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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 family Mymaridae, commonly known as fairyflies (Blackbourn 1935), includes the smallest members of the Chalcidoidea (Hymenoptera) (Annecke & Doutt 1961;Noyes & Valentine 1989;Lin et al. 2007;Huber et al. 2009;Huber & Noyes 2013). The name fairyflies refers to their usually slender bodies, very small size, and their delicate, fringed wings ). ...
... The name fairyflies refers to their usually slender bodies, very small size, and their delicate, fringed wings ). Some are less than 0.19 mm long, being the smallest of all winged insects (Noyes & Valentine 1989;Huber & Noyes 2013), while a few others occurring in the Southern Hemisphere may reach 6.0 mm in length, thus being considered giants of the world mymarids (Noyes & Valentine 1989;Gibson 1997;Huber & Noyes 2013;Pricop 2013;Huber 2017). Mymaridae are cosmopolitan, being an abundant group of Hymenoptera almost everywhere, particularly throughout the tropical and temperate regions (except Antarctica) (Annecke & Doutt 1961;Triapitsyn & Berezovskiy 2001). ...
... The name fairyflies refers to their usually slender bodies, very small size, and their delicate, fringed wings ). Some are less than 0.19 mm long, being the smallest of all winged insects (Noyes & Valentine 1989;Huber & Noyes 2013), while a few others occurring in the Southern Hemisphere may reach 6.0 mm in length, thus being considered giants of the world mymarids (Noyes & Valentine 1989;Gibson 1997;Huber & Noyes 2013;Pricop 2013;Huber 2017). Mymaridae are cosmopolitan, being an abundant group of Hymenoptera almost everywhere, particularly throughout the tropical and temperate regions (except Antarctica) (Annecke & Doutt 1961;Triapitsyn & Berezovskiy 2001). ...
Article
A review of the known Mymaridae (Hymenoptera: Chalcidoidea) in Egypt is presented based on the available literature records and new collections made during 2015–2021. Twenty-two named species in 13 genera are known to occur in Egypt, of which three, Arescon Walker, Dicopus Enock, and Stephanodes Enock, are recorded for the first time from the country, each with a single species. Three species, Anagrus frequens Perkins, Camptoptera papaveris Foerster, and Erythmelus flavovarius (Walker), are also recorded here for the first time from Egypt. A new species, Arescon splendidus Gadallah, Edmardash & Abul-Sood, n. sp., is described and illustrated. The genus Polynema Haliday is excluded from the present study and will be treated separately when fresh specimens of the two species described previously from Egypt are collected from the respective type localities. An identification key based on females for all the Egyptian species of Mymaridae is provided.
... Bristled wing models and non-dimensional parameters. Figure 1a displays a schematic diagram of the bristled wing of a fairyfly, Tinkerbella nana (re-drawn from Ref. 3 ). During flight, the distal part of the wing, where the bristles are nearly parallel to the direction of the wing span, moves faster than the proximal part and generates most of the aerodynamic force. ...
... The aspect ratio (AR) of the wing is defined as the ratio of the wing span (the cylinder length L) to the chord length c. According to the micrographs of the real bristled wings in Ref. 3 , we considered AR = 1.5-2.5 as biologically relevant. To examine the 3-D effect caused by the finite span, we also constructed a 2-D bristled wing model, which is a chordwise transection of the 3-D model and has been widely used by previous researchers (e.g., Refs. ...
... From Fig. 11, we can also see that the flow velocities around the two cylinders closest to the leading edge (cylinders No. 1 and 2) are higher than those around the rest; and the flow fields around cylinders No. 3, 4, 5, and 8 are almost identical, implying that the flow around a cylinder in the bristled wing is affected mainly by www.nature.com/scientificreports/ the two cylinders above and the two cylinders below it. Therefore, the flow velocities around different cylinders in the inner chord (i.e., cylinders No. [3][4][5][6][7][8][9][10][11][12][13], are reduced to the same degree, resulting in the equally low drag coefficients of the cylinders. However, for the leading edge of the wing (see cylinder No. 1 in Fig. 11), there is no cylinder above. ...
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The smallest insects fly with bristled wings at very low Reynolds numbers ( Re ) and use the drag of the wings to provide the weight-supporting force and thrust. Previous studies used two-dimensional (2-D) models to study the aerodynamic force and the detailed flow field of the bristled wings, neglecting the three-dimensional (3-D) effect caused by the finite span. At high Re , the 3-D effect is known to decrease the aerodynamic force on a body, compared with the 2-D case. However, the bristled wing operates at very low Re , for which the 3-D effect is unknown. Here, a 3-D model of the bristled wing is constructed to numerically investigate the detailed flow field and the aerodynamic force of the wing. Our findings are as follows: The 3-D effect at low Re increases the drag of the bristled wing compared with that of the corresponding 2-D wing, which is contrary to that of the high- Re case. The drag increase is limited to the tip region of the bristles and could be explained by the increase of the flow velocity around the tip region. The spanwise length of the drag-increasing region (measuring from the wing tip) is about 0.23 chord length and does not vary as the wing aspect ratio increases. The amount of the drag increment in the tip region does not vary as the wing aspect ratio increases either, leading to the decrease of the drag coefficient with increasing aspect ratio.
... Thus, the compound eyes of large insects contain 1000 times as many ommatidia as those of the smallest flying insects: Anax junius (Odonata, Aeschnidae) has up to 29 247 ommatidia per eye (Sherk, 1978), and the smallest flying insect Kikiki huna (Hymenoptera, Mymaridae) (Huber and Noyes, 2013) has only 25 ommatidia per eye. The following scaling range can be estimated for individual insect orders. ...
... The following scaling range can be estimated for individual insect orders. The number of ommatidia in the largest Hymenoptera is 640 times as great as that in the smallest representatives: 16 000 in Xylocopa latipes (Apidae) (Jander and Jander, 2002) and 25 in Kikiki huna (Huber and Noyes, 2013); for Coleoptera the range is 920 times: 29 450 in Augosoma centaurus (Scarabaeidae) (Rensch, 1959) and 32 in Scydosella mysawasensis (Ptiliidae) (Makarova et al., 2019); for Lepidoptera it is 219 times: about 27 000 in Sphinx convolvuli (Sphingidae) (Mazokhin-Porshnyakov, 1965) and 123 in Stigmella microtheriella (Nepticulidae) (Fischer et al., 2012). ...
... The minimum diameter of facets in many of the smallest microinsects with body length less than 0.4 mm, namely Scydosella musawasensis, Cylindrosella sp. (Coleoptera, Ptiliidae) (Makarova et al., 2019), Kikiki huna (Hymenoptera, Mymaridae) (measured in micrographs in Huber and Noyes, 2013), and Megaphragma caribbea (Hymenoptera, Trichogrammatidae), is also about 6 μm, i.e., far below the theoretical limit (Barlow, 1952). ...
... Various microorganisms tested for producing metal nanoparticles are: the bacterium Pseudomonas fluorescens (60 nm Au NPs), the yeast Yarrowia lipolytica (30 nm Au NP), the sponge Acanthella elongate (13 nm NPs), the algae Stoechospermum marginatum (50 nm NPs), and the fungus Candida albicans [219][220][221]. As an example, Figure 16 show an efficient biosynthesis approach to produce porous structures of bacterial cellulose nanofibers (20-40 nm in diameters) by foaming a mannitol-based media with a bacterial suspension of Gluconoacetobacter xylinus [253,254]. ...
... Various microorganisms tested for producing metal nanoparticles are: the bacterium Pseudomonas fluorescens (60 nm Au NPs), the yeast Yarrowia lipolytica (30 nm Au NP), the sponge Acanthella elongate (13 nm NPs), the algae Stoechospermum marginatum (50 nm NPs), and the fungus Candida albicans [219][220][221]. As an example, Figure 16 show an efficient biosynthesis approach to produce porous structures of bacterial cellulose nanofibers (20-40 nm in diameters) by foaming a mannitol-based media with a bacterial suspension of Gluconoacetobacter xylinus [253,254]. Insects also contain unique nanostructures implicated in specific physical and physiological functions ( Figure 17). Adhesion, chemical sensing, and response, color vision and manipulation, movement, mechano-sensation, and thermoregulation are the mentioned functions [255,256]. ...
... To construct a bacterial cellulose foam, a suspension of G. xylinus in growth media is foamed. © Nature, 2018[254]. ...
Article
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Nanomaterials are becoming important materials in several fields and industries thanks to their very reduced size and shape-related features. Scientists think that nanoparticles and nanostructured materials originated during the Big Bang process from meteorites leading to the formation of the universe and Earth. Since 1990, the term nanotechnology became very popular due to advances in imaging technologies that paved the way to specific industrial applications. Currently, nanoparticles and nanostructured materials are synthesized on a large scale and are indispensable for many industries. This fact fosters and supports research in biochemistry, biophysics, and biochemical engineering applications. Recently, nanotechnology has been combined with other sciences to fabricate new forms of nanomaterials that could be used, for instance, for diagnostic tools, drug delivery systems, energy generation/storage, environmental remediation as well as agriculture and food processing. In contrast with traditional materials, specific features can be integrated into nanoparticles, nanostructures, and nanosystems by simply modifying their scale, shape, and composition. This article first summarizes the history of nanomaterials and nanotechnology. Followed by the progress that led to improved synthesis processes to produce different nanoparticles and nanostructures characterized by specific features. The content finally presents various origins and sources of nanomaterials, synthesis strategies, their toxicity, risks, regulations, and self-aggregation.
... At least 106 of these families include species with adults less than 1 mm long, but species with body lengths smaller than 0.5 mm are to be found only in three families: Mymaridae, Trichogrammatidae (Hymenoptera), and Ptiliidae (Coleoptera). The smallest known insects are the wingless males of the parasitoid fairy wasp Dicopomorpha echmepterygis Mockford, 1997 (Mymaridae) (139 μm) (Mockford, 1997), while the smallest flying insects belong to the genera Kikiki Huber et Beardsley, 2000 (Mymaridae;158 μm) and Megaphragma Timberlake, 1924 (Trichogrammat idae; 170 μm) (Huber and Noyes, 2013;Polilov, 2016). The smallest flying non-parasitoid insects are featherwing beetles of the genus Scydosella Hall, 1999 (Ptili idae) (300 μm) (Polilov, 2016). ...
... Ommatidia usually represent the individual pixels of an image, so their number limits the total number of images the eye can form, or its spatial information capacity, and they can be counted in microscope images. They range from ~20 ommatidia in one of the smallest flying insects, the fairyfly Kikiki huna (body length=158µm; Huber and Beardsley 2000;Huber and Noyes 2013) to >30,000 in large dragonflies, which have enhanced vision for hunting prey (Cronin et al. 2014). ...
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The arthropod compound eye is the most prevalent eye type in the animal kingdom, with an impressive range of shapes and sizes. Studying its natural range of morphologies provides insight into visual ecology, development, and evolution. In contrast to the camera-type eyes we possess, external structures of compound eyes often reveal resolution, sensitivity, and field of view if the eye is spherical. Non-spherical eyes, however, require measuring internal structures using imaging technology like MicroCT (μCT). Thus far, there is no efficient tool to automate characterizing compound eye optics. We present two open-source programs: (1) the ommatidia detecting algorithm (ODA), which automatically measures ommatidia count and diameter, and (2) a μCT pipeline, which calculates anatomical acuity, sensitivity, and field of view across the eye by applying the ODA. We validate these algorithms on images, images of replicas, and μCT scans from eyes of ants, fruit flies, moths, and a bee.
... nov. members, are typically characterized by their wings bearing long marginal setae (Huber and Noyes, 2013). This feature is commonly observed in several families of Thysanoptera, in Coleoptera (Ptiliidae), in parasitic Hymenoptera (Mymaridae, Trichogrammatidae) and some Lepidoptera (Epermeniidae, Nepticulidae, Pterophoridae). ...
... In the last few decades, the function of these bristles in tiny insects was investigated and different roles were proposed. This fringe is mainly believed to serve an aerodynamic purpose during wing-wing interactions in clap and fling movements (Horridge, 1956;Ellington, 1984;Huber and Noyes, 2013;Santhanakrishnan et al., 2014;Jones et al., 2016). These setae can also contribute to the wings' movement and posture, i.e., folding and unfolding of the wings (Ellington, 1980;Grimaldi and Engel, 2005) as commonly seen in thrips. ...
... Sunada et al. (2002) investigated the role of these setae based on a model of a thrip's wing; they concluded that no advantages are provided by the bristles to the aerodynamics of a 'single wing' during lift and drag. Huber and Noyes (2013) discussed the body characteristics of tiny flying insects, more specifically parasitic wasps (Mymaridae). They hypothesized that the long marginal setae may contribute to reduce turbulence and decrease the force needed for drag on a rapidly flapping wing. ...
Article
Four new protopsyllidioid species, Paraprotopsyllidium spinosum gen. et sp. nov., Angustipsyllidium minutum gen. et sp. nov., Burmapsyllidium setosum gen. et sp. nov., and Maliawa akrawna gen et sp. nov. are described from the mid-Cretaceous Burmese amber, and assigned to a new family, Paraprotopsyllidiidae fam. nov., that we establish. These taxa are characterized by their narrowed fore and hind wings, bearing a fringe of long setae; the possible functions of these bristles are explored in this paper. The population dynamics, the potential feeding diet and the biogeographical distribution of this family are briefly discussed.
... The absence in both species examined here is likely linked with exceptionally small size and perhaps indicates a functional transition during miniaturization, as this vestiture may create a boundary layer around larger animals, which may be unnecessary in miniscule ones. Alternatively, the loss of this vestiture may be caused by size or other properties of the epidermal cells underlying the cuticle (Huber & Noyes, 2013). Another groundplan feature of Hymenoptera is the presence of paddle-shaped or spatulate setae (Basibuyuk & Quicke, 1995); we here refer to these as 'compressed' setae as they vary from wide throughout their length, or with a narrow stalk and apical widening, which would represent the true 'spatulate' condition These setae are presumed to function in grooming behaviors and are retained in Megaphragma as compressed setae, an especially important retention as the strigil has been lost. ...
Article
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.
... In the smallest ptiliid species (S. musawasensis), the facet diameter was 6.73 ± 0.23 mm, which is less than one fifth of what is seen in larger staphylinid coleopterans (Meyer-Rochow, 1972). Data obtained for the facet numbers and diameters of Ptiliidae are similar to those from the eyes of the smallest hymenopterans: the eye of the smallest flying mymarid hymenopteran Kikiki huna for instance contains around 25 facets of about 6.6 ± 0.48 mm diameter (counted from Huber and Noyes, 2013). ...
... The smallest flying insect K. huna (Mymaridae) has at most 25 facets per eye (counted from SEM micrographs in Huber and Noyes, 2013). Some species of the genus Megaphragma have about 30 facets per eye (Megaphragma caribea 32 ± 3, M. mymaripenne 29 ± 1, M. amalphitanum 29 ± 1), whereas in the related miniature wasp T. evanescens, despite the considerable similarities of the ultrastructural organization with that of Megaphragma (Makarova et al., 2015), the number of facets is almost 4.5 times higher (Fischer et al., 2010). ...
Article
The coleopteran family Ptiliidae (featherwing beetles) includes some of the smallest insects known with most of the representatives of this family measuring less than 1 mm in body length. A small body size largely determines the morphology, physiology, and biology of an organism and affects the organization of complex sense organs. Information on the organization of the compound eyes of Ptiliidae is scarce. Using scanning electron microscopy we analyzed the eyes of representatives of all subfamilies and tribes and provide a detailed description of the eye ultrastructure of four species (Nephanes titan, Porophila mystacea, Nanosella sp. and Acrotrichis grandicollis) using transmission electron microscopy. The results are compared with available data on larger species of related groups of Staphyliniformia and scale quantitative analyses are performed. The eyes of Ptiliidae consist of 15–50 ommatidia 6–13 μm in diameter and all conform to the apposition acone type of eye with fused rhabdoms of banded organization. Each ommatidium has the typical cellular arrangement present also in the eyes of larger staphyliniform beetles, but strongly curved lenses, short cones, reduced pigment cells, a high density of pigment granules and certain modifications of the rhabdom seem typical of ptiliid eyes. Allometric analyses show that as body size decreases, the number of facets drops more steeply than their average size does.
... Insect material from the Swedish Malaise Trap Project (SMTP) was examined in search for specimens Fairyflies (Mymaridae) are among the smallest winged insects (Huber & Noyes 2013). The body size of the imago varies from 0.13-6.0 ...
... The body size of the imago varies from 0.13-6.0 mm (Gibson 1997, Huber & Noyes 2013, Pricop 2013, Huber 2017. The family has a worldwide distribution and contains around 100 genera and 1400 nominal extant species (Huber & Greenwalt 2011). ...
Article
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Fairyflies (Mymaridae) of the genera Polynema and Stephanodes collected all over Swe-den by the Swedish Malaise Trap Project (SMTP) were examined. Five Polynema species were identified as known taxa: P. (Doriclytus) euchariforme Haliday, 1833, P. (Polynema) flavipes Walker, 1848, P. (Polynema) fuscipes Haliday, 1833, P. (Polynema) gracile (Nees ab Esenbeck, 1834), and P. (Polynema) pusillum Haliday, 1833. Stephanodes similis (Foer-ster, 1847) was the only occurring species for this genus, of which a brief description of Swedish material is provided. The five species of Polynema are redescribed and, in addition , illustrated notes are given on the five unnamed morphospecies of Polynema, in order to provide information for future taxonomic revisions of this genus. A preliminary identification key to all these taxa is provided.