit fly (Drosophila melanogaster) and the two available brain atlases of Polyneoptera-the desert locust (Schistocerca gregaria) and the Madeira cockroach (Rhyparobia maderae). This study paves the way for detailed analyses of neuronal circuitry and promotes cross-species brain comparisons. We discuss differences in brain organization between holometabolous and polyneopteran insects. Identification of ramification sites of the visual neurons integrated into the atlas supports previous claims about homologous structures in the optic lobes of flies and mantises.

Amazons Are Back: Absence of Males in a Praying Mantis from Uruguayan Savannas.

With a broad distribution throughout South America, Brunneria subaptera (Saussure) (Coptopterygidae) is the most abundant praying mantis species in Uruguay, mostly associated with grassland vegetation. Their body coloration can vary between green or brown sepia. The recent catalog of Uruguayan mantids showed that all specimens of B. subaptera from the collections in Uruguay were females, suggesting an absence of males in natural populations. The aim of this study was to determine the occurrence of female-only populations of B. subaptera in Uruguay. Moreover, we analyzed the genetic variation and estimated the time of the most recent common ancestor of the species. We performed bimonthly samplings in two localities in Uruguay for 1 year and we sexed the individuals. We extracted DNA from the abdominal tissue of adult females and sequenced a fragment of the mitochondrial cox1 gene to assess the genetic diversity and to estimate diversification times. We did not find any males in either of the studied localities. The age estimates revealed a recent origin of the species (2.33 Ma), and there was no genetic differentiation between the individuals from Uruguayan localities. This absence of males of B. subaptera documented in the sampled locations suggests that the species reproduces by thelytokous parthenogenesis. This study opens promising venues for future research into reproductive strategies and polymorphism in South-American praying mantises that inhabit one of the largest areas of grasslands in the world, currently under dramatic deterioration and reduction.

Animal locomotion: Wing-like femoral lobes help orchid mantid nymphs glide.

The femoral lobes of the orchid mantis give this fierce predator a flower-like appearance, but they also assist in gliding, showing that form can match function in more ways than one.

Biology, ecology, and biogeography of eremic praying mantis Blepharopsis mendica (Insecta: Mantodea).

Background: Blepharopsis mendica (Fabricius, 1775) is a large mantid species found from the Canary Islands across North Africa, the Middle East, and Pakistan. Research on this species has been limited, especially in Iran, despite the country's potential significance for studying its biology and distribution. Adults of this species are easily recognizable by their marble-white pattern and rhomboidal leaf-like pronotum. They are sit-and-wait predators that inhabit various open environments. Methods: Field observations were conducted across various regions of the Egyptian Flower mantis (Blepharopsis mendica) global distribution, with a focus on Morocco, Tunisia, and Iran. Distribution data for B. mendicawere gathered from fieldwork, museum collections, online biodiversity databases, and publications, totaling 593 occurrence points. Ecological niche modeling was performed using environmental data, and various models were evaluated for suitability. Phylogeographic analyses involved DNA sequencing and construction of a haplotype network to examine genetic relationships between populations. Divergence time estimation and biogeographical range expansion models were applied to explore historical distribution shifts of the species across different regions. The study provided comprehensive insights into the biology, distribution, and genetic history of B. mendica. Results: We provide information on the life cycle, ootheca, defense behavior, habitat, and biogeography of the Egyptian Flower mantis Blepharopsis mendica. This mantid is an overwintering univoltine species with nymphs emerging in summer and becoming adults in spring. In the wild, females start oviposition in April and can lay their first ootheca within a week after mating. The species is distributed from the Canary Islands to Pakistan in the dry belt. Thus, its distribution is associated with xeric areas or desert and semi-desert habitats. Phylogeographic analyses revealed three major genetic lineages, (i) in the Maghreb, (ii) from Egypt via Arabia to Iran (with internal substructures), and (iii) likely in Pakistan; the estimated onset of differentiation into these lineages is of Pleistocene age. Defense behavior involves flying away or extending wings broadly and lifting forelegs. Performing laboratory breeding, we documented life cycle and color changes from first instar to adulthood. Due to overwintering, the last larval instar needs considerably longer than the others. At 25 °C (±2), average adult life span was 118 days (±6 SD) for females (range: 100-124) and 46 days (±5 SD) for males (range: 39-55), with a significant difference among sexes. On average, oothecae contained 32.3 eggs (±10.1 SD) and the mean incubation period was 36.8 days (±2.9 SD). We did not find evidence of parthenogenesis. In general, the biology of B. mendica shows a variety of adaptations to its often extreme and little predictable type of habitat.

Mantodea (Insecta) in the Invertebrate Collection of Instituto Nacional de Pesquisas da Amaznia (INPA), Manaus, Brazil.

Increasingly, scientific collections play a fundamental role in the conservation and analysis of biodiversity as a witness and repository of ecosystem organisms at risk of losing their species. The Invertebrate Collection of the Instituto Nacional de Pesquisas da Amaznia (INPA) safeguards biological information, mainly from the Amazon biome, one of the most diverse and threatened ecosystems on the planet. Mantodea specimens deposited dry in the INPA Collection were reviewed. A total of 2,721 specimens were found deposited until the year 2019. A total of 194 species and 81 genera were found with representatives of all the neotropical and several "Old-World" families. Families such as Mantidae, Photinaidae, and Thespidae represented the highest percentages of specimens collected within the sample, with 30%, 11%, and 9%, respectively. Four genera and fourteen species are recorded for the first time for Brazil, bringing the number of species to 247. This officially makes Brazil the most diverse country in terms of Mantodea fauna on the planet. The Collection stands out for being the most diverse reported in Latin America, even with several species yet to be described. Presently it has seven primary type specimens and 26 secondary type specimens. The excellent state of conservation of the specimens, the number of specimens, and the diversity represented have made the INPA Collection an international benchmark for Amazonian and Neotropical studies on Mantodea.

Petal-shaped femoral lobes facilitate gliding in orchid mantises.

To glide in forest canopies, arboreal vertebrates evolved various skin-derived aerodynamic structures, such as patagial membranes or webbing, but no comparable structure has been reported from wingless arboreal arthropods.1,2,3 Orchid mantises (Hymenopus coronatus) have been traditionally considered a textbook example of flower mimicry for ∼200 years due to their highly expanded, petal-shaped femoral lobes. However, the empirical evidence substantiating the petal-mimicry function of the femoral lobes has not been entirely conclusive.4,5,6 Observational and experimental evidence suggests that these lobes do not contribute to flower mimicry for luring pollinators6,7 and likely serve other functions.7,8 After observing their aerial escape initiated with active jumping, we hypothesized that orchid mantises can glide and that their femoral lobes are used for gliding. Through behavioral investigations and morphological analyses, we show that orchid mantis nymphs are excellent gliders, exhibiting the shallowest gliding trajectories observed in terrestrial invertebrates.9,10,11,12,13 The lobe extensions on their femoral segments are cambered airfoils, which increase the mantis projected area by ∼36% and play a vital role in the aerodynamic underpinning of the observed gliding. Despite a 165-fold increase in body mass throughout ontogeny, older female mantis nymphs maintained a persistent gliding capability. We further showed a notable 40%-56% reduction in wing loading attributed to the positive size allometry of these lobes, indicating a clear promotion of gliding throughout ontogeny. This is the first documentation of gliding-adapted "leg wings" in a wingless arthropod. The evolution of such structures is potentially common among arboreal arthropods and demands a systematic re-examination.

Crypsis by background matching and disruptive coloration as drivers of substrate occupation in sympatric Amazonian bark praying mantises.

Background matching and disruptive coloration are common camouflage strategies in nature, but few studies have accurately measured their protective value in living organisms. Amazon's Bark praying mantises exhibit colour patterns matching whitish and greenish-brown tree trunks. We tested the functional significance of background matching and disruptive coloration of different praying mantis morphospecies (white, grey and green) detected by DNA barcoding. Through image analysis, avian visual models and field experiments using humans as potential predators, we explored whether the background occupation of mantises provides camouflage against predation. Data were obtained for individuals against their occupied tree trunks (whitish or greenish-brown) and microhabitats (lichen or bryophyte patches), compared to non-occupied trunks. White and grey mantises showed lower colour contrasts against occupied trunks at the scale of tree trunk, with no differences in luminance contrasts. Conversely, green mantises showed lower colour and luminance contrasts against microhabitats and also exhibited high edge disruption against greenish-brown trunks. The camouflage of white and green mantis models against colour-matching trunks increased search time and reduced encounter distance of human predators. We highlight the importance of camouflage strategies at different spatial scales to enhance individual survival against predators. Specifically, we present a stunning study system to investigate the relationship of phylogenetically related species that use camouflage in sympatry.

Massive horizontal gene transfer and the evolution of nematomorph-driven behavioral manipulation of mantids.

To complete their life cycle, a wide range of parasites must manipulate the behavior of their hosts.1 This manipulation is a well-known example of the "extended phenotype,2" where genes in one organism have phenotypic effects on another organism. Recent studies have explored the parasite genes responsible for such manipulation of host behavior, including the potential molecular mechanisms.3,4 However, little is known about how parasites have acquired the genes involved in manipulating phylogenetically distinct hosts.4 In a fascinating example of the extended phenotype, nematomorph parasites have evolved the ability to induce their terrestrial insect hosts to enter bodies of water, where the parasite then reproduces. Here, we comprehensively analyzed nematomorphs and their mantid hosts, focusing on the transcriptomic changes associated with host manipulations and sequence similarity between host and parasite genes to test molecular mimicry. The nematomorph's transcriptome changed during host manipulation, whereas no distinct changes were found in mantids. We then discovered numerous possible host-derived genes in nematomorphs, and these genes were frequently up-regulated during host manipulation. Our findings suggest a possible general role of horizontal gene transfer (HGT) in the molecular mechanisms of host manipulation, as well as in the genome evolution of manipulative parasites. The evidence of HGT between multicellular eukaryotes remains scarce but is increasing and, therefore, elucidating its mechanisms will advance our understanding of the enduring influence of HGT on the evolution of the web of life.

Evolutionary genomics of camouflage innovation in the orchid mantis.

The orchid mantises achieve camouflage with morphological modifications in body color and pattern, providing an interesting model for understanding phenotypic innovation. However, a reference genome is lacking for the order Mantodea. To unveil the mechanisms of plant-mimicking body coloration and patterns, we performed de novo assembly of two chromosome-level genomes of the orchid mantis and its close relative, the dead leaf mantis. Comparative genomic analysis revealed that the Scarlet gene plays an important role in the synthesis of xanthommatin, an important pigment for mantis camouflage coloration. Combining developmental transcriptomic analysis and genetic engineering experiments, we found that the cuticle was an essential component of the 'petal-like' enlargement, and specific expression in the ventral femur was controlled by Wnt signaling. The prolonged expression of Ultrabithorax (Ubx) accompanied by femoral expansion suggested that Ubx determines leg remodeling in the early developmental stage. We also found evidence of evolution of the Trypsin gene family for insectivory adaptation and ecdysone-dependent sexual dimorphism in body size. Overall, our study presents new genome catalogs and reveals the genetic and evolutionary mechanisms underlying the unique camouflage of the praying mantis, providing evolutionary developmental insights into phenotypic innovation and adaptation.

Exploring the Mitogenomes of Mantodea: New Insights from Structural Diversity and Higher-Level Phylogenomic Analyses.

The recently reorganized classification of Mantodea has made significant progress in resolving past homoplasy problems, although some relationships among higher taxa remain uncertain. In the present study, we utilized newly sequenced mitogenomes and nuclear gene sequences of 23 mantid species, along with published data of 53 mantises, to perform familial-sampling structural comparisons of mantodean mitogenomes and phylogenomic studies. Our rstructural analysis revealed generally conserved mitogenome organizations, with a few cases of tRNA gene rearrangements, including the detection of trnL2 duplication for the first time. In our phylogenetic analysis, we found a high degree of compositional heterogeneity and lineage-specific evolutionary rates among mantodean mitogenomes, which frequently corresponded to several unexpected groupings in the topologies under site-homogeneous models. In contrast, the topologies obtained using the site-heterogeneous mixture model fit the currently accepted phylogeny of Mantodea better. Topology tests and four-cluster likelihood mapping analyses further determined the preferred topologies. Our phylogenetic results confirm the monophyly of superfamilial groups Schizomantodea, Amerimantodea, Heteromantodea, Promantidea, and Mantidea and recover the early-branching relationships as (Mantoidoidea + (Amerimantodea + (Metallyticoidea + Cernomantodea))). Additionally, the results suggest that the long-unresolved phylogenetic position of Majangidae should be placed within Mantidea, close to Mantoidea, rather than within Epaphroditoidea. Our findings contribute to understanding the compositional and structural diversity in mantodean mitogenomes, underscore the importance of evolutionary model selection in phylogenomic studies, and provide new insights into the high-level phylogeny of Mantodea.

On plate 24 by Maria Sibylla Merian in Metamorphosis insectorum surinamensium, and some types of Prioninae (Coleoptera: Cerambycidae).

The status of the two specimens of Prioninae seen on Merian's plate 24 (1705) is discussed, and considered as being Mallodon spinibarbis (Linnaeus, 1758). Prionus similis Schönherr (1817) is revalidated and placed in incertae sedis in Prioninae. Information on the types of Cerambyx spinibarbis Linnaeus, 1758, C. cinnamomeus Linnaeus, 1758, and C. melanopus Linnaeus, 1767 are provided.

Ultrastructure of the nebenkern during spermatogenesis in the praying mantid Hierodula membranacea.

Spermatogenesis leads to the formation of functional sperm cells. Here we have applied high-pressure freezing in combination with transmission electron microscopy (TEM) to study the ultrastructure of sperm development in subadult males of the praying mantid Hierodula membranacea, a species in which spermatogenesis had not previously been studied. We show the ultrastructure of different stages of sperm development in this species. Thorough examination of TEM data and electron tomographic reconstructions revealed interesting structural features of the nebenkern, an organelle composed of fused mitochondria that has been studied in spermatids of other insect species. We have applied serial-section electron tomography of the nebenkern to demonstrate in three dimensions (3D) that this organelle in H. membranacea is composed of two interwoven mitochondrial derivatives, and that the mitochondrial derivatives are connected by a zipper-like structure at opposing positions. Our approach will enable further ultrastructural analyses of the nebenkern in other organisms.

Novel gene rearrangements in mitochondrial genomes of four families of praying mantises (Insecta, Mantodea) and phylogenetic relationships of Mantodea.

The mitochondrial genome (mitogenome) plays an important role in phylogenetic studies of many species. The mitogenomes of many praying mantis groups have been well-studied, but mitogenomes of special mimic praying mantises, especially Acanthopoidea and Galinthiadoidea species, are still sorely lacking in the NCBI database. The present study analyzes five mitogenomes from four species of Acanthopoidea (Angela sp., Callibia diana, Coptopteryx sp., Raptrix fusca) and one of Galinthiadoidea (Galinthias amoena) that were sequenced by the primer-walking method. Among Angela sp. and Coptopteryx sp., three gene rearrangements were detected in ND3-A-R-N-S-E-F and COX1-L2-COX2 gene regions, two of which were novel. In addition, individual tandem repeats were found in control regions of four mitogenomes (Angela sp., C. diana, Coptopteryx sp., G. amoena). For those, plausible explanations were derived from the tandem duplication-random loss (TDRL) model and the slipped-strand mispairing model. One potential motif was found in Acanthopidae that was seen as a synapomorphy. Several conserved block sequences (CBSs) were detected within Acanthopoidea that paved the way for the design of specific primers. Via BI and ML analysis, based on four datasets (PCG12, PCG12R, PCG123, PCG123R), the merged phylogenetic tree within Mantodea was reconstructed. This showed that the monophyly of Acanthopoidea was supported and that the PCG12R dataset was the most suitable for reconstructing the phylogenetic tree within Mantodea.

Extrinsic and intrinsic musculature of the raptorial forelegs in Mantodea (Insecta) in the light of functionality and sexual dimorphism.

Prehensile raptorial forelegs are prey capturing and grasping devices, best known for praying mantises (Mantodea) within insects. They show strong morphological and behavioral adaptations toward a lifestyle as generalist arthropod predators. In the past, few species of Mantodea were investigated, concerning morphological variability of the raptorial forelegs. Especially the knowledge of foreleg anatomy in the light of functional and comparative morphology is scarce. Our comparative approach is based on the, for arthropods very common, "female-biased sexual size dimorphism" (SSD) that occurs in almost every Mantodea species. Within Mantodea, this SSD is likely leading to a shift of the exploited ecological niche between male and female individuals due to changes in, for example, the possible prey size; which might be reflected in the chosen ecomorphs. In this context, we analyzed the musculature of the raptorial forelegs of female and male specimens in five different species with varying SSD, using high-resolution microcomputed tomography and dissection. We were able to confirm the presence of 15 extrinsic and 15 intrinsic muscles-including one previously undescribed muscle present in all species. Thus, presenting a detailed description and illustrative three-dimensional anatomical visualization of the musculature in Mantodea. Interestingly, almost no observable differences were found, neither between species, nor between the sexes. Furthermore, we homologized all described muscles, due to their attachment points, to the comprehensive nomenclature established by Friedrich and Beutel (2008), discussed potential functionality of the muscles and possible homologies to the neuropteran Mantispa styriaca (Büsse et al., 2021) and the newly introduced leg nomenclature by Aibekova et al. (2022). By elucidating the anatomy, particularly in the context of functionality and SSD, our results complement previous knowledge of the raptorial forelegs, and facilitate a better understanding of the underlying biomechanical system of the predatory strike, and ultimately, a future comparison to other insect taxa.

Functional morphology of the praying mantis male genitalia (Insecta: Mantodea).

Male genitalia in praying mantids are highly complex, but we know little of how they function. I combined the micro-computed tomography of a copulating pair of the European mantis (Mantis religiosa) with public videos of copulation in various species of Mantodea and an analysis of literature. The function of each major element is reviewed. Copulation is divided into three phases: opening, anchoring and deposition. The opening is achieved by pulling the female subgenital plate with the male apical process. Multiple cases of female cooperation or resistance were observed and one case of coercion by the male. In species with the reduced apical process, female cooperation is mandatory. The male subgenital plate may participate in the opening as an integral part of the genitalia. After the opening, the conformation of the genitalia drastically changes, revealing activity of the genital papilla. Tight grasp on female genitalia is maintained solely by the clamp on the right phallomere, despite the overall complexity and predictions of sexual conflict theory. Other prominent elements show rhythmic motions, but their functions are not entirely clear and evidently involve spermatophore deposition, female stimulation or rival sperm removal. The opening and anchoring are similar in Mantodea and Blattodea, but achieved with non-homologous elements.

Three new species of longhorned beetles (Coleoptera: Cerambycidae) with notes on additional taxa.

The differences between Methia enigma Martins, 1981 and M. argentina Bruch, 1918 (Cerambycinae: Methiini) are discussed, and a new country record for the latter is provided. Sexual dimorphism in Pseudosparna Mermudes and Monné, 2009 (Lamiinae: Acanthocinini) is reported for the first time, and two new species of Pseudosparna (Lamiinae: Acanthocinini) are described from Panama: Pseudosparna dimitrisi sp. nov. and Pseudosparna marilenae sp. nov. The female of Pseudosparna mantis Devesa & Santos-Silva, 2020 is described for the first time. Eranina dariae sp. nov. (Lamiinae: Hemilophini) is described from French Guiana. The type locality of Bisaltes (Bisaltes) bimaculatus Aurivillius, 1904 (Lamiinae: Apomecynini) is reported, and a new country record is provided for the species. Aerenea subsulcicollis Breuning, 1948 (= A. sulcicollis subsulcicollis) (Lamiinae: Compsosomatini) is synonymized with A. sulcicollis Melzer, 1932, and a new country record is provided. A new state record is provided for Schmidarius kondratieffi Santos-Silva, Heffern, Botero & Nascimento, 2020 (Lamiinae: Hemilophini). Tucumaniella Breuning, 1943 (Apomecynini) is synonymized with Cacostola Fairmaire & Germain, 1859 (Onciderini), and Tucumaniella brevipes Breuning, 1943 is synonymized with Cacostola volvula (Fabricius, 1787).

Studies in Australian Mantodea: New synonyms, notes on distributions, and an updated checklist.

Despite recent advances in Mantodean systematics that have resolved many historical problems within the order, the Australian fauna still contains many erroneously recorded or incorrectly placed species. An investigation of these problematic species and related taxa necessitated a number of taxonomic changes: Exparoxypilus Beier is a new synonym of Paraoxypilus Saussure, Paraoxypilus verrauxii Wood-Mason and Exparoxypilus africanus Beier are new synonyms of Paraoxypilus verreauxii Saussure, Pthersigena Hinton is a new synonym of Phthersigena Stål, Austromantis alboterminata Sjöstedt is a new synonym of Austromantis albomarginata Sjöstedt, Deiphobe australiana Giglio-Tos is a new synonym of Coenomantis kraussiana Saussure, Notomantis Tindale is a new synonym of Pseudomantis Saussure, Hierodula dimorpha Werner is a new synonym of Pseudomantis albofimbriata (Stål), Notomantis chlorophana Tindale is a new synonym of Pseudomantis hartmeyeri Werner, Pseudomantis papua (Werner) comb. nov. is transferred from Hierodula, Orthodera marginata (Saussure) is reinstated as a valid species, and Ciulfina liturgusa Giglio-Tos and Orthodera gunnii Le Guillou are considered nomina dubia. In addition, notes are provided on the nomenclature, systematics, distribution, and ecology of several other Australian taxa, and a synonymic checklist of Australian Mantodea is provided. Finally, the origins and diversification of the Australian fauna are discussed in light of these and other recent changes.

The genus Holaptilon Beier, 1964: discussion on the poorly known boxer mantis genus, with a new species (Mantodea, Gonypetidae) .

The boxer mantis genus Holaptilon Beier, 1964 (Mantodea: Gonypetidae: Gonypetinae) is known to include two flightless species with a similar body size and morphology. Type species of the genus, H. pusillulum Beier, 1964, is reported from Israel and Jordan. An interesting species of the genus was recently identified from Iran (H. brevipugilis Kolnegari, 2018). In this research, a new species of H. yagmur Yilmaz & Sevgili sp. n. is described from Turkey. This new species was collected from Karacadag (Sanliurfa), a volcanic mountain in the Southeastern Anatolia region. Its typical habitat is steppic stony areas. It indicates that the species may be cryptic or nocturnal since it was collected from under stones and ground traps. This finding indicates that the genus is more widespread than previously known and may have more species yet to be discovered. The existence of the genus Holaptilon was reported for the first time in Turkey. Meanwhile, while reviewing the praying mantis list of Turkey, we determined that the samples collected from Agri, which are given in the Mantodea list of Turkey, were misdiagnosed as Armene robusta Mishchenko, 1956 and actually belong to the genus Holaptilon, and that Turkey currently hosts 20 praying mantis species/subspecies.

Scydmaenus Latreille of Australia: Revision of species in subgenera Choleropsis Franz, Kingius Franz, Mascarensia Franz, Parallomicrus Franz, Scottiscydmaenus Franz, and description of Ascydmaenus subgen. n. (Coleoptera, Staphylinidae, Scydmaeninae).

The Como National Park (CNP), located in the north-eastern part of the Ivory Coast, is strongly influenced by the river Como, which is eponymous for the park. Gallery forests along the river, forest islands, open grasslands, bush and tree savannas of various densities offer diverse habitats that harbor both forest and savannah praying mantis species. We present a preliminary checklist of praying mantids (Insecta: Mantodea) of the CNP. Specimens were collected in 2000 and 20172019, resulting in 35 species in 13 families. Two species, Ischnomantis werneri (Giglio-Tos 1916) and Solygia sulcatifrons (Audinet-Serville 1838), were recorded for the first time in Ivory Coast. Additional 47 species, which are expected to occur in the CNP, are listed here.