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Wax-producing insects. (a) Shivaphis celti, (b) Geisha distinctissima, (c) Psylla morimotoi, (d) Epicopeia hainesii hainesii. Scale bars: 10 mm. 

Wax-producing insects. (a) Shivaphis celti, (b) Geisha distinctissima, (c) Psylla morimotoi, (d) Epicopeia hainesii hainesii. Scale bars: 10 mm. 

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Many insects masquerade as parts of plants, such as bark or leaves, or mimic poisonous organisms in order to defend themselves against predators. However, recent studies indicate that plants may mimic insects and other arthropods to deter herbivores. Here, I report visually similar white structures of plants and arthropods in Japan and suggest they...

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... fi eld observations, I also found and noted di- verse wax-producing insects, including woolly aphids [e.g. Shivaphis celti Das (Fig. 2a), Colopha kansugei (Uye), and Colophina clematis (Shinji)], scale insects [e.g. Orthezia yasushii Kuwana, Takahashia japonica (Cockerell) and Ceroplastes ceriferus (Fabricius)], fl atid nymphs [Geisha distinctissima (Walker)] (Fig. 2b), nymphs of jumping plant lice (e.g. Anomoneura mori Schwarz, Psylla mori- motoi Miyatake) (Fig. 2c), ...
Context 2
... I also found and noted di- verse wax-producing insects, including woolly aphids [e.g. Shivaphis celti Das (Fig. 2a), Colopha kansugei (Uye), and Colophina clematis (Shinji)], scale insects [e.g. Orthezia yasushii Kuwana, Takahashia japonica (Cockerell) and Ceroplastes ceriferus (Fabricius)], fl atid nymphs [Geisha distinctissima (Walker)] (Fig. 2b), nymphs of jumping plant lice (e.g. Anomoneura mori Schwarz, Psylla mori- motoi Miyatake) (Fig. 2c), delphacid nymphs [Saccha- rosydne procerus (Matsumura)], sawfl y larvae (Eriocampa mitsukurii Rohwer, E. kurumivora Togashi), lepidopteran caterpillars [e.g. Epicopeia hainesii hainesii Holland (Fig. 2d), Psychostrophia melanargia ...
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... celti Das (Fig. 2a), Colopha kansugei (Uye), and Colophina clematis (Shinji)], scale insects [e.g. Orthezia yasushii Kuwana, Takahashia japonica (Cockerell) and Ceroplastes ceriferus (Fabricius)], fl atid nymphs [Geisha distinctissima (Walker)] (Fig. 2b), nymphs of jumping plant lice (e.g. Anomoneura mori Schwarz, Psylla mori- motoi Miyatake) (Fig. 2c), delphacid nymphs [Saccha- rosydne procerus (Matsumura)], sawfl y larvae (Eriocampa mitsukurii Rohwer, E. kurumivora Togashi), lepidopteran caterpillars [e.g. Epicopeia hainesii hainesii Holland (Fig. 2d), Psychostrophia melanargia Butler, Epipomponia nawai (Dyar), and Samia cynthia pryeri (Butler)], and ladybird larvae (Scymnus ...
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... nymphs [Geisha distinctissima (Walker)] (Fig. 2b), nymphs of jumping plant lice (e.g. Anomoneura mori Schwarz, Psylla mori- motoi Miyatake) (Fig. 2c), delphacid nymphs [Saccha- rosydne procerus (Matsumura)], sawfl y larvae (Eriocampa mitsukurii Rohwer, E. kurumivora Togashi), lepidopteran caterpillars [e.g. Epicopeia hainesii hainesii Holland (Fig. 2d), Psychostrophia melanargia Butler, Epipomponia nawai (Dyar), and Samia cynthia pryeri (Butler)], and ladybird larvae (Scymnus Kugelann, Hyperaspis Redten- subepidermal air spaces (e.g. leaf mottling) (Lev-Yadun, 2006a, 2014aYamazaki & Lev-Yadun, 2015). Thus, plants most often use red, brown, black and white patterns to mimic arthropods ...
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... in lowland and mountainous areas, and were found on buds (C. yasushii), stems (C. clematis, lepi- dopteran caterpillars) or the undersides of leaves (S. celti, A. mori, P. morimotoi, Eriocampa spp., E. hainesii haine- sii). White colonies of woolly aphids and jumping plant lice on the underside of leaves looked like trichome-cov- ered leaves (Fig. 2a). Stems covered with wax-producing fl atid nymphs, which resembled white plant stems, were abundant in spring and early summer (Fig. ...
Context 6
... of leaves (S. celti, A. mori, P. morimotoi, Eriocampa spp., E. hainesii haine- sii). White colonies of woolly aphids and jumping plant lice on the underside of leaves looked like trichome-cov- ered leaves (Fig. 2a). Stems covered with wax-producing fl atid nymphs, which resembled white plant stems, were abundant in spring and early summer (Fig. ...

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... (ii) Antimicrobial activity has been recently demonstrated by Sahayaraj et al. [60]. (iii) The foam's multiple scattering of light in all directions causes the white foam colour [1,61], which might function as a physical signal to herbivorous insects and mammals commonly avoiding such whitish structures in nature already occupied by other phytophagous species [62]. (iv) The foam reflection protects from bright light (1800-2000 lux) and UV radiation. ...
Article
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Aphrophora alni spittlebug nymphs produce a wet foam from anal excrement fluid, covering and protecting themselves against numerous impacts. Foam fluid contact angles on normal (26°) and silanized glass (37°) suggest that the foam wets various substrates, including plant and arthropod surfaces. The pull-off force depends on the hydration state and is higher the more dry the fluid. Because the foam desiccates as fast as water, predators once captured struggle to free from drying foam, becoming stickier. The present study confirms that adhesion is one of the numerous foam characteristics resulting in multifunctional effects, which promote spittlebugs' survival and render the foam a smart, biocompatible material of biological, biomimetic and biomedical interest. The sustainable ‘reuse' of large amounts of excrement for foam production and protection of the thin nymph integument suggests energetic and evolutionary advantages. Probably, that is why foam nests have evolved in different groups of organisms, such as spittlebugs, frogs and fish.
... Moreover, several types of pseudo-variegation cue regarding potential competition with pathogens and herbivores that arrived earlier, or about cannibalistic or predatory herbivores. Therefore, Lev-Yadun & Niemelä (2017) suggested that such leaves were probably the defended models (Fig. 10) that were later mimicked by plants with mutations that caused various variegated phenotypes (Fig. 11) (see also Yamazaki, 2017;Lev-Yadun, 2013a, 2019. Pseudo-variegation types induced by various pathogens and herbivores that not only consume plants, but actually also defend plants from further herbivory, should be considered the extended phenotypes of both these plants and of the inducers that in this way defend their plant habitat. ...
... The various types of proposed defensive animal mimicry by plants comprise two general types: direct animal mimicry, and mimicry of cues about animal action (Lev-Yadun, 2016b, 2017). The direct type includes mimicry of butterfly eggs (Benson et al., 1975;Gilbert, 1980;Shapiro, 1981a, b;Williams & Gilbert, 1981;de Castro et al., 2018), slugs or snails (Rothschild, 1974(Rothschild, , 1984, bees or wasps (Lev-Yadun & Ne'eman, 2012), caterpillars (Rothschild, 1974(Rothschild, , 1984Lev-Yadun & Inbar, 2002;Aviezer & Lev-Yadun, 2015;Lev-Yadun, 2015a, 2016bYamazaki, 2016;Quicke, 2017) (Fig. 16), ant columns (Lev-Yadun & Inbar, 2002;Lev-Yadun, 2009d) (Fig. 17), aphids (Lev-Yadun & Inbar, 2002;Lev-Yadun, 2016b;Lev-Yadun & Niemelä, 2017;Yamazaki, 2017), beetles (Yamazaki & Lev-Yadun, 2014), gall midges (Polte & Reinhold, 2013), arthropod wing movement (Lev-Yadun, 2013b), eye spots (Aviezer & Lev-Yadun, 2015) and snake mimicry (Aviezer & Lev-Yadun, 2015;Mwafongo et al., 2017). Animal activity cue mimicry includes release of an aphid alarm pheromone (Gibson & Pickett, 1983), variegation that looks like tunnelling (Smith, 1986;Lev-Yadun, 2003a;Lee, 2007;Campitelli et al., 2008;Soltau et al., 2009;Yamazaki, 2010), leaf margins that look as if they have suffered chewing damage (Niemelä & Tuomi, 1987;Dirzo, 2002;Quicke, 2017), trichome cover that looks like spider webs or arthropod silk (Yamazaki & Lev-Yadun, 2015) (Fig. 18), animaldung-shaped plants (Wiens, 1978), and carrion and dung odours . ...
Article
A common idea is that resisting or blocking herbivore attacks by structural, chemical and molecular means after they have commenced is the first line of plant defence. However, these are all secondary defences, operating only when all the various methods of avoiding attack have failed. The real first line of plant defence from herbivory and herbivore-transmitted pathogens is avoiding such attacks altogether. Several visual, chemical and ‘statistical’ methods (and commonly their combined effects) have been proposed to allow avoidance of herbivore attacks. The visual types are camouflage, masquerade, aposematic coloration of toxic or physically defended plants (including Müllerian/Batesian mimicry), undermining herbivorous insect camouflage, delayed greening, dazzle and trickery coloration, heterophylly that undermines host identification, leaf movements, and signalling that colorful autumn leaves are soon to be shed. The mimicry types include: herbivore damage, insects and other animals, fungal infestation, dead/dry leaves or branches, animal droppings, and stones and soil. Olfactory-based tactics include odour aposematism by poisonous plants, various repelling volatiles, mimicry of faeces and carrion odours, and mimicry of aphid alarm pheromones. The ‘statistical’ methods are mast fruiting, flowering only once in many years and being rare. In addition to the theoretical aspects, understanding these mechanisms may have considerable potential for agricultural or forestry applications.
... Wax production is a common defensive strategy used by plants [1], herbivores [2,3] and predators [4,5] to reduce their susceptibility to their natural enemies. Furthermore, predators may indirectly appropriate wax from other producers. ...
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Background: Larvae of the minute aphidophagous Scymnus nubilus Mulsant (Coleoptera: Coccinellidae) are common predators in apple orchards, covered by a wax layer that might act as a defense mechanism against natural enemies. However, the costs and benefits of protection conferred by wax remain to be assessed. We tested the following hypothesis: there is a trade-off in wax producing ladybeetles between the protection conferred by wax cover and the physiological or behavioral costs associated with its production. We predict that: (1) wax production is an efficient defensive mechanism (against intraguild predation), (2) wax production is associated with detrimental physiological (growth, reproduction) or behavioral effects (behavioral compensation: increased biomass consumption). Results: Tests were carried out in the laboratory with wax and waxless larvae of S. nubilus, with and without lacewing larvae of Chrysoperla agilis (Neuroptera: Chrysopidae) being used as a potential intraguild predator of the coccinellid. Waxless individuals were more susceptible to intraguild predation by lacewing larvae. Adults originating from waxless larvae were lighter than the ones originating from wax larvae, suggesting a metabolic cost resulting from a constant need of wax production. Body-weight gain and conversion efficiency were lower in waxless larvae. Biomass consumption was similar, showing that waxless larvae did not compensate for the physiological cost by eating more aphid biomass. Conclusion: The results indicate the potential existence of a trade-off between growth and protection associated with wax production.
... Defensive visual Batesian animal mimicry by plants, which was recently reviewed by Lev-Yadun (2016 and by Quicke (2017), is therefore given here only in brief, exists in several forms: (1) butterfly egg mimicry (Benson et al. 1975;Gilbert 1980Gilbert , 1982Shapiro 1981a, b;Williams and Gilbert 1981;Lev-Yadun 2016de Castro et al. 2018), (2) ant mimicry (Lev-Yadun and Inbar 2002;Lev-Yadun 2009d;Mwafongo et al. 2017;de Castro et al. 2018), (3) aphid mimicry (Lev-Yadun and Inbar 2002;Yamazaki 2017), (4) caterpillar mimicry (Rothschild 1974(Rothschild , 1984Benson et al. 1975;Lev-Yadun and Inbar 2002;Aviezer and Lev-Yadun 2015;Lev-Yadun 2015c;Yamazaki 2016Yamazaki , 2017 (Fig. 3), (5) beetle mimicry (Yamazaki and Lev-Yadun 2014), (6) spider web mimicry (Yamazaki and Lev-Yadun 2015;Yamazaki 2017) (Supplementary Fig. S7), (7) snake mimicry (Aviezer and Lev-Yadun 2015;Mwafongo et al. 2017), (8) arthropod wing movement mimicry (Lev-Yadun 2013b), (9) bee and wasp mimicry (Lev-Yadun and Ne'eman 2012), (10) animal chewing damage mimicry (Niemelä and Tuomi 1987;Brown and Lawton 1991;Rivero-Lynch et al. 1996;Dirzo 2002;Lev-Yadun 2016Quicke 2017) or tunneling damage mimicry (Smith 1986;Lev-Yadun 2003b, 2015aSoltau et al. 2009;Yamazaki 2010), (11) gall midge mimicry (Polte and Reinhold 2013), (12) eye spot mimicry (Aviezer and Lev-Yadun 2015), (13) Bird droppings (Pannell and Farmer 2016), and (14) a nonvisual mimicry of feces and carrion odor mimicry (Lev-Yadun et al. 2009b) (Fig. 4). When I saw it I thought for several seconds that it was a caterpillar. ...
... Defensive visual Batesian animal mimicry by plants, which was recently reviewed by Lev-Yadun (2016 and by Quicke (2017), is therefore given here only in brief, exists in several forms: (1) butterfly egg mimicry (Benson et al. 1975;Gilbert 1980Gilbert , 1982Shapiro 1981a, b;Williams and Gilbert 1981;Lev-Yadun 2016de Castro et al. 2018), (2) ant mimicry (Lev-Yadun and Inbar 2002;Lev-Yadun 2009d;Mwafongo et al. 2017;de Castro et al. 2018), (3) aphid mimicry (Lev-Yadun and Inbar 2002;Yamazaki 2017), (4) caterpillar mimicry (Rothschild 1974(Rothschild , 1984Benson et al. 1975;Lev-Yadun and Inbar 2002;Aviezer and Lev-Yadun 2015;Lev-Yadun 2015c;Yamazaki 2016Yamazaki , 2017 (Fig. 3), (5) beetle mimicry (Yamazaki and Lev-Yadun 2014), (6) spider web mimicry (Yamazaki and Lev-Yadun 2015;Yamazaki 2017) (Supplementary Fig. S7), (7) snake mimicry (Aviezer and Lev-Yadun 2015;Mwafongo et al. 2017), (8) arthropod wing movement mimicry (Lev-Yadun 2013b), (9) bee and wasp mimicry (Lev-Yadun and Ne'eman 2012), (10) animal chewing damage mimicry (Niemelä and Tuomi 1987;Brown and Lawton 1991;Rivero-Lynch et al. 1996;Dirzo 2002;Lev-Yadun 2016Quicke 2017) or tunneling damage mimicry (Smith 1986;Lev-Yadun 2003b, 2015aSoltau et al. 2009;Yamazaki 2010), (11) gall midge mimicry (Polte and Reinhold 2013), (12) eye spot mimicry (Aviezer and Lev-Yadun 2015), (13) Bird droppings (Pannell and Farmer 2016), and (14) a nonvisual mimicry of feces and carrion odor mimicry (Lev-Yadun et al. 2009b) (Fig. 4). When I saw it I thought for several seconds that it was a caterpillar. ...
... Defensive visual Batesian animal mimicry by plants, which was recently reviewed by Lev-Yadun (2016 and by Quicke (2017), is therefore given here only in brief, exists in several forms: (1) butterfly egg mimicry (Benson et al. 1975;Gilbert 1980Gilbert , 1982Shapiro 1981a, b;Williams and Gilbert 1981;Lev-Yadun 2016de Castro et al. 2018), (2) ant mimicry (Lev-Yadun and Inbar 2002;Lev-Yadun 2009d;Mwafongo et al. 2017;de Castro et al. 2018), (3) aphid mimicry (Lev-Yadun and Inbar 2002;Yamazaki 2017), (4) caterpillar mimicry (Rothschild 1974(Rothschild , 1984Benson et al. 1975;Lev-Yadun and Inbar 2002;Aviezer and Lev-Yadun 2015;Lev-Yadun 2015c;Yamazaki 2016Yamazaki , 2017 (Fig. 3), (5) beetle mimicry (Yamazaki and Lev-Yadun 2014), (6) spider web mimicry (Yamazaki and Lev-Yadun 2015;Yamazaki 2017) (Supplementary Fig. S7), (7) snake mimicry (Aviezer and Lev-Yadun 2015;Mwafongo et al. 2017), (8) arthropod wing movement mimicry (Lev-Yadun 2013b), (9) bee and wasp mimicry (Lev-Yadun and Ne'eman 2012), (10) animal chewing damage mimicry (Niemelä and Tuomi 1987;Brown and Lawton 1991;Rivero-Lynch et al. 1996;Dirzo 2002;Lev-Yadun 2016Quicke 2017) or tunneling damage mimicry (Smith 1986;Lev-Yadun 2003b, 2015aSoltau et al. 2009;Yamazaki 2010), (11) gall midge mimicry (Polte and Reinhold 2013), (12) eye spot mimicry (Aviezer and Lev-Yadun 2015), (13) Bird droppings (Pannell and Farmer 2016), and (14) a nonvisual mimicry of feces and carrion odor mimicry (Lev-Yadun et al. 2009b) (Fig. 4). When I saw it I thought for several seconds that it was a caterpillar. ...
Article
Several types of defensive Batesian mimicry seem to be much more common in plants than was historically and is currently considered. It is based either on visual aspects (shape, coloration, and even movement), on odors, and on combinations of both these sensing modalities. Various characters that seem to function as defensive Batesian mimicry, may also simultaneously take part in pollination, physiological functions, or in other defensive mechanisms. The defended models for the visual Batesian mimics in plants belong to several categories: (1) spiny, thorny and prickly plant species, (2) mechanically or chemically defended parts of the same individual plant, or other members of the same species (auto mimicry), (3) colorful and chemically defended plants, (4) dangerous animals (aggressive, toxic), (5) fungal attacks, (6) animal action and animal damage cues, and (7) oozing defensive white latex. Olfactory defended models include: (1) toxic plants, (2) animal alarm pheromones, and (3) animal carrion and feces odors. Many more descriptive, genetic, phylogenetic and experimental studies have to be done in order to better understand the role of defensive Batesian mimicry in plant biology.
... However, as spittlebugs do not consume leaf tissue, this explanation should not apply to them. Yamazaki (2017) suggested that the foam's white colour might function as a signal to herbivorous insects and mammals to avoid plants attacked recently. He claimed that the colour 'white' is indicative of reduced plant quality to herbivores. ...
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1. Biofoam in spittlebugs has traditionally been seen as a defence against predation and a microclimate that reduces the risks of overheating and drying out. This study addresses the possible role of the foam as a light attenuator. 2. Nymphs exhibit higher mortalities when reared under brighter light (1800–2000 lux) than under less bright light (600–800 lux). At all developmental stages, photoavoidance is strongest when the nymphs are depleted of foam covers. First‐ and second‐instar nymphs appear to be the most vulnerable to exposures by bright light. 3. Smaller bubbles are more effective as light attenuators than are larger ones. As younger instars possess smaller canals from which bubbles are released and, furthermore, exhibit higher concentrations of proteins in bubble liquid, they can produce smaller‐sized bubbles and their foams are more effective at reducing light than are those of older nymphs. 4. The findings of this study show that most of the visible and UV radiation is reflected by the foam: transmittance of visible light was 15% at 600 nm and 12% at 350 nm. 5. These results demonstrate that spittlebug foam also possesses properties that render it an effective barrier against potentially damaging solar radiation.
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Resprouting is a functional trait in species which occur in fi re-prone ecosystems. These plants can resprout from aerial buds and by recruiting belowground bud bank using carbohydrates allocated in roots as resource. In this study, we present morpho-anatomical features and chemical composition related to the resprouting potential of two species of Eugenia L. in an area of the Cerrado (Brazilian savanna) under regeneration, after the clear-cutting of Pinus sp. with the later burning of pine needles layer. We used standard histological techniques for belowground organs analysis and aerial buds protection degree. Belowground buds in layer from soil surface down to 10 depth were counted and the chemical analyses were performed on roots. In all aerial buds, there were relevant protection traits. The belowground organ is a sobole and the number of buds in its upper portion varied from 24 to 517 between individuals of both species. Phenolic compounds, fl avonoids, starch and other carbohydrates were detected in roots. The protection of aerial buds, the large number of belowground buds and the storing and protective compounds may have favored the resprouting of the species in the area.