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A New Species of Praying Mantis from Peru Reveals Impaling as a Novel Hunting Strategy in Mantodea (Thespidae: Thespini)
Abstract
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.
Supplementary resource (1)
... This family comprises 32 genera (Rivera & Svenson 2016), but few have been studied in revisionary works (e.g., Rivera et al. 2011;Agudelo & Rafael 2014;Scherrer 2014;Maldaner et al. 2015). This family comprises species with cryptic coloration, usually dull-colored or yellowish (Rivera 2010;Rivera & Callohuari 2019). All the females are apterous, and they resemble nymphs at first glance. ...
... obs.). Similar behavior was recently recorded by Rivera & Callohuari (2019) in a species of Carrikerella. ...
... They are generalist predators and caged adults feed on all kinds of small and softbodied prey, such as flies, moths, cockroaches, crickets, and grasshoppers. Adults and more advanced nymphs capture prey utilizing the usual sweep hunting strategy (see Rivera & Callohuari 2019) with the spines of fore femur and tibia. On the other hand, tiny and fragile caged first instar nymphs were observed to utilize only the dorsal spine of the fore tibia to capture prey. ...
The identification of species in many genera of Neotropical Mantodea is a difficult task because of a lack of revisionary works with keys and illustrations of male genitalia. One of these genera is Pseudomiopteryx Saussure, 1870, which is known to be composed of 10 species and one subspecies of small (19–25mm) brownish praying mantises that live in the leaf litter of tropical forests of Central America, the Amazon, and the Andean tropics. The genus is revised and the known valid species are redescribed based mainly on type-material. The results yield 11 valid species, including a new species from Costa Rica: Pseudomiopteryx riverai sp. n. Pseudomiopteryx amazonensis Toledo-Piza, 1968 is a junior synonym of Pseudomiopteryx guyanensis Chopard, 1911, and the subspecies Pseudomiopteryx bogotensis gorgonae Ariza & Salazar, 2005 was raised to species level. A syntype specimen of Pseudomiopteryx decipiens, Pseudomiopteryx guyanensis, and Pseudomiopteryx infuscata Saussure & Zehntner, 1894 was herein designated as the lectotype. New information on geographical distributions and natural history were given, and descriptions of the female, ootheca, and first instar nymph of Pseudomiopteryx spinifrons were presented. The geographical distributions of some species are extended. A key to males of all species is given, as well as detailed figures of male genitalia.
... 2), Blepharopsis mendica (Fabricius, 1775) (Korsakofi 1949, board II, figs 1 and 2, and board IV, fig. 1), Callibia diana (Avendaño & Sarmiento 2011), Cardioptera viridipennis (Heitzmann-Fontenelle 1969), Thesprotiella simpira (Rivera & Callohuari, 2020) (Rivera & Callohuari 2020, fig. 6g and i), Didymocorypha lanceolata (Fabricius, 1798) (Wieland 2013, p. 163, figs 428-431), Empusa pennata (Bugnion 1923, figs 22, 28 and 29), Helvia cardinalis Stål, 1877 (Heynderycx 2000), Hierodula patellifera (Oshima 2021(Oshima , 2022a, Hierodula sp. ...
... 2), Blepharopsis mendica (Fabricius, 1775) (Korsakofi 1949, board II, figs 1 and 2, and board IV, fig. 1), Callibia diana (Avendaño & Sarmiento 2011), Cardioptera viridipennis (Heitzmann-Fontenelle 1969), Thesprotiella simpira (Rivera & Callohuari, 2020) (Rivera & Callohuari 2020, fig. 6g and i), Didymocorypha lanceolata (Fabricius, 1798) (Wieland 2013, p. 163, figs 428-431), Empusa pennata (Bugnion 1923, figs 22, 28 and 29), Helvia cardinalis Stål, 1877 (Heynderycx 2000), Hierodula patellifera (Oshima 2021(Oshima , 2022a, Hierodula sp. ...
There is a great knowledge gap regarding the morphology of immatures of Mantodea, making taxonomic identification of the nymphs difficult, therefore hindering their inclusion in other studies. The present work recognizes general ontogenetic patterns shared by praying mantises through the comparative analysis of the post-embryonic changes in the external morphology of the nymphal instars of 10 Neotropical species, representing nine genera in five families. Nymphs were reared in the laboratory, their exuviae were collected and vouchers fixed. A total of 369 specimens and 495 exuviae were studied, and a total of 7000 images of live nymphs, at each instar, were taken. Shared developmental patterns were recognized for the form of the spines of the forelegs, proximal discoidal spine of the forefemora, wing pads, female genitalia, cerci, body adornments such as denticles, lobes and apophyses, and crypsis. Based on the recognition of a set of key differences in these features, two distinct, successive forms are recognized for mantis nymphs: the protonymph, the form of the first instar, and the deuteronymph, the form of the remaining instars. In turn, two successive forms are identified for the deuteronymphs: the mesonymph, the form of the early deuteronymphs, and the metanymph, the form of the late deuteronymphs. Literature data provide good support for the occurrence of the observed patterns also in many other and highly diverse lineages of Mantodea, suggesting this might be a generalized feature, fixed in the ontogenetic system of the order. If so, these nymphal forms can be considered semaphoronts of Mantodea, which are more practical and more relevant references than instars for comparisons between praying mantis nymphs. An original key to recognizing the different postembryonic semaphoronts and sexes of Mantodea is provided. The discovery has wide application in phylogenetic and taxonomic studies, providing an important framework for the study of nymphs of Mantodea. It is recommended that taxonomic studies provide descriptions for proto-, meso- and metanymphs, and include their features in identification keys.
... We investigated 437 Mantodea specimens, and found them to represent 44 species in 28 genera. The following list contains nine new Peruvian records compared to the most recent checklist (Rivera & Vergara-Cobián (2017), with additions by Rivera & Callohuari (2019) and Svenson & Rodrigues (2019)) and one species new to science. (Fig. 12a, b), while external and other genital characters are rather similar across specimens. ...
... ). Recently,Rivera & Callohuari (2019) andSvenson & Rodrigues (2019) added additional taxa to the list. FIGURES 130-135. ...
We present here the first illustrated checklist of the praying mantids (Mantodea) collected at the Panguana Field Station in Central Peru over the course of 50 years. The examination of over 430 specimens obtained mainly by light-trapping, but also other methods, revealed 44 species in 28 genera. Mantoida brunneriana, Mantoida cf. argentinae, Pseudomiopteryx cf. decipiens, Angela trifasciata, Liturgusa neblina, Cardioptera squalodon, Metriomantis cf. pilosella, Acontista festae, and Heterovates pardalina are new Peruvian records. Microphotina panguanensis n. sp. is new to science and the first species of the genus Microphotina described from the Western Amazon. The checklist of confirmed Peruvian Mantodea species is raised to 80. Thus, more than half of the currently known Peruvian Mantodea species is found at Panguana. We discuss the reasons for this diversity and comment on putative additional species which might be sampled if collection efforts are intensified. The results highlight the Conservation value of ACP Panguana for Western Amazonian Mantodea.
... An increase in the length of the coxa and femur has been previously related to an increase in strike reachability and faster movements (Oufiero et al., 2016), which could be potentially related to a specific hunting strategy related to environmental mimicry. Multiple species also have morphologically unique forelegs (e.g., those of Amorphoscelis, Idolomantis, Toxodera; see Fig. 7), which often indicate different predatory specializations (e.g., Rivera & Callohuari, 2020), which could have been key to the diversification of Mantidea. ...
Mantodea (praying mantises) is a group of exclusively predatory insects, which, together with nonraptorial blattodeans (cockroaches and termites) and groups exclusively found in the fossil record, form the group Dictyoptera. A central characteristic of Mantodea is the specialization of their first pair of legs as raptorial grasping appendages, but the evolution from walking to raptorial legs is not yet fully understood. Here, we trace the evolution of the raptorial appendages in Dictyoptera through time using a morphometric (morphospaces) approach. We also describe two new mantodean nymphs preserved in amber from the Cretaceous and Eocene, which expand the scarce mantodean fossil record. Blattodean and mantodean appendages appear distinct in morphospace, but several appendages of fossil non‐mantodeans can be considered raptorial, providing a potential transitional link between walking and raptorial morphotypes. Therefore, we discuss potential mantodean affinities for other predatory fossil dictyopterans. We examine changes across extant mantodeans, characterized by a straightening of the tibia especially associated with the rise of the diversification of the Mantidea and discuss whether a thickening of the femur could reflect an early adaptation to cursorial hunting.
... Mantises are well known to catch small birds, especially hummingbirds, as they fly midair (Fisher 1994;Hildebrand 1949). They are the only known insect to kill their prey with their legs (Rivera and Callohuari 2019). There is only a record of Hierodula patellifera (Serville), predating on crimsonbacked sunbird from India (Browne 1899). ...
With more than one million named species, insects are the most diverse terrestrial creatures on the planet, representing around 75% of the global fauna. They contribute to invaluable ecosystem functions such as nutrient cycling, pollination, and seed dispersals and serve as a significant food source, aid in biocontrol of other organisms (such as predators, parasites), and maintain soil structure and fertility. Meantime, they are profoundly beneficial as pollinators. They often compete with humans, as pests of agriculture and stored products and as vectors of life-threatening diseases. Owning four of the globally recognised biodiversity hotspots—Himalaya, Indo Burma, Western Ghats and Sri Lanka, and Sundaland—India is represented by 67,111 insect species in four classes of subphylum Hexapoda: Collembola (345 species), Protura (20 species), Diplura (18 species), and Insecta (66,728 species), 64.8% of the overall faunal diversity of the country. Eight insect orders—Coleoptera, Lepidoptera, Hymenoptera, Diptera, Hemiptera, Orthoptera, Thysanoptera, and Odonata—form the majority (94%) of the insects in the country. This chapter further updates India’s known insect diversity, emphasising its diversity in ecosystems (such as aquatic, mangroves, soil, and forests) and biogeographic zones (Himalaya, Trans-Himalaya, Desert, and Islands). Additionally, information has been provided on the potential of insects in food security, pollination, forest pests, and their significance to medical and veterinary in context with Indian fauna. The chapter also includes a list of 27 insect species identified as invasive aliens in India and reports 22 species as threatened in the IUCN Red List from India.
Keywords: Ecosystem diversity, Pollination, Food security, Threatened insects
... Because of a multiplicity of possible morphological interpretations for leg armament, we outline literature for the five primary functions: attachment, sensation, grooming, communication and object manipulation. Beginning with the last category, it is well known that legs and leg armament can be used for prey capture in insects through clasping between two or more segments (Oufiero, 2019;Olmi et al., 2020), the increase in friction or adhesion via arrays of various hair-like structures (Zhang & Weirauch, 2013), impalement (Rivera & Callohuari, 2020) or even 'stomping' (Betz & Mumm, 2001) among other mechanisms. Notably, tarsal setation, in opposition with the tarsal annuli, has been suggested to be used for grasping in ants (Cassil et al., 2006). ...
Miniaturization strongly affects functional morphology. Whereas some anatomical structures are barely affected by scaling, others can fundamentally change as the body becomes ever smaller. No prior study has focused on the effect of miniaturization on grooming and attachment structures in Hymenoptera, which can be highly diverse and complex. Through comparative description of the legs of the extremely small wasps of the families Mymaridae and Trichogrammatidae, we evaluate the functional and phylogenetic patterns concerning possible functional effects of miniaturization. On the one hand, the studied species retain some features characteristic of other Chalcidoidea, while on the other, they display some parallelisms associated with miniaturization in leg structure. These observations support a two‐stage morphocline of miniaturization, wherein the first stage is characterized by the preservation of structural complexity and retention of all basic functions, as for instance in examined Megaphragma and the females of Dicopomorpha. The second stage is characterized by a significant simplification, with the loss of redundant non‐essential functions, as observed for the males of Dicopomorpha, which have grossly reduced leg structures, including total loss cleaning devices. Whether these stages are ordered or unordered should be evaluated in future study. Functional optimization of attachment in male Dicopomorpha is indicated by the highly derived mushroom‐shaped tarsi, complemented by novel grappling spurs on the hindfeet, possibly for copulation. Our observations underline adaptive trade‐offs in the expression of complex and multifunctional leg structures at extreme scales.
... Praying mantises as vertebrate predators were reported more than would be expected and were mostly observed preying on birds. Mantises not only eat fledglings but are also well known to capture small birds, especially hummingbirds as they hover in mid-air (Fisher, 1994;Hildebrand, 1949;Lorenz, 2007;Murray, 1958), and are the only known insect to impale their prey with its legs (Rivera & Callohuari, 2019). Malacostraca were also documented more than expected and were documented mainly from Brazil and the United States, where they were underrepresented, | 9 VALDEZ with a bias towards Central America and the Caribbean. ...
Aim
Arthropods as vertebrate predators is a generally overlooked aspect in ecology due to the cryptic nature of these events, the relatively small size of arthropods and the difficulty in finding published data. This study represents the largest global assessment of arthropods preying on vertebrates to provide a conceptual framework, identify global patterns and provide a searchable database.
Location
Global.
Time period
Present.
Major taxa studied
Arthropods and vertebrates.
Methods
A systematic literature review was conducted.
Results
Over 1,300 recorded observations were collated from 89 countries. Arthropod predators were from 6 classes and 83 families. Vertebrate prey were from 5 classes and 163 families. Spiders represented over half of all predatory events and were the main predator for all vertebrates except birds, which were mostly preyed upon by praying mantises. Forty percent of all prey were amphibians, specifically frogs. Depredated reptiles were nearly all lizards, half of mammal prey were bats, nearly a third of fish were Cypriniformes and half of bird prey were passerines. Predation by spiders was mainly documented from the U.S., Brazil and Australia, and biased mostly everywhere except the U.S.; insect predatory events were mainly documented from Europe, Australia and the Americas, and biased toward North America; amphibian events were mainly from the Americas and strongly biased everywhere, except for the U.S. and Australia; reptiles events were recorded mostly from the Americas and Australia, and biased towards the U.S. and Australia; predation on birds were mainly from the Americas, Australia and Europe, and biased towards Central America and Europe; and mammal events were mostly reported from North and Central America, Australia, and Asia, and strongly biased everywhere except Brazil.
Main conclusions
This study demonstrates that arthropods are underestimated predators of vertebrates. Recognizing and quantifying these predator–prey interactions is vital for identifying patterns and the potential impact of these relationships on shaping vertebrate populations and communities.
... Praying mantises as vertebrate predators were reported more than would be expected and were mostly observed preying on birds. Mantises not only eat fledglings but are also well known to capture small birds, especially hummingbirds as they hover in mid-air (Fisher, 1994;Hildebrand, 1949;Lorenz, 2007;Murray, 1958), and are the only known insect to impale their prey with its legs (Rivera & Callohuari, 2019). Malacostraca were also documented more than expected and were documented mainly from Brazil and the United States, where they were underrepresented, | 9 VALDEZ with a bias towards Central America and the Caribbean. ...
Preprint is now published in Global Ecology and Biogeography. Updated and published version available here:
https://www.researchgate.net/publication/343258316_Arthropods_as_vertebrate_predators_A_review_of_global_patterns
A recent reinvigorated effort into examining morphology and ecology of Nearctic Mantodea has led to the description of nearly 10 new species in the past decade. Expeditions to southern Arizona, combined with iNaturalist observations, suggested that S. wheelerii sensu Anderson is a complex of at least two cryptic species across its extensive range. Herein I describe one new species: Stagmomantis clauseni sp. nov., the Sonoran Tiger Mantis, from the Sonoran and Chihuahuan deserts, utilizing citizen science contributions, comparative morphology, and captive observations to describe its ontogeny and ethology. In addition, I also redescribe Stagmomantis wheelerii (Thomas 1875) and document the holotypes of Stagmomantis colorata (Hebard 1922) for the first time. The development of Stagmomantis clauseni sp. nov. also warranted the description of new hatching and signaling systems in Mantodea. For ease of both citizen and professional scientists a dichotomous key is provided for members of the subgenus Nigralora including characters in life.
This study presents a new nature-inspired optimization algorithm, namely the Mantis Search Algorithm (MSA), inspired by the unique hunting behavior and sexual cannibalism of praying mantises. In brief, MSA consists of three optimization stages, including the search for prey (exploration), attack prey (exploitation), and sexual cannibalism. Those operators are simulated using various mathematical models to effectively tackle optimization challenges across diverse search spaces. The performance of MSA is rigorously tested on fifty-two optimization problems and three real-world applications (five engineering design problems, and the parameter estimation problem of photovoltaic modules and fuel cells) to show its versatility and adaptability to different scenarios. To disclose the MSA’s superiority, it is compared to two categories from the rival optimizers: the first category involves well-established and highly-cited optimizers, like Differential evolution; and the second category contains recently-published algorithms, like African Vultures Optimization Algorithm. This comparison is conducted using several performance metrics, the Wilcoxon rank-sum test and the Friedman mean rank to disclose the MSA’s effectiveness and efficiency. The results of this comparison highlight the effectiveness of this new approach and its potential for future optimization applications.
Praying mantids (class Insecta, order Mantodea) are a group of predatory insects comprising approximately 2500 described species, that occur across all continents except Antarctica, with the greatest species diversity in tropical and subtropical regions. Mantids predominantly prey on other invertebrates but are known to feed on small vertebrates. During April and May 2021, we observed mantid feeding events in Manujan County, Kerman Province in southern Iran. Two distinct feeding events were observed where female European Mantids (Mantis religiosa) preyed on Purple Sunbird (Cinnyris asiaticus) and Crested Lark (Galerida cristata) nestlings. In addition, we collated information from online searches of mantids feeding on nestlings elsewhere in the world, revealing two more observations. In Taiwan, a Giant Asian Mantid (Hierodula patellifera) was recorded preying on a nestling Warbling White-eye (Zosterops japonicus) and in Brazil, a mantid (Stagmatoptera sp.) was recorded feeding on a nestling White-throated Seedeater (Sporophila albogularis). To date, the only existing scientific evidence of praying mantids feeding on passerine nestlings was recorded in 1922. We propose two potential explanations for the observed trophic interactions between mantids and passerine nestlings: (1) during egg production female mantids, especially those in poor physical condition, may opportunistically feed on nestlings in order to increase fecundity via nutrient gain and (2) mantids may initially be attracted toward the nest by parasitic or coprophagous insects, as a result of poor nest sanitation, and subsequently prey on nestlings after detecting movements. Our unusual observations represent the first records of praying mantids feeding on nestling passerines in nearly 100 years.
The extremely derived morphology and behaviour of extant praying mantises combined with a scarce record of fossil relatives introduce significant challenges to tracing their evolution from Palaeozoic stem‐dictyopterans. Extant members of Chaeteessidae, Mantoididae and Metallyticidae could be invaluable to resolving the mantodean tree, yet their inclusion in phylogenetic analyses led to conflicting hypotheses due to their highly disparate respective morphologies. In this contribution, we present Labradormantis guilbaulti gen. et sp.n., a new fossil species described from both fore‐ and hind‐wing imprints discovered in the Redmond Mine locality (Late Cretaceous, Cenomanian, Redmond Formation; Labrador, Canada). The examination of its hind‐wing AA2* supports the hypothesis that this structure, unique to Chaeteessidae among extant mantises, is a true vein and that its occurrence represents a plesiomorphy for Mantodea. A parsimony analysis including newly coded wing‐related characters further established that L. guilbaulti gen. et sp.n. displays a unique combination of plesiomorphic and apomorphic character states that situates it within the extinct family Baissomantidae. This dataset resolved the phylogenetic relationships of early‐diverging extant lineages as (Chaeteessidae (Mantoididae (Metallyticidae, Artimantodea))), and suggested that the Eocene Lithophotina floccosa Cockerell might be a close relative of extant metallyticids. It also indicated a trend towards increased modularity within mantis fore‐wings, in contrast with a trend towards increased morphological integration in their hind‐wings, both of which are potentially associated with improved flight performance for modern mantises. This study emphasizes the importance of fossils for resolving phylogenetic relationships and for introducing transitional phenotypes to infer ancient evolutionary trends of extant derived clades.
Popa spurca
(Stål) (Deroplatyidae) is a well-known species within the African Mantodea. Because this species is often kept as pets, several citizen-based platforms provide information on the rearing of this species. There is however no scientific information available on the distribution and biology of this species. The aim of this study was to use museum records to establish the historic distribution of P. spurca in southern Africa and specifically South Africa, and to investigate its biology under captive breeding and rearing conditions. A total of 212 specimen records of P. spurca were recorded during a survey of all Mantodea in the national museum collections in South Africa. The results suggested that the distribution of this genus seems to be largely associated with the Savanna biome in South Africa. The average lifespan of a P. spurca individual was 332 ± 62 days adult longevity was significantly longer in females. Oothecae contained an average of 84 ± 30 eggs and the mean incubation period was 35 ± 4 days. This study contributes to the understanding of the historic distribution and biology of P. spurca which has never been studied before in southern Africa.
Observations in unmanipulated, semi-natural conditions were made of a single individual of the praying mantid Hierodulatenuidentata , while hunting and capturing an unusual prey for this kind of insect, guppy fish, Poeciliareticulata . This repetitive fishing behavior, recorded daily, is reported here for the first time and discussed in relation to the adaptive behavioral plasticity of praying mantids. We speculate regarding learning from experience as a hunting strategy.
Digitization of natural history collections is a major challenge in archiving biodiversity. In recent years, several approaches have emerged, allowing either automated digitization, extended depth of field (EDOF) or multi-view imaging of insects. Here, we present DISC3D: a new digitization device for pinned insects and other small objects that combines all these aspects. A PC and a microcontroller board control the device. It features a sample holder on a motorized two-axis gimbal, allowing the specimens to be imaged from virtually any view. Ambient, mostly reflection-free illumination is ascertained by two LED-stripes circularly installed in two hemispherical white-coated domes (front-light and back-light). The device is equipped with an industrial camera and a compact macro lens, mounted on a motorized macro rail. EDOF images are calculated from an image stack using a novel calibrated scaling algorithm that meets the requirements of the pinhole camera model (a unique central perspective). The images can be used to generate a calibrated and real color texturized 3Dmodel by ‘structure from motion’ with a visibility consistent mesh generation. Such models are ideal for obtaining morphometric measurement data in 1D, 2D and 3D, thereby opening new opportunities for trait-based research in taxonomy, phylogeny, eco-physiology, and functional ecology.
Th e present article aims to describe a new species of lichen-mantis: Carrikerella amazonica sp. nov., based on female specimens captured in various Amazonian locations of Colombia. Scope: is the third species known of the genus currently. Methodology: all specimens were captured manually in their typical habitat on tree trunks. Results: notes are included regarding the Carrikerella genus and its species, as well as its natural habitat and behavior. Conclusions: It is concluded that their species inhabit the tropical rain forest of the lowlands.
The Atlantic Forest is one of the world’s top biodiversity hotspots, but the diversification processes of its biota are still poorly known, with competing models attributing dominant roles to either Quaternary climatic changes or geographic barriers. Many studies identify the Doce river as a major phylogeographic break, but the reasons for this phenomenon are highly debated. Here we test the predictions of the refugial and barrier models for a common species of praying mantis, Miobantia fuscata, focusing in the areas immediately south and north of the Doce river. Our analyses show high intraspecific genetic diversity, deep coalescence times and no evidence for recent population expansion. Phylogeographic structure is inconsistent with a refugial hypothesis. Significant gene flow between northern and southern populations also conflicts with a strong role for geographic barriers. This study highlights the need for considering invertebrate taxa to infer recent landscape changes, and points towards a more complex picture of genetic diversification in the Atlantic Forest.
The present article aims to describe a new species of lichen-mantis: Carrikerella amazonica sp. nov., based on female specimens captured in various Amazonian locations of Colombia. Scope: is the third species known of the genus currently. Methodology: all specimens were captured manually in their typical habitat on tree trunks. Results: notes are included regarding the Carrikerella genus and its species, as well as its natural habitat and behavior. Conclusions: It is concluded that their species inhabit the tropical rain forest of the lowlands.
We present a preliminary checklist of the praying mantises (Insecta: Mantodea) of Peru. A total of 67 species are reported, nine of which constitute new records from that country. The following new nomenclatural procedures are introduced: Musoniella margharethae Battiston and Picciau, 2008 is transferred to Musonia as Musonia margharethae (Battiston and Picciau, 2008) (comb. n.), and Phyllovates brevicollis Orofino, Ippolito and Lombardo, 2006 is considered a new synonym (syn. n.) of Pseudovates peruviana (Rehn, 1911). A new genus, Piscomantis gen. n., is established to accommodate Galapagia peruana Beier, 1935, which is now referred to as Piscomantis peruana (Beier, 1935) (comb. n.). Ten species are removed from the Peruvian mantis fauna and two nomina nuda are proposed. An overview of the biogeography and ecology of Peruvian praying mantises is presented.
The crustacean order Stomatopoda comprises seven superfamilies of mantis shrimps, found in coastal waters of the tropics and subtropics. These marine carnivores bear notable raptorial appendages for smashing or spearing prey. We investigated the evolutionary relationships among stomatopods using phylogenetic analyses of three mitochondrial and two nuclear markers. Our analyses recovered the superfamily Gonodactyloidea as polyphyletic, with Hemisquilla as the sister group to all other extant stomatopods. A relaxed molecular clock, calibrated by seven fossil-based age constraints, was used to date the origin and major diversification events of stomatopods. Our estimates suggest that crown-group stomatopods (Unipeltata) diverged from their closest crustacean relatives about 340 Ma (95% CRI [401–313 Ma]). We found that the specialized smashing appendage arose after the spearing appendage ∼126 Ma (95% CRI [174–87 Ma]). Ancestral state reconstructions revealed that the most recent common ancestor of extant stomatopods had eyes with six midband rows of hexagonal ommatidia. Hexagonal ommatidia are interpreted as plesiomorphic in stomatopods, and this is consistent with the malacostracan ground-plan. Our study provides insight into the evolutionary timescale and systematics of Stomatopoda, although further work is required to resolve with confidence the phylogenetic relationships among its superfamilies.
This study provides a comprehensive review of historical morphological nomenclature used for praying mantis (Mantodea) morphology, which includes citations, original use, and assignment of homology. All referenced structures across historical works correspond to a proposed standard term for use in all subsequent works pertaining to praying mantis morphology and systematics. The new standards are presented with a verbal description in a glossary as well as indicated on illustrations and images. In the vast majority of cases, originally used terms were adopted as the new standard. In addition, historical morphological topographical homology conjectures are considered with discussion on modern interpretations. A new standardized formulation to present foreleg femoral and tibial spines is proposed for clarity based on previous works. In addition, descriptions for methods of collection, curation, genital complex dissection, and labeling are provided to aid in the proper preservation and storage of specimens for longevity and ease of study. Due to the lack of consistent linear morphometric measurement practices in the literature, we have proposed a series of measurements for taxonomic and morphological research. These measurements are presented with figures to provide visual aids with homologous landmarks to ensure compatibility and comparability across the Order. Finally, our proposed method of pinning mantises is presented with a photographical example as well as a video tutorial available at http://mantodearesearch.com.
We review 147 incidents of the capture of small birds by mantids (order Mantodea, family Mantidae). This has been documented in 13 different countries, on all continents except Antarctica. We found records of predation on birds by 12 mantid species (in the genera Coptopteryx, Hierodula, Mantis, Miomantis, Polyspilota, Sphodromantis, Stagmatoptera, Stagmomantis, and Tenodera). Small birds in the orders Apodiformes and Passeriformes, representing 24 identified species from 14 families (Acanthizidae, Acrocephalidae, Certhiidae, Estrildidae, Maluridae, Meliphagidae, Muscicapidae, Nectariniidae, Parulidae, Phylloscopidae, Scotocercidae, Trochilidae, Tyrannidae, and Vireonidae), were found as prey. Most reports (.70% of observed incidents) are from the USA, where mantids have often been seen capturing hummingbirds attracted to food sources in gardens, i.e., hummingbird feeders or hummingbird-pollinated plants. The Ruby-throated Hummingbird (Archilochus colubris) was the species most frequently reported to be captured by mantids. Captures were reported also from Canada, Central America, and South America. In Africa, Asia, Australia, and Europe, we found 29 records of small passerine birds captured by mantids. Of the birds captured, 78% were killed and eaten by the mantids, 2% succeeded in escaping on their own, and 18% were freed by humans. In North America, native and non-native mantids were engaged in bird predation. Our compilation suggests that praying mantises frequently prey on hummingbirds in gardens in North America; therefore, we suggest caution in use of large-sized mantids, particularly non-native mantids, in gardens for insect pest control.
We present the description of a new genus and new species of praying mantis, Hondurantemna chespiritoi
gen. n. & sp. n. This species of cryptic mantis, collected in National parks in Mexico and Honduras,
remained unknown despite its considerable body size. Based on a phylogenetic analysis with molecular
data and traditional morphological analysis, we place this new genus within Antemninae, a monotypic
Mantidae subfamily. We update the subfamily concept for Antemninae and provide a key to the two genera.
We describe the external morphology of immatures and adults of the new species as well as the genital
complexes of both sexes and the ootheca of Antemna rapax. The observed morphological changes between
immature and adult females suggests that the selection for an alternate strategy for crypsis is a response
to size increase of the abdomen during development. Immatures exploit a stick/branch habitat based on
their morphological appearance while adult females appear as a leaf to disguise the profile of the body.
We perform phylogenetic analyses of the ‘polymorphic earless praying mantises’, a heterogeneous assemblage comprising c. 55% of mantodean diversity in the Neotropics. Bayesian and maximum-likelihood were implemented on a DNA dataset of 9949 aligned nucleic acid characters comprising ten mitochondrial and nuclear genes. Our analyses largely resolved congruent relationships with high levels of support for higher-level taxonomic groups, but revealed extensive inconsistencies between the resolved topology and morphology-based classification systems. The polymorphic earless praying mantises, now granted superfamily status as the Acanthopoidea stat. n., comprises 8 families, 15 subfamilies and 18 tribes. Our newly revised organization required the following taxonomic changes: (i) Thespidae sensu n., including subfamilies Pseudopogonogastrinae subfam. n., Pseudomiopteryginae sensu n., Bantiinae subfam. n., Miobantiinae sensu n. and Thespinae sensu n. (tribes Musoniellini trib. n. and Thespini sensu n.); (ii) Angelidae stat. n. et sensu n.; (iii) Coptopterygidae stat. n.; (iv) Liturgusidae sensu n.; (v) Photinaidae stat. n., including Macromantinae stat. n., Cardiopterinae stat. n., Photiomantinae subfam. n. and Photinainae sensu n. (tribes Microphotinini trib. n., Orthoderellini stat. n. and Photinaini sensu n.); (vi) Stenophyllidae stat. n.; (vii) Acontistidae stat. n.; and (viii) Acanthopidae sensu n. Our new system also resulted in the reassignment of various genera to new and existing higher-level taxa, the exclusion of old world genera otherwise traditionally classified among the Thespidae, Liturgusidae and Angelidae, the confirmation of Stenophylla Westwood as member of this clade, and the revalidation of Paradiabantia Piza stat. r. We provide diagnoses for all suprageneric taxa using external morphological characters and male genitalia. A key to higher-level groups is provided. We incorporate egg case structural variation as a novel approach for taxon delineation.
This review presents a detailed account of taxonomic studies of praying mantids (Mantodea) from the Neotropical region. The current knowledge for each genus is presented along with a brief review of the advances in systematics for the order with emphasis on the Neotropical fauna. Additional comments on the limitations in taxonomic studies in mantids are also discussed. Resumen Esta revisión presenta un registro detallado de los estudios taxonómicos de las "mantis religiosas" (Mantodea) de la región Neotropical. Se presenta un resumen del estado del conocimiento de la taxonomía de cada género así como también una breve revisión de los avances recientes en el estudio de su sistemática y comentarios adicionales sobre las limitaciones en estudios taxonómicos en el orden Mantodea.
Genitalia are rapidly evolving morphological structures most likely under sexual selection. Due to their internal nature they are often hidden inside the body, thus morpho-functional studies of animal genitalia are broadly lacking. Males of some bushcricket taxa bear paired genital appendices called titillators, the exact function of which is unknown since they are obscured inside the female body during pairing. To investigate titillator morphology and possible function during copulation, we studied the bushcricket Metrioptera roeselii (Hagenbach, 1822) using a novel combination of independent, yet complementary, techniques. Copulating pairs were snap-frozen and scanned by X-ray micro-computed tomography (μCT) to visualize the coupling of male and female genitalia in situ. Video recordings of copulating pairs also showed rhythmical insertion of male titillators into the female's genital chamber, where they percuss a softened structure on the female's subgenital plate. Movements did not induce damage to the female's structure, which lacks any sclerotized genital counterparts. Instead, scanning electron microscopy and histological sections show the female subgenital plate to be covered with two different types of sensory receptors at the contact zone between the male's titillator and the female genital chamber. We interpret the non-harmful function of the titillator processes, the lack of a genital counter-structure and the presence of sensory cells on the female's subgenital plate as indicators of a copulatory courtship function of titillators, subject to sexual selection by female choice.
Copyright © 2015. Published by Elsevier Ltd.
We present a quantitative method for describing how heterochronic changes in ontogeny relate to phyletic trends. This is a step towards creating a unified view of developmental biology and evolutionary ecology in the study of morphological evolution. Using this representation, we obtain a greatly simplified and logical scheme of classification. We believe that this scheme will be particularly useful in studying the data of paleontology and comparative morphology and in the analysis of processes leading to adaptive radiation. We illustrate this scheme by examples drawn from the literature and our own work.
The taxonomy of the Neotropical “ant-mantis” Mantillica Westwood, 1889 is revised and the external morphology is re-described. The genus Mantillica (Thespidae: Oligonicinae: Oligonicini) contains only one currently known species, M. nigricans Westwood, 1889. Mantillica sialidea Westwood, 1889 is transferred to the genus Thrinaconyx Saussure as Thrinaconyx sialidea (Westwood, 1889) comb. n. A description of the female and the male external genitalia of M. nigricans is presented, including some aspects of its ant-mimicry adaptation. Remarkable characteristics of Mantillica include the short pronotum and notable myrmecomorphic features in nymphs and females. This species likely belongs to a Batesian mimic complex that include the aggressive Neoponera apicalis (Latreille) ant models. The known distribution of Mantillica is restricted to the Brazilian Amazon, which suggests M. nigricans as endemic of this region, probably with additional populations in the Amazonian areas of Colombia, Peru and Venezuela.
With approximately 2400 described species distributed worldwide (Ehrmann, 2002), praying mantises (Mantodea) exhibit extensive variation in morphological adaptations and life history strategies, typically in connection to their strict predatory habits. Praying mantis diversity is frequently under-appreciated, mostly because of their sedentary and highly cryptic lifestyle, often resorting to various forms of mimicry and mimesis, resembling sticks, flowers, tree bark, bird droppings, pebbles, moss, lichen, and green and dead leafs. Because of these attributes, they are difficult to collect and observe in the field. Certain aspects of praying mantis ecology, behavior and physiology have historically received more attention than others, though most studies focus on a handful of species from temperate regions (see Prete et al ., 1999). In contrast, tropical regions (which hold the bulk of Mantodea biodiversity) have received limited attention by researchers, that attention often punctuated. Their apparent lack of economic importance, rarity in collections and the ongoing “taxonomic impediment”, have all conspired to impede taxonomic and evolutionary studies. The consequences of these deficiencies are far reaching. For example, the current dearth of detailed taxonomic treatments and identification keys prevent the accurate assessment of regional faunas (Rivera, 2010). Further, the lack of a solid taxonomic foundation precludes the formal documentation of relevant aspects of natural history, a discipline that has also faced a steady decline since the early 20 th Century (Hampton & Wheeler, 2012), but nonetheless is fundamental for fostering scientific inquiry and hypotheses formulation. The diversity of ecological strategies and adaptations of mantises, their role as predators, and their well-known (but little understood) cannibalistic sexual behaviors, make of these charismatic insects also outstanding model organisms to explore broader questions in ecology and evolutionary biology.
The Neotropical praying mantis genus Miobantia Giglio-Tos, 1917 currently includes six species with a complex taxonomic history. Although frequently found in the atlantic forest of Brazil, little is known about these species. Several obstacles make it difficult identifying these mantids, including high levels of sexual dimorphism and both sexes known only for one of the six currently known species. The taxonomic boundaries among the species of Miobantia are proposed in this work, through a cladistic analysis using 99 morphological characters, and the analysis of sequence variation of approximately 700 bp of the mithocondrial gene COI for association of dimorphic sexes and nymphs. Additionally, an investigation of intraspecific variation is conducted based on a large number of specimens of M. fuscata (Giglio-Tos) in oder to choose the most relevant features for the separation of the species to be used in taxonomic descriptions and identification keys. The male of M. aptera Giglio-Tos, and the females of M. ciliata (Stål) and M. fuscata are described for the first time, and redescription of the males of these and the female of M. aptera, are provided based on additional data. Miobantia nebulosa (Giglio-Tos) is herein redescribed (male genitalia included) and proposed as a junior synonym of M. rustica (Fabr.). Additionally, four new species are here described, totaling nine species for the genus: M. immanis n. sp. (Espírito Santo, Brazil), M. arctissima n. sp. (Espírito Santo, Brazil), M. sulista n. sp. (Santa Catarina and Rio Grande do Sul, Brazil; San Bernardino, Paraguay), and M. nordestina n. sp. (Bahia, Brazil); both sexes of all new species are described, except for M. arctissima, which remain known by males only. Identification keys and figures of diagnostic features are provided for both males and females of all valid species. The species distribution is mapped according to literature data and 50 new records.
Collections of biological specimens are fundamental to scientific understanding and characterization of natural diversity. This paper presents a system for liberating useful information from physical collections by bringing specimens into the digital domain so they can be more readily shared, analyzed, annotated and compared. It focuses on insects and is strongly motivated by the desire to accelerate and augment current practices in insect taxonomy which predominantly use text, 2D diagrams and images to describe and characterize species. While these traditional kinds of descriptions are informative and useful, they cannot cover insect specimens "from all angles" and precious specimens are still exchanged between researchers and collections for this reason. Furthermore, insects can be complex in structure and pose many challenges to computer vision systems. We present a new prototype for a practical, cost-effective system of off-the-shelf components to acquire natural-colour 3D models of insects from around 3mm to 30mm in length. Colour images are captured from different angles and focal depths using a digital single lens reflex (DSLR) camera rig and two-axis turntable. These 2D images are processed into 3D reconstructions using software based on a visual hull algorithm. The resulting models are compact (around 10 megabytes), afford excellent optical resolution, and can be readily embedded into documents and web pages, as well as viewed on mobile devices. The system is portable, safe, relatively affordable, and complements the sort of volumetric data that can be acquired by computed tomography. This system provides a new way to augment the description and documentation of insect species holotypes, reducing the need to handle or ship specimens. It opens up new opportunities to collect data for research, education, art, entertainment, biodiversity assessment and biosecurity control.
Heterochrony can be defined as change to the timing or rate of development relative to the ancestor. Because organisms generally change in shape as well as increase in size during their development, any variation to the duration of growth or to the rate of growth of different parts of the organism can cause morphological changes in the descendant form. Heterochrony takes the form of both increased and decreased degrees of development, known as “peramorphosis” and “paedomorphosis,” respectively. These are the morphological consequences of the operation of processes that change the duration of the period of an individual’s growth, either starting or stopping it earlier or later than in the ancestor, or by extending or contracting the period of growth. Heterochrony operates both intra- and interspecifically and is the source of much intraspecific variation. It is often also the cause of sexual dimorphism. Selection of a sequence of species with a specific heterochronic trait can produce evolutionary trends in the form of pera- or paedomorphoclines. Many different life history traits arise from the operation of heterochronic processes, and these may sometimes be the targets of selection rather than morphological features themselves. It has been suggested that some significant steps in evolution, such as the evolution of vertebrates, were engendered by heterochrony. Human evolution was fuelled by heterochrony, with some traits, such as a large brain, being peramorphic, whereas others, such as reduced jaw size, are paedomorphic.
A brief account of the history of insect morphology is given. Different techniques and analytical methods used in current projects on insect morphology and phylogeny and their optimized combined application are described. These include fixation, dissection, maceration, histology (microtome sectioning), scanning electron microscopy (SEM), transmission electron microscopy (TEM), serial block-face scanning electron microscopy (SBFSEM), focused ion beam scanning electron microscopy (FIB/SEM), confocal laser scanning microscopy (CLSM), bleaching, micro-computed tomography (μCT), computer-based three-dimensional reconstruction, focus stacking of digital images, geometric morphometrics and the storage of morphological metadata. The role of insect morphology in the “age of phylogenomics” is discussed.
The concept of heterochrony has long had a central place in evolutionary theory. During their long history, heterochrony and several associated concepts such as paedomorphosis and neoteny have often been contentious and they continue to be criticized. Despite these criticisms, we review many examples showing that heterochrony and its associated concepts are increasingly cited and used in many areas of evolutionary study. Furthermore, major strides are being made in our understanding of the underlying genetic and developmental mechanisms of heterochrony, and in the methods used to describe heterochronic changes. A general theme of this accumulating research is that some of the simplistic notions of heterochrony, such as terminal addition, simple rate genes, and “pure” heterochronic categories are invalid. However, this research also shows that a more sophisticated view of the hierarchical nature of heterochrony provides many useful insights and improves our understanding of how ontogenetic changes are translated into phylogenetic changes.
We utilized a long-term data base collected over a broad geographic range to examine predator size - prey size relationships for 18 species of marine fish predators from continental shelf waters off the northeast US coast. Regression analysis was used to illustrate interspecific variation in ontogenetic patterns of prey size use, gape allometries, and ratio-based trophic niche breadths. Size- based feeding strategies were assessed through comparison of frequency distributions of relative prey sizes eaten and were related to general predator feeding tactics and gape morphology. The results demonstrated that the range of prey sizes eaten expanded with increasing predator body size for each of the marine predators examined, leading to asymmetric predator size - prey size distribu- tions. Absolute maximum prey size and slopes of maximum prey size versus predator size varied widely among predator taxa. Distinct size-based feeding strategies were evident, as diets of some predators were dominated by prey that were 10 to 20% of predator size, whereas other predators frequently consumed prey >50% of predator size. Gape sizes and allometric relationships with body size were also diverse among predators and often were closely associated with maximum prey sizes. Ratio-based trophic-niche breadths generally did not expand with predator ontogeny and tended to narrow for the largest predators, which may be common for animal taxa.
Insecticide fogging is often used to document arthropod species richnessin forest canopies, but this technique may not effectively sample invertebratesthat are concealed within a variety of microhabitats. We quantified the effectsof fogging on invertebrates in canopy epiphyte mats of a Costa Rican cloudforest by extracting arthropods from 18 paired pre- and post-fogging samples.Mean abundance and morphospecies richness of living arthropods were respectivelyreduced by 33 and 30% in epiphyte material after fogging, but most organismssurvived the treatment. Relative abundances of major taxa were unaffected byfogging. Herbivores were less abundant after fogging than other trophic groups,and the median body length of non-mite arthropods present in epiphytes wassignificantly smaller after fogging. Examination of seven post-fogging samplesshowed that many arthropods killed by insecticide remained trapped within theepiphyte material. These results provide the first quantitative assessment of aspecific component of arboreal arthropod biodiversity that is missed by thefogging technique.
Ambush predation is characterized by an animal scanning the environment from a concealed position and then rapidly executing a surprise attack. Mantis shrimp (Stomatopoda) consist of both ambush predators ('spearers') and foragers ('smashers'). Spearers hide in sandy burrows and capture evasive prey, whereas smashers search for prey away from their burrows and typically hammer hard-shelled, sedentary prey. Here, we examined the kinematics, morphology and field behavior of spearing mantis shrimp and compared them with previously studied smashers. Using two species with dramatically different adult sizes, we found that strikes produced by the diminutive species, Alachosquilla vicina, were faster (mean peak speed 5.72±0.91 m s(-1); mean duration 3.26±0.41 ms) than the strikes produced by the large species, Lysiosquillina maculata (mean peak speed 2.30±0.85 m s(-1); mean duration 24.98±9.68 ms). Micro-computed tomography and dissections showed that both species have the spring and latch structures that are used in other species for producing a spring-loaded strike; however, kinematic analyses indicated that only A. vicina consistently engages the elastic mechanism. In the field, L. maculata ambushed evasive prey primarily at night while hidden in burrows, striking with both long and short durations compared with laboratory videos. We expected ambush predators to strike with very high speeds, yet instead we found that these spearing mantis shrimp struck more slowly and with longer durations than smashers. Nonetheless, the strikes of spearers occurred at similar speeds and durations to those of other aquatic predators of evasive prey. Although counterintuitive, these findings suggest that ambush predators do not actually need to produce extremely high speeds, and that the very fastest predators are using speed to achieve other mechanical feats, such as producing large impact forces.
Epiphyte mats (contiguous pieces of live and dead epiphytes perched upon branches of trees) are a conspicuous component of tropical cloud forests and harbor diverse meso-and microarthropod communities. We investigated differences in arthropod assem-blage structure between the vegetative (green) and humic (brown) portions of epiphyte mats in a lower montane forest in Monteverde, Costa Rica. Because of qualitative differences between the two substrates, we hypothesized that they would support different arthropod communities and that variation in community parameters would be linked to the quantity of brown material present in a mat sample. The green fraction contained twice as many individuals and species per gram dry mass than the brown fraction. Morphospecies composition was very similar between green and brown portions, but the relative abundance of several taxa differed significantly between the substrates. Contrary to our prediction, total arthropod abundance and richness in a sample were not correlated with the proportion of brown material present. In laboratory trials, the most common morphospecies of oribatid mite in this system showed a preference for brown substrates.
1.
A series of preliminary experiments are conducted to determine the most convenient quantity of food mantids must receive in order to ensure a long life span and a high level of striking.
2.
In other series of experiments the dynamic components of the strike are analysed. Preparatory movements tend to place the animal in an optimum position for striking and probably integrate a mechanism for estimating distances. Movements during the strike itself include the sudden displacement of the predatory legs and the lunge, i.e. a sudden thrust which takes place simultaneously with the movement of the predatory legs. An especial study of this latter component is presented. Computation previous to strike has to take this lunge into account in order to hit successfully.
3.
The maximum motivation distance to strike lies between 70–80% of the foreleg length. Optimum hitting is achieved at a distance that represents 30–40% of foreleg extension. The gap between these two distances, i.e. 70–80% and 30–40%, is filled by the lunge. When the lunge is prevented by immobilizing the animal, a dramatic increase in the number of misses occurs.
4.
There is a relationship between the maximum motivation distance to strike and optimum distance to hit, on the one hand, and the salient anatomical features of the grasping mechanism and the length of the forelegs, on the other hand.
X-ray micro-CT is a powerful tool to visualize without damage details of the inner structures of beetles, the largest order of insects with a hard external skeleton. This contribution shows the three-dimensional (3D) reconstruction of the head morphology of three rove beetle species (Insecta, Coleoptera, Staphylinidae)--Noddia sp., Creophilus maxillosus, and Hesperosoma sp.--using X-ray microtomography at a spatial resolution of 6 μm. The details of skeletal muscle fiber insertions are described, giving a comprehensive overview of mandible mobility and organization. With the support of 3D rendering, we discuss the relationship among the mandible forms, the development of the muscles controlling the movement, and the head morphology. The well-developed posterior part of the head capsule is always accompanied by a well-developed mandible, a large adductor muscle, and a large apodeme for the wide areas of the muscle fiber attachment. In Noddia sp., muscles connected to the posterolateral angle of the head capsule are mainly short muscles, whereas in Creophilus maxillosus, the latter are mainly long muscles, and in Hesperosoma sp. no mandible adductor muscle fibers are present on the posterolateral angle of the head capsule. These results offer new invaluable information regarding the biting functions of beetle mandibles and the trend of their morphological change during their long-term evolution.
After a brief synopsis of the history of mantodean classification, a re-organized systematic arrangement of extant praying mantids is provided. To overcome past homoplasy problems, a phylogenetic framework based on male genital structure was used, supplemented by published morphological, chromosomal and molecular data. As already noticed by previous authors, external morphology is highly homoplastic and does not provide useful systematic tools above subfamily level. In contrast, the morphology of male external genitalia is largely congruent with the results of recent molecular phylogenies, but contradicts the most widely used past systems. Additionally, some genital structures widely used for taxonomic purposes could be shown to be not homologous, most notably the distal process. Evolutionary transitions of the distal process and the phalloid apophysis across the mantodean phylogenetic tree are identified and named. The phalloid apophysis of many derived mantodeans shows a tendency towards bifurcation into an anterior and a posterior lobe. This and other observed genital traits are hypothesized to be an adaptation of males towards a stable copulatory grasp in groups exhibiting increased sexual dimorphism, associated with an increased risk for the male to be cannibalized during copulation. Genital characters allowed most genera to be unambiguously assigned to the major clades (superfamilies) recovered by our genital and previous molecular data. The few exceptions concern genera with secondarily simplified genitalia lacking diagnostic structures. Taxonomic literature is very heterogeneous, and several subfamilies yet lacking any modern revisionary treatment will need further refinement. To account for phylogenetic constraints, i.e. correct for past polyphyletic groupings, the number of families was elevated to 29, and the number of subfamilies to 60. We establish the new family Leptomantellidae, the new subfamilies Brancsikiinae and Deiphobinae, the new tribes Leptomiopterygini, Hagiomantini, Gonypetellini, Bolbellini, Epsomantini, Neomantini, Amantini, Armenini, Danuriellini, Deiphobini, Cotigaonopsini, Didymocoryphini, Oxyelaeini, Heterochaetulini, Rhodomantini and Pseudoxyopsidini, and the new subtribes Amphecostephanina, Bolbina, Tricondylomimina, Gonypetyllina, Antistiina, Toxomantina and Tarachomantina. New morphological diagnoses are provided for the currently recognized families. Despite a few yet to be solved problems, this work offers the urgently needed working base for future studies in Mantodean systematics, life
history and ecology.
Fabrication goes for a quick spin
Most 3D printing techniques involve adding material layer by layer. This sets some limitations on the types of applications for which 3D printing is suitable, such as printing around a preexisting object. Kelly et al. present a different method for manufacturing by rotating a photopolymer in a dynamically evolving light field (see the Perspective by Hart and Rao). This allowed them to print entire complex objects through one complete revolution, circumventing the need for layering. The method may be particularly useful for high-viscosity photopolymers and multimaterial fabrication.
Science , this issue p. 1075 ; see also p. 1042
The Mantodea, or praying mantises, are easily recognized by their most apparent apomorphy ‐ the prothoracic pair of legs having evolved into massive raptorial legs for grasping prey. Praying mantises, although encompassing only 2494 known species, display amazing morphological diversity. The group has conquered every possible habitat except cold and aquatic environments, thus having to adapt to habitats from the African sandy deserts to rainforests, and from living on the ground up to the canopy. Comprehensive taxonomic studies were published, treating large numbers of species and revising their taxonomy by applying both molecular and morphological data. This chapter shows a simplified phylogenetic tree indicating which large‐scale monophyletic groups have been recently revised. Molecular and morphological studies both show that the Mantodea tend to form ecomorphs. Whenever a new habitat is conquered, selection drives the mantodean body into a suitable shape.
Three-dimensional printing technology has shown its importance in many fields. In this study, the potential of this technique in zoological systematics was assessed. For the first time, 3D printed models were incorporated in the description of a new genus as a complement to pictures and drawings to illustrate complex 3D structures and to be used in education. Hereby, we also tested the performances of different printing materials and suggest resin as the most suitable option for the zoological field. As a case study, Labrys chinensis gen. nov., sp. nov. was described using an integrative approach: detailed morphology based on light- and electron microscopy, phylogenetic position as revealed from two ribosomal RNA genes, generic traits were tested for homoplasy, and the intra- and interpopulation variations of four sampled populations were analyzed. The new genus belongs to the subfamily Tylenchinae, family Tylenchidae in the infraorder Tylenchomorpha. It is characterized by a unique labial plate that has four narrow lobes with tips detached from the adjacent cuticle, laterally broad elongated amphidial apertures, a strong sclerotized excretory duct, a round spacious postvulval uterine sac, and a spicule with a sharp protrusion at the blade.
Functional systems, such as feeding mechanics, often involve the evolution of several components of the musculoskeletal system that are moved in coordination to capture prey. Because these systems often involve the quick movement of several structures, some feeding systems have been hypothesized to be stereotypic. While the motor activity patterns are often stereotyped, the subsequent kinematics can be variable, many times in response to variation in prey stimulus (e.g., prey position). Patterns of feeding kinematics have been well studied among vertebrates, with less attention on invertebrate systems. The goal of this study was to examine the amount of stereotypy in the feeding strike kinematics of praying mantises. We filmed 8 juvenile ghost praying mantises (Phyllocrania paradox) at 1000 Hz, across several days within instar 7. We digitized several points that represent the movement of the coxa, trochanter-femur and tibia of the raptorial foreleg to obtain a set of kinematics including angles and angular velocities of the joint, as well as body lunge. Using the coefficient of variation, we found less stereotypy in the approach stage of the strike compared to the sweep. Using Bonferroni corrected Pearson's correlations of kinematics with prey position we found few traits related to prey position with the exception of some kinematics of the coxa joint and the amount of lunge used during the strike. Our results suggest that several components of the praying mantis strike are stereotypic, while others exhibit flexibility to ensure successful capture of the prey.
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.
These thousand eyes have also been looking upon naturalists for quite a while, but only few have looked back. Usually they were zoologists interested in specific groups which live on, in or under bryophytes; in the role these animals play during initial land colonization by cryptogams; in freshwater associations and in diverse other aspects. Botanists have published far fewer observations, these dealing mainly with fertilization or spore dispersal by invertebrates. Although scattered and uneven, taken as a whole the compiled data offer suggestive insights into the relationships between bryophytes and invertebrates, especially in regard to their co-evolution.
The Neotropical lichen mimicking mantid genus Pseudopogonogaster Beier, 1942 is revisited and re-described. The examination of the type species, P. mirabilis Beier, 1942, the first described species of this lineage, allowed us to determine that those species currently included in Pseudopogonogaster and Calopteromantis Terra, 1982 are all congeneric. As a result, the genus Calopteromantis is now considered to be a junior synonym of Pseudopogonogaster. Accordingly, Calopteromantis hebardi Terra, 1982 and Calopteromantis marulandae Salazar, 2002 are transferred to Pseudopogonogaster. The species Calopteromantis otongica Lombardo & Ayala, 1998 from Ecuador was also found to be a synonym of P. mirabilis. In addition, a new species from northwestern Peru, Pseudopogonogaster kanjaris, is described herein, thus constituting a new genus and species record for this country. Analysis of the distribution of the six recognized species of Pseudopogonogaster suggests that this genus is endemic to the mountain forests of Colombia, Ecuador and northern Peru and its members are specialized inhabitants of epiphytic vegetation, exhibiting unique morphological and chromatic adaptations to these habitats. The newly available information on Pseudopogonogaster also allowed to determine that Calopteromantis terrai Jantsch, 1994, described from Mato Grosso, Brazil, is actually a member of the genus Eumiopteryx Giglio-Tos, 1915 (espidae: Pseudomiopteriginae) and, thus, is transferred to this genus as E. terrai (Jantsch, 1994) comb.n.
The history of high-level classification of praying mantids is presented. The various historical stages that helped to construct our current knowledge on Mantodea taxonomy and systematic are highlighted through an extensive literature review, from pre-Linnaean times to the present. A provisional checklist of the names associated to suprageneric ranks (family, subfamily and tribe) is also provided, in addition to some remarks on the future efforts that need to be conducted in Mantodea research.
A comprehensive taxonomic sampling of Mantodea (praying mantises), covering virtually all higher-level groups, was assembled to reconstruct the phylogeny of the order. Sequence data were generated from five mitochondrial and four nuclear loci (12S rRNA, 16S rRNA, 18S rRNA, 28S rRNA, Histone III, Cytochrome Oxidase I & II, NADH dehydrogenase subunit 4, and Wingless) for 329 mantis exemplars along with seven cockroach and eight termite species. Only seven of 14 families, 14 of 33 subfamilies, and seven of 14 tribes were recovered as monophyletic, indicating that phylogeny is largely incongruent with classification. Mapping biogeographical regions on the phylogeny demonstrated that our results adhere closer to biogeographical distributions than to classification. Specific patterns in distribution suggest that major morphological convergences have confounded taxonomists’ ability to reconstruct natural groups. A major revision of higher-level relationships is in order through a comprehensive investigation of morphology and molecular data. We found that major mantis lineages diverged prior to and during the isolation of geographical regions and subsequent ecomorphic specializations within these regions may have led to convergences in morphology. Divergence time estimation places the origin of Mantodea at the beginning of the Jurassic with most modern mantises originating on Gondwana in the Cretaceous. The first major divergence among modern mantises occurred as a result of the north–south splitting of South America and Africa. Subsequent divergences resulted from the breakup of Gondwana. The position of the Indian subcontinent appears to be central to the diversification of Afrotropical and Indomalayan mantises while Antarctica may have served as the conduit for the mantis invasions into South America and Australasia. When India separated from Antarctica and drifted north it distributed mantis lineages back into the Afrotropics and carried a diverse taxonomic assemblage to Asia.
Tropical cloud forests are functionally important ecosystems, but are severely threatened due to deforestation and fragmentation. Epiphyte mats, accumulations of live vegetation and dead organic matter on tree trunks and branches, are a conspicuous component of cloud forests and harbor diverse assemblages of meso- and microarthropods. We compared the morphospecies richness, composition, and abundance of arthropods in epiphyte mats between primary and secondary forests of Monteverde, Costa Rica, and at two nearby replicate sites. Epiphyte mats were thinner and less structurally diverse in secondary forest. We collected ca 36,000 micro- and mesoarthropods from epiphyte mats in the 2-yr study. Whereas arthropod morphospecies richness did not differ among forest types, arthropod abundance was significantly higher in secondary forest due to larger numbers of ants, especially Solenopsis spp. Arthropod assemblages showed a high degree of taxonomic overlap both within and between primary and secondary forests (Jaccard abundance-based similarity = 0.93-0.96). Although characteristics of the arthropod fauna proved to be similar among sites and between forest types, there was a significant temporal effect: arthropod morphospecies richness in epiphyte mats generally was lower in the dry season (February-May), when many taxa probably became dormant or sought shelter against desiccation in deeper portions of mats. © 2007 The Author(s) Journal compilation © 2007 by The Association for Tropical Biology and Conservation.
Abstract. -We present a quantitative method for describing how heterochronic changes in ontogeny relate to phyletic trends. This is a step towards creating a unified view of developmental biology and evolu- tionary ecology in the study of morphological evolution. Using this ...
Despite several recent analyses on the phylogeny of Neuroptera some questions still remain to be answered. In the present analysis we address these questions by exploring a hitherto unexplored character complex: the tentorium, the internal cuticular support structure of the insect head. We described in detail the tentoria of representatives of all extant neuropteran families and the muscles originating on the tentorium using 3D microCT images and analyzed differences in combination with a large published matrix based on larval characters. We find that the tentorium and associated musculature are a source of phylogenetically informative characters. The addition of the tentorial characters to the larval matrix causes a basad shift of the Sisyridae and clearly supports a clade of all Neuroptera except Sisyridae and Nevrorthidae. A sister group relationship of Coniopterygidae and the dilarid clade is further corroborated. A general trend toward a reduction of the dorsal tentorial arms and the development of laminatentoria is observed. In addition to the phylogenetic analysis, a correlation among the feeding habits, the development of the maxillary muscles, and the laminatentoria is demonstrated.
Heterochrony, or the evolution of ontogeny, has been well studied in embryology and skeletal development, providing insight into morphological and genetic mechanisms of evolution.1-5 However, heterochronic studies of behavior and cognition lag behind in comparison. In a recent study we investigated the ontogeny of social behavior and cognition in humans' closest living relatives, chimpanzees (Pan troglodytes) and bonobos (Pan paniscus). These two species are estimated to have had a chimpanzeelike common ancestor between 0.86 and 1.8 mya.6,7 Bonobos have been argued to exhibit morphological indications of paedomorphism relative to chimpanzees, especially in the cranium, and to exhibit paedomorphic behavior as adults.6-11 We found that bonobos exhibit developmental delays relative to chimpanzees in several aspects of their social behavior and cognition. Here, we describe how placing these results in the framework of heterochrony contributes to understanding behavioral and cognitive differences between adults of these two species and to our knowledge of hominid evolution in general.
Coordination of the complete capture sequence of the praying mantis has been studied in detail using several specially developed photographic techniques.
The mantis was able to attack prey throughout a large three-dimensional capture zone by changing body orientation relative to its perch. This orientation centred prey on the median plane and brought it within an attack zone relative to the prothorax. Alignment with the median plane simplifies the attack since the prey can then be localized using only two dimensions. The attack comprised several stereotyped components which together formed a single movement sequence of all six legs. Although too rapid for visual feedback, a simple mechanism permits steering of these movements to capture prey at particular locations within the attack zone.
These findings are contrasted with those from studies of mantis visual behaviour and a simple mechanism is suggested for how prey location is encoded to produce steering of the attack.
As we celebrate the visionary genius of Carl Linnaeus, it is time to analyse how professional taxonomy interfaces with the rest of biology and beyond. Where next for Linnaeus's heirs, asks H. C. J. Godfray?
The Orthoptera of Costa Rica, part I: Mantidae
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Ontogênese da perna raptoria em “louva-a-deus” (Mantodea): um estudo comparativo de alometria
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Neue und seltene Mantodeen aus deutschen Museen
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Ann Nat Hist Mus Wien 52:126-154