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Scanning electron microscopy micrographs of Mantispa styriaca, forelegs. a, c anterior and posterior view of closed raptorial foreleg; b, d anterior and posterior view of open raptorial foreleg; e ventral view of femur. As area of setae, Avs anteroventral spines, Btn basitarsal notch, Ce cutting edge, Cx coxa, Datp distal anterotibial protrusion, Dptp distal posterotibial protrusion, Fe femur, Pvc posteroventral carina, Sbs subbasal spine, Ta tarsus, Ti tibia, Tr trochanter
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The insect leg is a multifunctional device, varying tremendously in form and function within Insecta: from a common walking leg, to burrowing, swimming or jumping devices, up to spinning apparatuses or tools for prey capturing. Raptorial forelegs, as predatory striking and grasping devices, represent a prominent example for convergent evolution wit...
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... The spring's elastic potential energy is turned into kinetic energy during the leg's stem phase, when the body is accelerated up and forward within every stride cycle. Resilin, Cpr56 F and other Cpr, including the elastic proteins in the leg muscles, are good candidates for this elastic storage capacity and may save energetic costs during locomotion as insect leg joints contain numerous elastic springs (Niederegger and Gorb, 2003;Gladun et al., 2009;Büsse et al., 2021;Appel et al., 2022, Fig. 4C-E, G-H). This elevated elastic capacity was, for example, shown in cockroach legs that have a leg resilience ranging from 60% to 75%. ...
... Mantispoidea represent either an intermediate lineage or a close relationship with Chrysopidae (green lacewings) in the order Neuroptera (Winterton et al., 2010;Wang et al., 2017;Winterton et al., 2018;Vasilikopoulos et al., 2020;Cai et al., 2023). They have well-known for the hypermetamorphic larval development and for most adults having praying mantis-like raptorial forelegs (Aspöck, U. & Aspöck, H., 2008;Engel et al., 2018;Ardila-Camacho et al., 2021;Büsse et al., 2021;Snyman et al., 2021). Currently, in Mantispoidea, there are about 530 extant species in 74 genera with a worldwide distribution known, as well as 141 extinct species in 96 genera from Upper Triassic to Miocene. ...
Mantispoidea (Insecta, Neuroptera) are well-known for the hypermetamorphic larval development and for most adults having praying mantis-like raptorial forelegs. Currently, this superfamily comprises Berothidae (beaded lacewings) and three raptorial mantispoid families: Rhachiberothidae (thorny lacewings), Dipteromantispidae and Mantispidae (mantidflies or mantid lacewings). It includes about 530 extant species in 74 genera, and 143 extinct species in 97 genera from the Upper Triassic to the Miocene. Berothidae, well-known for some adults with scale-like setae on the female wings, currently includes 119 extant species in 26 genera, and 59 extinct species in 37 genera from the Upper Triassic to the Eocene. Rhachiberothidae represents the relictual group of Mantispoidea, comprising only 14 extant species across four genera, which are confined to the Afrotropical Region. However, the abundant fossil records (28 species and 21 genera), especially from the Cretaceous, highlight the past glory of this family. The Cretaceous dipteran-like family Dipteromantispidae, comprising ten species across nine genera, remains enigmatic in terms of its phylogenetic status. Mantispidae, the largest family within Mantispoidea, comprises approximately 395 extant species across 44 genera and 46 extinct species in 30 genera, ranging from the Lower Jurassic to the Miocene. This family is distinctive from other raptorial families for its highly mobile and variously elongated prothorax. Here, we describe several new genera and species from mid-Cretaceous Kachin amber, Myanmar: Three new species of Osmyloberotha Khramov, 2021 (O. dispersa sp. nov., O. chenzuyini Zhuo, Li & Liu sp. nov. and O. magnimaculata sp. nov.) in Berothidae, with a detailed redescription of O. simpla Khramov, 2021; two new genera and three new species (Paraberothinae: Paraberothoides longispina Li, Zhang & Liu gen. et sp. nov.; Subfamily unknown: Dicranoberotha zhangzhiqiae Zhuo, Li & Liu gen. et sp. nov., Dicranoberotha liumohanae Zhuo, Li & Liu gen. et sp. nov.) of Rhachiberothidae, three new species (Enigmadipteromantispa dilatata Li, Nakamine, Yamamoto & Liu sp. nov., Kurtodipteromantispa relicta Li, Nakamine, Yamamoto & Liu sp. nov. and Kurtodipteromantispa univenula sp. nov.) of Dipteromantispidae, as well as four new genera and five new species (Symphrasinae: Parvosymphrasites aploneura Li, Lin, Liu & Wang gen. et sp. nov. and Proplega evelynae Zhuo, Li & Liu gen. et sp. nov.; Subfamily unknown: Haplacantha robusta gen. et sp. nov., Haplacantha tenuis gen. et sp. nov. and Mesomantispoides felixoporcus gen. et sp. nov.) of Mantispidae. Sinuijumantispa So & Won, 2022, previously classified within Drepanicinae, is now recognized as a stem group of Mantispidae, which is possibly related to Mesomantispoides gen. nov. or Mesomantispinae. Our new findings offer crucial insights into the taxonomy, morphological diversity and evolution of Mantispoidea.
... Garcia-Castineiras et al. 1978;Fujimori 1978;Gast and Lee 1978;Giurginca et al. 2015). This protocol was applied and enabled to find functional significances of material properties in many studies on arthropod endocuticle (Wang et al. 2019), wings Ma et al. 2022), foot attachment devices and legs (Peisker et al. 2013;Rebora et al. 2018;Petersen et al. 2018;Friedrich and Kubiak 2018;Jandausch et al. 2018;Beutel et al. 2020;Büsse et al. 2021;Li et al. 2021;Büscher et al. 2021), thorax structure (Casey et al. 2022), antennae (Saltin et al. 2019), mouthparts Michels and Gorb 2015;Büsse and Gorb 2018;Weihmann and Wipfler 2019;Matsumura et al. 2021a, b;Lehnert et al. 2021;Sun et al. 2021;Wei et al. 2022) or genitals (Matsumura et al. 2014(Matsumura et al. , 2017(Matsumura et al. , 2020a(Matsumura et al. , b, 2021aKamimura et al. 2021). In addition, this protocol was cross-validated by employing AFM-Nanoindentation at least for hair of foot attachment devices in a lady beetle (Peisker et al. 2013). ...
The radula, a chitinous membrane spiked with teeth, is the molluscan autapomorphy for the gathering and processing of food. The teeth, as actual interfaces between the organism and the ingesta, act as load transmitting regions and have to withstand high stresses during foraging-without structural failure or high degrees of wear. Mechanisms contributing to this were studied previously in paludomid gastropods from Lake Tanganyika. For some species, gradients in hardness and Young's modulus along the teeth were detected, enabling the bending and relying of teeth onto the next row, distributing the stresses more equally. The here presented study on one of them-Lavigeria grandis-aims at shedding light on the origin of these functional gradients. The mechanical properties were identified by nanoindentation technique and compared to the elemental composition, determined by elemental dispersive X-ray spectroscopy (EDX, EDS). This was done for the complete radular (mature and immature tooth rows), resulting in overall 236 EDX and 700 nanoindentation measurements. Even though teeth showed regional differences in elemental composition, we could not correlate the mechanical gradients with the elemental proportions. By applying confocal laser scanning microscopy (CLSM), we were finally able to relate the mechanical properties with the degree of tanning. CLSM is a common technique used on arthropod cuticle, but was never applied on radular teeth before. In general, we found that nanoindentation and CLSM techniques complement one another, as for example, CLSM is capable of revealing heterogeneities in material or micro-gradients, which leads to a better understanding of the functionalities of biological materials and structures.
... Raptorial foreleg As Rhachiberothidae, Symphracinae, and Calomantispinae, the procoxal surface of Doratomantispinae is smooth, lacking the posterodistal impressed (present in Drepanicinae) or flattened (present in Drepanicinae and Mantispinae) area, or with a proximal annular groove (present in Mantispinae) (Ardila-Camacho et al., 2021;Lambkin, 1986a, b;Li et al., 2020a). The protrochanter of Doratomantispinae is similar to that of Drepanicinae and Calomantispinae, being subconical with the "trochanterfemur complex" association less developed than in Mantispinae (Ardila-Camacho et al., 2021;Büsse et al., 2021). ...
Abstract
Doratomantispinae is an extinct mantispid subfamily reported only from mid-Cretaceous Myanmar amber, which includes two
genera, Doratomantispa Poinar in Poinar & Buckley, 2011 and Paradoxomantispa Lu et al., 2020, and six species. Here, we
present an overview of the genera and species of the subfamily, with descriptions of seven new species, namely D. arcimaculata
sp. nov., D. gaoyuhei sp. nov., D. zhangwenjuni sp. nov., D. zhangzhiqiae sp. nov., D. zhuozhengmingi sp. nov., D. yumeiyingae
sp. nov., and P. mahaiyingae sp. nov.. We also synonymize Lonchomantispa Shi, Yang & Ren in Shi et al., 2020 with Dorato�mantispa, proposing the new combination: D. longa (Shi, Yang & Ren in Shi et al., 2020) comb. nov.. The adult morphology of
Doratomantispinae is studied in detail and comprehensively compared with many other mantispoids. We found some putative
apomorphies supporting the monophyly of Doratomantispinae and its close relationship with the derived mantispid clade. This
subfamily represents a lineage with the richest species diversity among mantispids known from Myanmar amber
... Rhachiberothinae and Drepanicinae possess a subconical trochanter, in Mantispinae it is tubular and curved, whereas in Calomantispinae it is ovoid. This podomere was described in Mantispa styriaca as strongly associated or nearly fused with the femur forming the "trochanter-femur complex" (Büsse et al. 2021). In such complex, the movement between both podomeres is restricted, allowing a higher freedom in the articulation between the coxa and the trochanter. ...
... The femoral integumentary specializations of the Calomantispinae are similarly arranged to those of Drepanicinae, although a smaller variety of processes and setae are present (Figs 22d, e). Posteroventral row is long-almost reaching the femoral base -, situated over a slightly laterally compressed "posteroventral carina" (Büsse et al. 2021), and composed by two spaced primary processes located on the distal femur half (Figs 22d, e, 27a-c, 29). variably sized secondary processes are present proximally, between, and distally to the primary processes. ...
... Mantispinae species have tibia noticeably shorter than the femur (Figs 22f-h); its length ratio to the femur varies between 0.5-0.6. The tibial base is markedly curved at the joint with the femur; there is on the tibial anterior surface, a patch of clavate setae, which extends over almost the entire length (Figs 28d, g); the distal margin of the tibia is produced into a blade-shaped process, referred to as a "distal anterotibial protrusion" by Büsse et al. (2021) (Fig. 28b); the ventral keel is well developed, thin, lacking prostrate setae, and with two lateral rows of trichoid setae, which is a similar condition to that of extant Rhachiberothinae (Figs 28d, g). ...
Adult external morphology of the extant raptorial Mantispoidea (Insecta: Neuroptera: Mantispidae and Rhachiberothidae) is compared emphasizing the morphology of the subfamily Symphrasinae as a key group to understand the phylogenetic relationships among the members of the superfamily. Plega dactylota Rehn, 1939 is thoroughly characterized in order to exemplify the morphology of the Symphrasinae. Additionally, following a review of the literature and examination of comparative material of Dilaridae, Berothidae, Rhachiberothidae and all Mantispidae subfamilies, a new interpretation of the components of the raptorial apparatus (i.e., head, prothorax, grasping forelegs, as well as integumentary specializations) is presented. Also, wing venation for these groups is reinterpreted, and new homology hypotheses for wing venation are proposed based on tracheation and comparative analyses. Given the high morphological divergence on the genital sclerites within the Mantispoidea, plus the confusing previous usage of neutral terminology and terms referring to appendages across taxonomic and morphological studies, we attempt to standardize, simplify, and situate terminology in an evolutionary context under the “gonocoxite concept” (multi-coxopod hypothesis). The remarkable morphological similarity of the genital sclerites of Symphrasinae and Rhachiberothidae (sensu U. Aspöck & Mansell 1994) with the Nallachinae (Dilaridae) was taken as a starting point to understand the morphology of other Mantispidae subfamilies. Based on these morphological comparisons, we provide a revised phylogenetic analysis of Mantispoidea. This new phylogenetic analysis supports a sister group relationship between the family Rhachiberothidae, comprising Rhachiberothinae and Symphrasinae, and the family Mantispidae, including the subfamily Mantispinae and its sister taxa Drepanicinae and Calomantispinae, which may represent a single subfamily. Based on these analyses, raptorial condition probably evolved a single time in these insects and subsequently became diversified in the two sister clades of the raptorial Mantispoidea.
The form of an animal's limbs has to balance multiple functions: locomotion, grasping, climbing, and jumping, among others. For cryptic animals, especially those that resemble elements of their habitat like sticks or grasses, the limbs may also be modified to enhance the camouflage. The performance of a limb in one category may require a tradeoff, reducing performance in another category. Praying mantises provide a diverse group of insects who all use their forelegs for one function, capturing prey, while some species use them as part of their camouflage. Here we use a large database of images of mantis species to capture the variation in morphology across the order, and to calculate the largest prey that their forelegs can hold. We find that the length and thickness of the femur and the length of the tibia comprise most of the variability across species. The majority of species have similar foreleg morphology, with two large groups extending into areas of the morphospace with thicker or thinner forelegs. A geometric relationship between dimensions of the foreleg and the optimal prey diameter maps directly onto the variability across species determined by principal components analysis; legs with thinner femurs and shorter tibia can't hold large prey, and the distribution of the species across the morphospace follows the gradient of optimum prey size. These results suggest that some species trade ability to grasp larger prey for benefits including crypsis, and the praying mantises are an ideal system for studying morphological and functional variation in limbs.
Prehensile raptorial forelegs are prey capturing and grasping devices, best known for praying mantises (Mantodea) within insects. They show strong morphological and behavioral adaptations toward a lifestyle as generalist arthropod predators. In the past, few species of Mantodea were investigated, concerning morphological variability of the raptorial forelegs. Especially the knowledge of foreleg anatomy in the light of functional and comparative morphology is scarce. Our comparative approach is based on the, for arthropods very common, "female-biased sexual size dimorphism" (SSD) that occurs in almost every Mantodea species. Within Mantodea, this SSD is likely leading to a shift of the exploited ecological niche between male and female individuals due to changes in, for example, the possible prey size; which might be reflected in the chosen ecomorphs. In this context, we analyzed the musculature of the raptorial forelegs of female and male specimens in five different species with varying SSD, using high-resolution microcomputed tomography and dissection. We were able to confirm the presence of 15 extrinsic and 15 intrinsic muscles-including one previously undescribed muscle present in all species. Thus, presenting a detailed description and illustrative three-dimensional anatomical visualization of the musculature in Mantodea. Interestingly, almost no observable differences were found, neither between species, nor between the sexes. Furthermore, we homologized all described muscles, due to their attachment points, to the comprehensive nomenclature established by Friedrich and Beutel (2008), discussed potential functionality of the muscles and possible homologies to the neuropteran Mantispa styriaca (Büsse et al., 2021) and the newly introduced leg nomenclature by Aibekova et al. (2022). By elucidating the anatomy, particularly in the context of functionality and SSD, our results complement previous knowledge of the raptorial forelegs, and facilitate a better understanding of the underlying biomechanical system of the predatory strike, and ultimately, a future comparison to other insect taxa.
Spiders are among the most diverse animals, which developed different morphological and behavioral traits for capturing prey. We studied the anatomy and functionality of the rare and apomorphic raptorial spider feet using 3D reconstruction modeling, among other imaging techniques. The evolutionary reconstruction of the raptorial feet (tarsus plus pretarsus) features using a composite tree of spiders, indicating that similar traits emerged three times independently in Trogloraptoridae, Gradungulinae, and Doryonychus raptor (Tetragnathidae). The characteristics defining the raptorial feet are an interlocked complex merging of the base of the elongated prolateral claw with the pretarsal sclerotized ring, with the former clasping against the tarsus. Raptorial feet even flex over robust raptorial macrosetae forming a reduced tarsal version of a catching basket to encase prey during hunting. Our results show that Celaeniini (Araneidae) and Heterogriffus berlandi (Thomisidae), taxa previously compared with raptorial spiders, lack the raptorial feet key characteristics and the tarsal-catching basket. We make predictions about the possible behavior of the abovementioned taxa that will need to be tested by observing living specimens. We conclude that multiple morphological tarsal and pretarsal micro-structures define the raptorial foot functional unit and recommend a comprehensive evaluation before assigning this configuration to any spider taxa.
The diversity of insects can be explained by their ability to establish various ecological niches, which includes the foraging from diverse sources. The cuticle-based feeding structures interact with the food and show adaptations in shape, material composition and mechanical properties to it. Some predatory species are extremely specialised to a very narrow spectrum of prey. In this study, we focus on the mouthparts of a very prominent ambush predator, the antlion larvae of Euroleon nostras . By nanoindentation, we tested the hardness and the Young’s modulus of the mouthparts, which are significantly harder and stiffer than other insect cuticle structures. To gain insight into the origins of the high values, we studied the degree of tanning using confocal laser scanning microscopy and determined the content of inorganic elements by energy dispersive spectroscopy. We could correlate the proportions of the transition metals and alkaline earth metals with the mechanical property values. We also conducted experiments on the breaking stress, the puncturing and biomechanical behaviour of the jaws, which highlighted their extraordinary strength. These findings are not only valuable for biologists, but also for material scientists, as they contribute to our understanding of the origins of mechanical property heterogeneities in insect cuticle.
