Hymenoptera and biomimetic surfaces: insights and innovations

• Vinicius Marques Lopez,
• Carlo Polidori and
• Rhainer Guillermo Ferreira

Beilstein J. Nanotechnol. 2024, 15, 1333–1352, doi:10.3762/bjnano.15.107

• development of self-cleaning surfaces and materials resistant to biofouling in marine environments. Legs Adhesive pads: The adhesive organ in Hymenoptera consists of a flexible cuticle pad (i.e., arolium) situated between the pretarsal claws, capable of unfolding and retracting with each step [125] (Figure 6A

Functional morphology of cleaning devices in the damselfly Ischnura elegans (Odonata, Coenagrionidae)

• Silvana Piersanti,
• Gianandrea Salerno,
• Wencke Krings,
• Stanislav Gorb and
• Manuela Rebora

Beilstein J. Nanotechnol. 2024, 15, 1260–1272, doi:10.3762/bjnano.15.102

• microscope. (a) Ventral view of tibia (Ti) and tarsus (Ta) showing the tibial grooming devices (arrows) located on the medial side of the tibia. C, claws. (b) Detail of a grooming device emerging from a well-developed socket (S). (c) Detail of the grooming devices observed from the ventral side of the leg

The effect of age on the attachment ability of stick insects (Phasmatodea)

• Marie Grote,
• Stanislav N. Gorb and
• Thies H. Büscher

Beilstein J. Nanotechnol. 2024, 15, 867–883, doi:10.3762/bjnano.15.72

• the substrate as the contact area of the pad is proportional to adhesion [10][11][12]. For rough substrates, the pads are complemented by a pair of rigid claws used for friction interlocking with surface asperities and ensuring attachment, but claws perform poorly on smooth surfaces [13]. The ability

• investigated various functions of this system and how it changes under certain conditions, such as substrate geometry [36], the presence or absence of claws [37], different surface characteristics of substrates [33][38][39], and the combined effect with pad fluids [40]. For these animals, whose lives strongly

• major influences of the claws [33][37]. On smooth surfaces (0 µm), smooth pads generate proper contact with the surface. Microrough surfaces interfere with the contact formation of smooth pads; however, the dimension of the nubs on the euplantulae yield different responses to the roughness because fine

Comparative analysis of the ultrastructure and adhesive secretion pathways of different smooth attachment pads of the stick insect Medauroidea extradentata (Phasmatodea)

• Julian Thomas,
• Stanislav N. Gorb and
• Thies H. Büscher

Beilstein J. Nanotechnol. 2024, 15, 612–630, doi:10.3762/bjnano.15.52

• ) at their distal ends. The pretarsus features the arolium (ar) situated between a pair of claws (cl). The euplantulae, the cuticle between them, and the arolium bear a rather smooth surface structure. The remaining surface of the tarsomeres, where no attachment pads are situated, is covered with setae

Insect attachment on waxy plant surfaces: the effect of pad contamination by different waxes

• Elena V. Gorb and
• Stanislav N. Gorb

Beilstein J. Nanotechnol. 2024, 15, 385–395, doi:10.3762/bjnano.15.35

• two distally located claws and adhesive pads situated on the ventral side of three (out of five) proximal tarsomeres (later referred to as basal, middle, and distal) (Figure 2a,b). In common with most beetles from the family Chrysomelidae [37], this species has hairy tarsal adhesive pads (according to

Growing up in a rough world: scaling of frictional adhesion and morphology of the Tokay gecko (Gekko gecko)

• Anthony J. Cobos and
• Timothy E. Higham

Beilstein J. Nanotechnol. 2022, 13, 1292–1302, doi:10.3762/bjnano.13.107

• Animals attach to surfaces in numerous ways, including claws, suction, and both wet and dry adhesion. In fact, some animals can utilize multiple attachment mechanisms [1][2], leading to multifunctionality across surfaces of varying roughness. Dry adhesion is found in many invertebrates and squamate

Effect of sample treatment on the elastic modulus of locust cuticle obtained by nanoindentation

• Chuchu Li,
• Stanislav N. Gorb and
• Hamed Rajabi

Beilstein J. Nanotechnol. 2022, 13, 404–410, doi:10.3762/bjnano.13.33

• : biomimetics; cuticle; locust; material properties; mechanical testing; nanoindentation; water content; Introduction Cuticle is a lightweight material that forms the whole exoskeleton of insects, from the flexible intersegmental membrane to the stiff jaws and claws. Cuticle of each insect body part has

Physical constraints lead to parallel evolution of micro- and nanostructures of animal adhesive pads: a review

• Thies H. Büscher and
• Stanislav N. Gorb

Beilstein J. Nanotechnol. 2021, 12, 725–743, doi:10.3762/bjnano.12.57

• (Figure 4) consists of five tarsomeres. It is equipped with two claws and an arolium on the pretarsus, as well as euplantulae on the proximal four to five tarsomeres [1][2][109][161][162][225]. Except for the euplantulae of some Aschiphasmatini (Aschiphasmatinae) that are covered with adhesive setae [109

• adhesive properties in their respective environments. Consequently, the convergent presence of the same AMS is primarily a result of the same environmental condition found and to establish the necessary functional principle. Functional principles Studies on different groups of insects have shown that claws

• generally contribute to the attachment on rough surfaces due to friction and mechanical interlocking [83][89][92][232][233][234][235][236]. The performance of claws depends on the radius of the claw tip in relation to the curvature of the surface irregularities [83][234][237][238]. However, in combination

A comparison of tarsal morphology and traction force in the two burying beetles Nicrophorus nepalensis and Nicrophorus vespilloides (Coleoptera, Silphidae)

• Liesa Schnee,
• Benjamin Sampalla,
• Josef K. Müller and
• Oliver Betz

Beilstein J. Nanotechnol. 2019, 10, 47–61, doi:10.3762/bjnano.10.5

• , Hauptstr.1, 79104 Freiburg, Germany 10.3762/bjnano.10.5 Abstract Our aim was to compare friction and traction forces between two burying beetle species of the genus Nicrophorus exhibiting different attachment abilities during climbing. Specifically, the interaction of adhesive hairs and claws during

• detected between males and females within each species. With claws intact, both species showed the highest forces on rough surfaces, although N. nepalensis with clipped claws performed best on a smooth surface. However, N. nepalensis beetles outperformed N. vespilloides, which showed no differences between

• smooth and rough surfaces with clipped claws. Both species demonstrated poor traction forces on micro-rough surfaces. Results concerning the impact of surface polarity were inconclusive, whereas roughness more strongly affected the attachment performance in both species. Nanotribometric analyses of the

When the going gets rough – studying the effect of surface roughness on the adhesive abilities of tree frogs

• Niall Crawford,
• Thomas Endlein,
• Jonathan T. Pham,
• Mathis Riehle and
• W. Jon P. Barnes

Beilstein J. Nanotechnol. 2016, 7, 2116–2131, doi:10.3762/bjnano.7.201

• possess claws as well as adhesive pads. Additionally, there are studies of plant surfaces that have evolved to be anti-adhesive as far as insects are concerned. The effects of surface roughness on animals with hairy pads (geckos, spiders, insects such as beetles) are reasonably predictable. When the

• rough surfaces. Where insufficient bending occurs (see Figure 8C), air bubbles are likely to be formed, with a consequent reduction in adhesion. Many climbing organisms utilise claws to climb on rough surfaces, which can interlock with asperities on vertical surfaces – this is seen in geckos [40

• ], spiders [41] and many insects [42][43]. However the effectiveness of a claw is usually dependent on the asperity size being larger than the claw tip diameter [44]. When the claws fail to interlock on the surface, staying attached relies on the adhesive pads of the organism [45]. On the basis of tests with

Influence of ambient humidity on the attachment ability of ladybird beetles (Coccinella septempunctata)

• Lars Heepe,
• Jonas O. Wolff and
• Stanislav N. Gorb

Beilstein J. Nanotechnol. 2016, 7, 1322–1329, doi:10.3762/bjnano.7.123

• hairy attachment devices of C. septempunctata. The tarsus is composed of three tarsomeres and two ventrally curved claws (Figure 1B–D,H–J). Only the first two tarsomeres (T1 and T2 in Figure 1) are ventrally covered by tenent setae. Different types of tarsal adhesive setae were distinguished: (1) setae

• ) in males. Tarsi of males were also ventrally covered by tenet setae types shown in (E, F), but have an additional type, which is terminated with discoidal terminal elements (K). CW, claws; T1, first proximal tarsomer; T2, second proximal tarsomer; T3, third proximal tarsomer. The arrows in (B–D) and

Functional diversity of resilin in Arthropoda

• Jan Michels,
• Esther Appel and
• Stanislav N. Gorb

Beilstein J. Nanotechnol. 2016, 7, 1241–1259, doi:10.3762/bjnano.7.115

• structures connecting claws and pulvilli to the terminal tarsomere [43][50]. In the pretarsus of the drone fly (Eristalis tenax) (Insecta, Diptera, Syrphidae), for example, membranous cuticle with large proportions of resilin forms a spring-like (or joint-like) element (Figure 3A–C) that makes the pulvilli

Aquatic versus terrestrial attachment: Water makes a difference

• Petra Ditsche and
• Adam P. Summers

Beilstein J. Nanotechnol. 2014, 5, 2424–2439, doi:10.3762/bjnano.5.252

• water make it harder to bring surfaces into close apposition. Most arthropods living in flowing water have well-developed tarsal claws, with which they hold onto rough surfaces [56]. These claws show a variety of different shapes and sizes (Figure 10) and are the most common attachment devices of

• aquatic macroinvertebrates in both running and still water [78]. The larvae of some taxa, such as mayflies and caddis larvae usually bear one claw at their tarsi, while many others like stoneflies or several aquatic beetles have two tarsal claws (Figure 10). Double claws might act in the same direction or

• in accordance to the clamp principle (or something intermediate). Free-living caddis larvae like Rhyacophila have additional claws like grapples on their posterior prolegs. Circlets of outwardly directed hooks imply the spacer principle. They occur on the prolegs in larvae of several Diptera taxa

Physical principles of fluid-mediated insect attachment - Shouldn’t insects slip?

• Jan-Henning Dirks

Beilstein J. Nanotechnol. 2014, 5, 1160–1166, doi:10.3762/bjnano.5.127

• of claws is limited to compliant surfaces in which the claws can insert, or rough surfaces with asperities larger than the diameter of the claw tip [15]. Hence, to stick to smooth and stiff natural substrates, such as stones or leaves, insects and other arthropods have to use adhesive pads (Figure 1

Insect attachment on crystalline bioinspired wax surfaces formed by alkanes of varying chain lengths

• Elena Gorb,
• Sandro Böhm,
• Nadine Jacky,
• Louis-Philippe Maier,
• Kirstin Dening,
• Sasha Pechook,
• Boaz Pokroy and
• Stanislav Gorb

Beilstein J. Nanotechnol. 2014, 5, 1031–1041, doi:10.3762/bjnano.5.116

• , insects use different structures for attachment, depending on the texture of the substrate. They usually apply their claws to interlock with surface irregularities on rough surfaces, when the diameter of the claw tip is smaller than the dimensions of typical surface asperities or cavities [1]. On smooth

• some types of trichomes, acting mainly at the macroscopic level, hinder the interlocking of insect claws. Additionally, plant-produced wet films on the surface, microscopic cuticular folds and epicuticular (deposited onto the plant cuticle) wax crystals reduce the adhesion of insect attachment pads

• ventrally curved claws with a claw tip diameter of about 4 μm [42] and hairy adhesive pads situated on the ventral side of the two first proximal tarsomeres (Figure 5a). Pads are covered with numerous tiny setae having various tip shapes, from sharp-pointed to spatula-like, ranging in width from ca. 1.8 to

Impact of cell shape in hierarchically structured plant surfaces on the attachment of male Colorado potato beetles (Leptinotarsa decemlineata)

• Bettina Prüm,
• Robin Seidel,
• Holger Florian Bohn and
• Thomas Speck

Beilstein J. Nanotechnol. 2012, 3, 57–64, doi:10.3762/bjnano.3.7

• cuticular folds. For attachment of L. decemlineata, as for many other insects, both adhesive pads and claws are responsible (Figure 3). The hairy adhesive pads of beetles show best attachment on smooth surfaces, or on surfaces with very large diameters of the asperities, as shown in experiments with insects

• having dissected claws [13][14][15][16][17]. For various insects it has been shown that roughness induced by wax crystals [7][18][19][20][21][22][23] or cuticular folds [8] strongly reduces attachment ability. Different mechanisms have been hypothesised [20][24], but the roughness hypothesis is most

• , claws generally improve grip [14][15] depending on the dimensions of the surface asperities and the insect’s claws [15][16]. If the diameter of the claw tip is smaller than the surface roughness, the claws can hook into the surface irregularities and the beetle thereby increases attachment forces. As in

The effect of surface anisotropy in the slippery zone of Nepenthes alata pitchers on beetle attachment

• Elena V. Gorb and
• Stanislav N. Gorb

Beilstein J. Nanotechnol. 2011, 2, 302–310, doi:10.3762/bjnano.2.35

• to the upward one, whereas clawless insects did not. These results led to the conclusion that, (i) due to the particular shape of lunate cells, the pitcher surface has anisotropic properties in terms of insect attachment, and (ii) claws were mainly responsible for attachment enhancement in the

• downward pitcher direction, since, in this direction, they could interlock with overhanging edges of lunate cells. Keywords: adhesive pads; claws; Coccinella septempunctata; insect–plant interactions; traction force; Introduction Pitcher-shaped trapping organs produced at the tips of tendrils are

• contact area caused by surface micro-roughness [19][22][25][26]. Also, due to the fragile and brittle nature of wax crystals and their small dimensions, insects cannot apply their claws for interlocking with crystals in order to climb up the pitcher wall [22][25][26]. As for the potential role of lunate