First assessment of the biodiversity of praying mantises (Insecta: Mantodea) in Cameroon with DNA barcoding.

Praying mantises are the apex insect predators in many ecosystems, nevertheless they receive relatively less recognition in biodiversity reviews. We report a first survey of diversity of praying mantises in Cameroon, which is situated in the Congo Basin region, one of the richest biodiversity hotspots. Combination of light trapping with manual collecting resulted in 495 specimens representing 62 species. A total of eight species are novel for the country, at least five species are likely undescribed. DNA barcodes of 72 specimens representing every collected species were obtained, curated, and submitted to NCBI database. For eight species, barcodes are published for the first time. A maximum likelihood phylogenetic tree was created using all available barcodes of Mantodea of Central African subregion. The results obtained during this study stress the importance of combining traditional and molecular approaches during biodiversity assessments of often neglected taxa, the latter aiding in uncovering new species, resolving unknown morphological divergencies and assigning conspecifics.

Taxonomic revision of the praying mantis subfamily Hierodulinae of China (Mantodea: Mantidae).

Species known from China in the praying mantis subfamily Hierodulinae are revised. A new species, Titanodula menglaensis sp. nov. is described. Hierodula tenuidentata Saussure, 1869 and Dracomantis mirofraternus Shcherbakov Vermeersch, 2020 are newly recorded from China. Two new synonyms are proposed: Titanodula formosana (Giglio-Tos, 1912) = Titanodula fruhstorferi (Werner, 1916), syn. nov. and Hierodula macrodentata Wang, Zhou Zhang, 2020 = Hierodula latipennis Brunner de Wattenwyl, 1893. Ootheca and male genitalia of the Chinese species are described and photographed. An identification key to genera and species of Hierodulinae from China is also provided. The current Chinese checklist contains 21 species.

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.

Mantodea (Insecta) of Uruguay: diversity and distribution.

Praying mantises (Order Mantodea) are charismatic insects thanks to their diverse and peculiar appearance and surprising behaviors. However, studies on this order are scarce, particularly in the Southern Neotropical Region. Here we present the first catalogue of praying mantises from Uruguay. We revised Mantodea specimens deposited at Colección de Entomología, Facultad de Ciencias (FCE-MN), Universidad de la República, Montevideo, Uruguay. We recorded information about distribution and notes on the biology of the species. Nineteen species are reported for Uruguay, including new family, genera or species for the country. We report the presence of six taxa for first time: the family Mantoididae, genus Pseudovates, Musoniella and Metaphotina, the species Mantoida beieri, Eumusonia livida, Musoniella argentina, Metaphotina brevipennis, Parastagmatoptera theresopolitana and Pseudovates iheringi. We could confirm four species previously reported for the country, but eight species are based on dubious records or species and we cannot confirm their current occurrence in this work. More studies are needed in the country for further investigations of Mantodea in the country and the region.

Mantidis Oötheca (mantis egg case) original species identification via morphological analysis and DNA barcoding.

ETHNOPHARMACOLOGICAL RELEVANCE: Mantidis Oötheca (mantis egg case; sangpiaoxiao) is a medicine from an insect source, which has been widely used in Asian countries. However, misidentification due to a lack of information given variations in the medicinal portion of the ootheca and morphological similarities of the ootheca as an egg chamber. AIM OF THE STUDY: Thus, this study aims to provide the first comprehensive data for discriminating authentic of Mantidis Oötheca. Here, we provide detailed ootheca morphology and their molecular information to accurately identify Mantidis Oötheca. MATERIALS AND METHODS: Oothecae of Tenodera angustipennis (Saussure, 1869), Tenodera sinensis (Saussure, 1871), Hierodula patellifera Serville, 1839, and Hierodula sp. were used in the comparative morphological, principal component analysis, and DNA barcoding. RESULTS: The morphological analyses revealed that the emergence area, outline, angle of distal end, width of air-filled layer, and weight are useful diagnostic characters. Using these quantitative and qualitative characteristics, we developed the effective identification key. Furthermore, our CO1 sequences from all individuals were monophyletic with high bootstrap values at genus and species levels. Moreover, morphological identification using our developed key among all studied individuals agreed with molecular identification results using CO1 barcoding data. CONCLUSIONS: These multilateral approaches, including morphological, statistical, and DNA barcoding methods are highly reliable identification tools. Moreover, our diagnostic key characteristics and molecular barcoding should aid in the accurate identification, authentication, and quality control of Mantidis Oötheca medicinal materials.

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.

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.

A Cretaceous-aged Palaeotropical dispersal established an endemic lineage of Caribbean praying mantises.

Recent phylogenetic advances have uncovered remarkable biogeographic histories that have challenged traditional concepts of dispersal, vicariance and diversification in the Greater Antilles. Much of this focus has centred on vertebrate lineages despite the high diversity and endemism of terrestrial arthropods, which account for 2.5 times the generic endemism of all Antillean plants and non-marine vertebrates combined. In this study, we focus on three Antillean endemic praying mantis genera, Callimantis, Epaphrodita and Gonatista, to determine their phylogenetic placement and geographical origins. Each genus is enigmatic in their relation to other praying mantises due to their morphological affinities with both Neotropical and Old World groups. We recovered the three genera as a monophyletic lineage among Old World groups, which was supported by molecular and morphological evidence. With a divergence at approximately 107 Ma, the lineage originated during the break-up of Gondwana. Ancestral range reconstruction indicates the lineage dispersed from an African + Indomalayan range to the Greater Antilles, with a subsequent extinction in the Old World. The profound ecomorphic convergence with non-Caribbean groups obscured recognition of natural relationships within the same geographical distribution. To the best of our knowledge, the lineage is one of the oldest endemic animal groups in the Greater Antilles and their morphological diversity and restricted distribution mark them as a critical taxon to conserve.

Comparative Mitogenomic Analyses of Praying Mantises (Dictyoptera, Mantodea): Origin and Evolution of Unusual Intergenic Gaps.

Praying mantises are a diverse group of predatory insects. Although some Mantodea mitogenomes have been reported, a comprehensive comparative and evolutionary genomic study is lacking for this group. In the present study, four new mitogenomes were sequenced, annotated, and compared to the previously published mitogenomes of other Mantodea species. Most Mantodea mitogenomes share a typical set of mitochondrial genes and a putative control region (CR). Additionally, and most intriguingly, another large non-coding region (LNC) was detected between trnM and ND2 in all six Paramantini mitogenomes examined. The main section in this common region of Paramantini may have initially originated from the corresponding control region for each species, whereas sequence differences between the LNCs and CRs and phylogenetic analyses indicate that LNC and CR are largely independently evolving. Namely, the LNC (the duplicated CR) may have subsequently degenerated during evolution. Furthermore, evidence suggests that special intergenic gaps have been introduced in some species through gene rearrangement and duplication. These gaps are actually the original abutting sequences of migrated or duplicated genes. Some gaps (G5 and G6) are homologous to the 5' and 3' surrounding regions of the duplicated gene in the original gene order, and another specific gap (G7) has tandem repeats. We analysed the phylogenetic relationships of fifteen Mantodea species using 37 concatenated mitochondrial genes and detected several synapomorphies unique to species in some clades.

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).

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.

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.

Reconstruction of two colonisation pathways of Mantis religiosa (Mantodea) in Germany using four mitochondrial markers.

Past and recent climatic changes induced shifts in species ranges. Mantis religiosa has also expanded its range across Germany within the past decades. To determine the ancestry of German M. religiosa we sequenced four mitochondrial genes (COI, COII, Cyt b, ND4) of European M. religiosa populations. We found an east, central and west European lineage of M. religiosa. These distinct lineages are consistent with genetic isolation by distance during glacial periods, and the re-colonization of northern parts of Europe by species from different refugia. Within Germany, we found haplotypes clustering to the central and west European lineage suggesting that M. religiosa immigrated from two directions into Germany. Mismatch distributions, and negative Tajima's D and Fu's Fs values indicate a current range expansion of the central and west European lineage. We hypothesise that ongoing global warming which increases the availability of thermally favourable areas in Germany for M. religiosa adds to its current range expansion. In conclusion, M. religiosa colonized Germany via two directions: west German populations descended from French populations and east German populations from Czech populations.

[Identification of original species of Mantidis Oötheca (Sangpiaoxiao) based on DNA barcoding].

Both market research and literature reports both found that the ootheca of mantodea was all used as medicine. However, Chinese Pharmacopoeia only records the ootheca of three mantis species. The clinical use of ootheca unrecorded in Chinese Pharmacopoeia, will pose potential risks to drug safety. It's urgent to identify the origin of Mantidis Oötheca. The current researches about original animal in Mantidis Oötheca are based on morphology and unanimous. DNA barcoding fill gaps of the traditional morphological identification, which is widely used in animal classification studies. This study first use DNA barcoding to analyze genetic distance among different Mantidis Oötheca types, align COI sequences between mantis and Mantidis Oötheca and construct the phylogeny tree. The result confirmed that Tenodera sinensis and Hierodula patellifera were the origin insects of Tuanpiaoxiao and Heipiaoxiao, respectively, and Statilia maculate and Mantis religiosa were the origin insects of Changpiaoxiao.