Figures 52-55 - uploaded by John Theodore Huber
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Dicopomorpha echmepterygis male, paratype, ventral. 52 habitus 53 head + prothorax + procoxa 54 apex of gaster 55 mesosoma + base of most legs and metasoma. Scale line = 20 μm, except Fig. 52 = 50 μm. 

Dicopomorpha echmepterygis male, paratype, ventral. 52 habitus 53 head + prothorax + procoxa 54 apex of gaster 55 mesosoma + base of most legs and metasoma. Scale line = 20 μm, except Fig. 52 = 50 μm. 

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A new genus and species of fairyfly, Tinkerbella nana (Hymenoptera: Mymaridae) gen. n. and sp. n., is described from Costa Rica. It is compared with the related genus Kikiki Huber and Beardsley from the Hawaiian Islands, Costa Rica and Trinidad. A specimen of Kikiki huna Huber measured 158 μm long, thus holding the record for the smallest winged in...

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... The hymenopteran ovipositor consists of two pairs of valvifers located at the base, which house the muscles controlling the ovipositor mechanism, along with three pairs of valvulae capable of sliding along each other [154] ( Figure 7A,B). The size of this structure can vary significantly, ranging from micrometers to the longest ovipositors documented in Arthropoda with lengths of over 100 mm [155,156], facilitating oviposition in diverse substrates such as wood, soil, or within other organisms [157] ( Figure 1F, Figure 7B). ...
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The extraordinary adaptations that Hymenoptera (sawflies, wasps, ants, and bees) exhibit on their body surfaces has long intrigued biologists. These adaptations, which enabled the immense success of these insects in a wide range of environments and habitats, include an amazing array of specialized structures facilitating attachment, penetration of substrates, production of sound, perception of volatiles, and delivery of venoms, among others. These morphological features offer valuable insights for biomimetic and bioinspired technological advancements. Here, we explore the biomimetic potential of hymenopteran body surfaces. We highlight recent advancements and outline potential strategic pathways, evaluating their current functions and applications while suggesting promising avenues for further investigations. By studying these fascinating and biologically diverse insects, researchers could develop innovative materials and devices that replicate the efficiency and functionality of insect body structures, driving progress in medical technology, robotics, environmental monitoring, and beyond.
... Agricultural lands often exhibit altered soil properties and reduced habitat complexity, leading to changes in arthropod communities compared to natural forest ecosystems (Huber et al., 2017). Conversely, forests typically support more diverse and stable arthropod populations due to their complex structure and less disturbed soil conditions (Huber et al., 2013). Seasonal variations also play a critical role in shaping these communities, with fluctuations in temperature, moisture, and food availability influencing arthropod activity and diversity (Trebicki et al., 2010). ...
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This study examines the seasonal variations in abundance and diversity of Hymenoptera soil arthropods in agricultural versus forest ecosystems. Soil samples were systematically collected monthly from February 2015 to January 2017 across three distinct seasons-winter, summer, and monsoon-from both agricultural and forested areas in Rajanpalle village, Telangana, India. Using the Shannon-Weiner Index, the research assessed species diversity, while the analysis of seasonal abundance involved evaluating the relative proportions of different Hymenoptera species. The results indicated significant seasonal fluctuations in abundance within the agricultural ecosystem, with notable declines in diversity as reflected by lower Shannon-Weiner Index values. Specifically, species like Camponotus sp. and Monomorium sp. showed decreased diversity during summer months. Conversely, the forest ecosystem displayed more consistent abundance patterns and higher diversity indices throughout the year, suggesting a more stable and diverse habitat. These findings underscore the adverse effects of agricultural practices on soil arthropod communities, emphasizing the importance of preserving forest environments to support higher biodiversity and ecosystem stability.
... Most studies on insect flight focus on macroscopic insects with membraned wings, such as flies, moths and dragonflies [6][7][8][9][10][11]. However, there is a myriad of miniature flying insects with body size less than 1 mm [12,13]; some as small as a few hundred micrometres, e.g. a species of fairyflies called Tinkerbella nana [14]. In contrast to macroscopic insects, the morphology, behaviour and flight of miniature insects and the fluid-structure interactions (FSIs) of their wings are much less well understood, despite having long fascinated scientists [15][16][17]. ...
... These wings are collectively called bristled wings. Studies of a wide range of miniature insects [14,[21][22][23][24][25][26] provide detailed data on the number, shape and micro-structure of the bristles, as well as the dimensions of the central wing pads. In addition, the flight characteristics of several species of bristled-winged insects have been quantified by high-speed videos, allowing for analysis and numerical simulations of flight kinematics and aerodynamics at extremely small size scales [26,27,[29][30][31][32]. ...
... (b) Number of bristles n plotted against the ratio l/B in insects, where l is the average bristle length. Measurements gathered or estimated from [14,[21][22][23][24][25][26][27][28] are tabulated in tables S1 and S2 in the electronic supplementary material. ...
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The smallest flying insects often have bristled wings resembling feathers or combs. We combined experiments and three-dimensional numerical simulations to investigate the trade-off between wing weight and drag generation. In experiments of bristled strips, a reduced physical model of the bristled wing, we found that the elasto-viscous number indicates when reconfiguration occurs in the bristles. Analysis of existing biological data suggested that bristled wings of miniature insects lie below the reconfiguration threshold, thus avoiding drag reduction. Numerical simulations of bristled strips showed that there exist optimal numbers of bristles that maximize the weighted drag when the additional volume due to the bristles is taken into account. We found a scaling relationship between the rescaled optimal numbers and the dimensionless bristle length. This result agrees qualitatively with and provides an upper bound for the bristled wing morphological data analysed in this study.
... Most studies on insect flight focus on macroscopic insects with membraned wings, such as flies, moths, and dragonflies [6][7][8][9][10][11]. However, there is a myriad of miniature flying insects with body size less than 1 mm [12,13]; some as small as a few hundred microns, e.g., a species of fairyflies called Tinkerbella nana [14]. In contrast to macroscopic insects, the morphology, behavior, and flight of miniature insects and the fluidstructure interactions (FSI) of their wings are much less well-understood, despite having long fascinated scientists [15][16][17]. ...
... These wings are collectively called bristled wings. Studies of a wide range of miniature insects [14,[19][20][21][22][23][24] provide detailed data on the number, shape, and micro-structure of the bristles, as well as the dimensions of the central wing pads. In addition, the flight characteristics of several species of bristled-winged insects have been quantified by high-speed videos, allowing for analysis and numerical simulations of flight kinematics and aerodynamics at extremely small size scales [24][25][26][27][28][29]. ...
... The Reynolds number (Re) of the bristled wing is usually estimated based on the wing chord length (Re c ) and has been reported to be Re c =O(1) − O (10) [16,32,34]. [14,[19][20][21][22][23][24][25]35] are tabulated in Tables S1 and S2 in the supplementary information. ...
Preprint
The smallest flying insects often have bristled wings resembling feathers or combs. We combined experiments and three-dimensional numerical simulations to investigate the trade-off between wing weight and drag generation. In experiments of bristled strips, a reduced physical model of the bristled wing, we found that the elasto-viscous number indicates when reconfiguration occurs in the bristles. Analysis of existing biological data suggested that bristled wings of miniature insects lie below the reconfiguration threshold, thus avoiding drag reduction. Numerical simulations of bristled strips showed that there exist optimal numbers of bristles that maximize the weighted drag when the additional volume due to the bristles is taken into account. We found a scaling relationship between the rescaled optimal numbers and the dimensionless bristle length. This result agrees qualitatively with and provides an upper bound for the bristled wing morphological data analyzed in this study.
... Many of the smallest flying insects have bristled wings (Fig. 9), e.g. thrips (Lewis, 1973), tiny beetles (Polilov, 2005) and fairyflies (Huber and Noyes, 2013). ...
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Approximately half of the existing winged-insect species are of very small size (wing length about 0.3-4 mm); they are referred to as miniature insects. Yet until recently, much of what we know about the mechanics of insect flight was derived from studies on relatively large insects, such as hoverflies, honey bees and hawkmoths. Because of their very small size, many miniature insects fly at a Reynolds number (Re) on the order of 10 or less. At such a low Re, the viscous effect of the air is very large: A miniature insect moves through the air as would a bumble bee move through mineral oil. Miniature insects must use new flapping mode and new aerodynamic mechanisms to fly. Over the past decade, much work has been done in the study of the mechanics of flight in miniature insects: novel flapping modes have been discovered and new mechanisms of aerodynamic force generation have been revealed; progress has also been made on the fluid-mechanics related flight problems, such as flight power requirements and flight dynamic stability. This article reviews these developments and discusses potential future directions.
... The number of ommatidia therefore determines the total number of images an eye can form, or its spatial information capacity. Ommatidia can be counted in micrographs, ranging from about 20 in the fairyfly Kikiki huna (body length = 158 µm) 21,22 to over 30,000 in large dragonflies 5 . Compound eyes further divide into two structural groups: apposition eyes, in which pigment cells between ommatidia restrict incoming light to a single rhabdom, such that lens size limits optical sensitivity (Fig. 1a), and superposition eyes, in which light travels through a clear zone that allows many facets to contribute to each point (Fig. 1b), thereby multiplying the final sensitivity. ...
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With a great variety of shapes and sizes, compound eye morphologies give insight into visual ecology, development, and evolution, and inspire novel engineering. In contrast to our own camera-type eyes, compound eyes reveal their resolution, sensitivity, and field of view externally, provided they have spherical curvature and orthogonal ommatidia. Non-spherical compound eyes with skewed ommatidia require measuring internal structures, such as with MicroCT (µCT). Thus far, there is no efficient tool to characterize compound eye optics, from either 2D or 3D data, automatically. Here we present two open-source programs: (1) the ommatidia detecting algorithm (ODA), which measures ommatidia count and diameter in 2D images, and (2) a µCT pipeline (ODA-3D), which calculates anatomical acuity, sensitivity, and field of view across the eye by applying the ODA to 3D data. We validate these algorithms on images, images of replicas, and µCT eye scans from ants, fruit flies, moths, and a bee.
... morphological adaptations, some of which set records among the insects. For example, the smallest insect on Earth is a fairy wasp of the genus Dicopomorpha (Mymaridae) 19 , while the longest egg-laying organ (the ovipositor, measured in absolute size) occurs in Darwin wasps of the genus Megarhyssa (Ichneumonidae) 20 . These two extreme examples have a major life history strategy in common: they are both parasitoids, carnivores that complete their entire life cycle feeding on just one individual prey item, the host 21 . ...
... The first transition between phytophagy and parasitoidism is estimated in the Late Triassic in the most recent common ancestor (MRCA) of Vespina (node 6, Fig. 2; Supplementary Figs. [18][19]. Parasitoidism thus evolved once and remained the dominant life strategy in Hymenoptera, with no subsequent major innovations in life history evolving until the Early Cretaceous around 140 Ma (Fig. 2). ...
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The order Hymenoptera (wasps, ants, sawflies, and bees) represents one of the most diverse animal lineages, but whether specific key innovations have contributed to its diversification is still unknown. We assembled the largest time-calibrated phylogeny of Hymenoptera to date and investigated the origin and possible correlation of particular morphological and behavioral innovations with diversification in the order: the wasp waist of Apocrita; the stinger of Aculeata; parasitoidism, a specialized form of carnivory; and secondary phytophagy, a reversal to plant-feeding. Here, we show that parasitoidism has been the dominant strategy since the Late Triassic in Hymenoptera, but was not an immediate driver of diversification. Instead, transitions to secondary phytophagy (from parasitoidism) had a major influence on diversification rate in Hymenoptera. Support for the stinger and the wasp waist as key innovations remains equivocal, but these traits may have laid the anatomical and behavioral foundations for adaptations more directly associated with diversification.
... The transition within Symphyta from phytophagy mediated by symbiotic fungi in wood wasps to parasitoidism in the orussoid-apocritan lineage denotes a radical change facilitating the subsequent radiation of the order as parasitoids [9,10]. Parasitoidism in Hymenoptera arose about 200-250 Ma [1,3,4], [30][31][32][33][34][35][36][37][38][39][40][41][42][43][44][45] Ma after the origin of the order. It should be noted that Peters et al. [3] include Orussidae within a clade separate from the remaining Symphyta. ...
... Proctotrupomorpha appears to have arisen around 200 Ma, and several lineages within the clade are associated with extreme miniaturization, especially in the Mymarommatidae and the chalcidoid Mymaridae ('fairy flies') and Trichogrammatidae, with fully winged specimens approx. 0.14-mm long [37,38]. It is among the Chalcidoidea, and in particular the egg parasitoids and Sternorrhyncha parasitoids, where the most important biological control agents, for example, Anagyrus lopezi (Encyrtidae), have been selected [39]; see also Box 2. ...
Article
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Parasitoid wasps are the most successful group of insect parasitoids, comprising more than half the known diversity of Hymenoptera and probably most of the unknown diversity. This life-style has enabled them to be used as pest control agents conferring substantial economic benefits to global agriculture. Major lineages of parasitoid wasps include Ichneumonoidea, Ceraphronoidea, Proctotrupomorpha and a number of aculeate families. The parasitoid life-style arose only once among basal Hymenoptera, in the common ancestor of the Orussidae and Apocrita some 200+ Ma ago. The ancestral parasitoid wasp was probably an idiobiont on wood-living beetle larvae. From this comparatively simple biology, Hymenoptera radiated into an incredible diversity of hosts and parasitoid lifestyles, including hyperparasitoidism, kleptoparasitoidism, egg-parasitoidism and polyembryony, in several instances co-opting viruses to subdue their hosts. Many lineages evolved beyond the parasitoid niche, becoming secondarily herbivorous or predatory nest provisioners and eventually giving rise to most instances of insect societies.
... In recent years, the anatomy of the smallest insects (Polilov, 2016(Polilov, , 2017Minelli and Fusco, 2019) and collembolans (Tullbergiidae) (Panina et al., 2019) has been studied in detail, but the miniaturization of arachnids has been studied little, despite the fact that mites are presumably the smallest terrestrial arthropods (Huber and Noyes, 2013;Dunlop, 2019). ...
... Acariform four-legged mites (Acariformes, Eriophyoidea) of the family Eriophyidae are some of the smallest mites (Dunlop, 2019), they are microscopic parasites of plants, closely related to the group of soil mites Nematalycidae (Bolton et al., 2017;Klimov et al., 2018Klimov et al., , 2022. Due to parasitism, Eriophyoidea went the way of miniaturization (Nuzzaci and Alberti, 1996), reaching almost the minimum size possible for multicellulars of 80e90 mm (Huber and Noyes, 2013). ...
Article
Miniaturization is one of the important trends in the evolution of terrestrial arthropods. In order to study adaptations to microscopic sizes, the anatomy of the smallest insects was previously studied, but not the anatomy of the smallest mites. Some of the smallest mites are Eriophyidae. In this study we describe for the first time the anatomy of the mite Achaetocoptes quercifolii, which is about 115 μm long. For this purpose, we used light, scanning, and transmission electron microscopy and performed 3D reconstructions. The anatomy of A. quercifolii is compared with the anatomy of larger representatives of Eriophyoidea. Despite the small size of the studied species, there is no considerable simplification of its anatomy compared to larger four-legged mites. A. quercifolii has a number of miniaturization effects similar to those found in microinsects: a strong increase in the relative volume of the reproductive system, an increase in the relative volume of the brain, reduction in the number and size of cells of the nervous system. As in some larger four-legged mites, A. quercifolii undergoes midgut lysis at the stage of egg production. On the other hand, in A. quercifolii a greater number of opisthosomal muscles are preserved than in larger gall-forming four-legged mites.
... Paraprotopsyllidiids are characterised by their narrowed wings with long marginal setae, a typical feature observed in many very minute flying insects. This design of the wings is hypothesised to play an aerodynamic role during wing movement and posture while decreasing the weight of the insect and providing the minimum required wing surface for effective flight, as commonly observed in some Diptera (Nymphomyiidae), Coleoptera (Ptiliidae), parasitic Hymenoptera (Mymaridae, Trichogrammatidae), Lepidoptera (Epermeniidae, Nepticulidae, Pterophoridae) and Thysanoptera (Horridge, 1956;Ellington, 1980Ellington, , 1984Wootton, 1992;Sunada et al., 2002;Huber & Noyes, 2013;Sato et al., 2013;Santhanakrishnan et al., 2014;Jones et al., 2015Jones et al., , 2016Cheng & Sun, 2018;Kasoju et al., 2018;Ford et al., 2019;Lyu et al., 2019;Yavorskaya et al., 2019;Zhao et al., 2019;Lee et al., 2020;Kolomenskiy et al., 2020;Jiang et al., 2022). Some additional functions of bristles have been suggested, i.e. mechanosensory purposes (Ai, 2013;Valmalette et al., 2015). ...
Article
With new fossils of Protopsyllidioidea discovered from amber, our knowledge of the biodiversity in the superfamily increases, and so does our understanding of the evolution of suborder Sternorrhyncha and its ‘basal’ groups. The new species Burmapsyllidium grimaldii Hakim, Azar & Huang sp. nov., assigned to the family Paraprotopsyllidiidae, is reported from the mid-Cretaceous Burmese amber, and described and illustrated.