Effect of foliar tomentosity on phenotypic plasticity in Bemisia tabaci (Hom., Aleyrodidae)

Some whitefly species, as well as members of other insect families, are known to develop different phenotypes under different environmental conditions. In this study, the identification of an environmental trigger and its sensitive stage for the development of the setose phenotype of Bemisia tabaci nymphs are reported. Quantification tests revealed phenotypic plasticity manifested by positive correlation between trichome density on tomentose leaves and the percentage of setose nymphs found upon them. Artificial simulation of leaf tomentosity provided sufficient stimulation to induce the setose nymph phenotype. The crawler was found to be the only sensitive stage for this induction. Polymorphism of the nymphal phenotypes correlated with additional biological features of both nymph and adult, which could be treated as proxies of fitness. Setose nymphs were smaller and developed (from eggs to adult) faster. Adults emerging from such nymphs were smaller and had shorter lives. Adaptivity of the detected phenotypic plasticity and of the correlating nymphal and adult features are discussed.     

Effects of plant and prey characteristics on the predatory behavior of Delphastus catalinae

Nymphal setosity of the whitefly Bemisia tabaci Gennadius (Homoptera: Aleyrodidae) has been reported to be induced by mechanical stimuli such as leaf tomentosity, and related to the predatory performance of the coccinellid Delphastus catalinae (Horn) (Coleoptera: Coccinellidae). In this study, a possible adaptive value of this phenomenon for the whitefly is shown through the combined effects of leaf and prey characteristics on the walking and predatory behaviors of the beetle. Leaf tomentosity significantly affected the walking patterns of the beetle and therefore its searching abilities, thus indirectly increasing the influence of prey distribution upon predator's efficiency. Moreover, while searching on tomentose leaves, the beetles showed preference for the smooth prey phenotype. This behavior was found dependent on the experience of the beetle in previous encounters. These results are pertinent to intraspecific competition between the different nymphal phenotypes and to the predatory efficiency of this beetle, which is utilized in biological control of whiteflies.     

Frequency-dependent herbivory by a leaf beetle,Phaedon brassicae,on hairy and glabrous plants of Arabidopsis halleri subsp. gemmifera

Frequency-dependent prey choice by natural enemies may influence the coexistence of multiple prey types, but little is known about whether frequency-dependent foraging choice occurs in herbivory on plants showing resistance polymorphism within a single population. Here we examined frequency-dependent foraging by a crucifer-feeding leaf beetle, Phaedon brassicae, on trichome-producing (hairy) and trichomeless (glabrous) plants coexisting within a natural population of the perennial herb Arabidopsis halleri subsp. gemmifera. Larvae of P. brassicae fed on hairy leaves showed slower growth than those fed on glabrous leaves. Although adult beetles consumed similar amounts of leaves when they were fed either hairy or glabrous leaves in no-choice conditions, our choice experiment showed that adult beetles fed at less than the proportionally expected level on hairy leaves compared to glabrous leaves when the hairy leaves were less or equally abundant. Both types of leaves were consumed at the proportionally expected levels when the hairy leaves were more abundant than the glabrous leaves. In a natural population, the leaf damage on the hairy plants was negatively correlated with the local proportion of the glabrous plants in a 1-m diameter patch across 2 years, while correlations between the leaf damage on the glabrous plants and their proportion differed between the 2 years. Additionally, we found five glucosinolates in leaves of A. halleri, but their accumulation did not differ between hairy and glabrous plants. Our experimental results indicate that hairy plants incur less herbivory by P. brassicae when glabrous plants are abundant. The field pattern provides evidence suggestive of frequency-dependent herbivory acting on hairy plants. The present study highlights one of the putative mechanisms of maintaining plant resistance polymorphism.     

Plant architecture and prey distribution influence foraging behavior of the predatory mite Phytoseiulus persimilis (Acari: Phytoseiidae)

The arrangement, number, and size of plant parts may influence predator foraging behavior, either directly, by altering the rate or pattern of predator movement, or, indirectly, by affecting the distribution and abundance of prey. We report on the effects of both plant architecture and prey distribution on foraging by the predatory mite, Phytoseiulus persimilis Athias-Henriot (Acari: Phytoseiidae), on cucumber (Cucumis sativus L.). Plants differed in leaf number (2- or 6-leafed), and there were associated differences in leaf size, plant height, and relative proportions of plant parts; but all had the same total surface area. The prey, the twospotted spider mite Tetranychus urticae Koch (Acari: Tetranychidae), were distributed either on the basal leaf or on all leaves. The effect of plant architecture on predator foraging behavior varied depending on prey distribution. The dimensions of individual plant parts affected time allocated to moving and feeding, but they did not appear to influence the frequency with which predators moved among different plant parts. Overall, P. persimilis moved less, and fed upon prey longer, on 6-leafed plants with prey on all leaves than on plants representing other treatment combinations. Our findings suggest that both plant architecture and pattern of prey distribution should be considered, along with other factors such as herbivore-induced plant volatiles, in augmentative biological control programs.     

Climbing of Leaf Trichomes by Eriophyid Mites Impedes Their Location by Predators

Eriophyid mites are plant parasites that are well adapted to hide away from predators. Tiny and wormlike, they can invade very narrow spaces in plants or form galls that, apart from other functions, serve as a shelter from predation. Previous observations showed that some free-living eriophyids as well as tetranychid mites spend their quiescence on the top of leaf trichomes. Here, I investigated climbing leaf trichomes by the eriophyid, Rhinophytoptus concinnus, and tested whether it enables the herbivores to avoid phytoseiid mites. Climbing behavior took place just prior to the quiescent period of juveniles. Larvae and nymphs raised the hind part of their stiffening bodies and walked, turning around on their axis. Having found a hair, juveniles attached their anal suckers to its tip, and, pushing back from a leaf surface or the base of the hair, they lifted their bodies up to become motionless. As revealed by the playback experiments with the phytoseiid mite, Typhloctonus tiliarum, predatory females needed much more time to find quiescent nymphs perching on hairs than those placed on a leaf surface. The time of nymph handling was similar in both situations. Also, a similar number of predators gave up feeding on nymphs in both locations. I conclude that climbing leaf trichomes enables the herbivorous mite to hide from predators. After detection, however, placement on trichomes does not give the quiescent juveniles any advantage over those placed on a leaf blade.     

Effects of the presence of a predatory fish and the phenotype of its prey (a shredding shrimp) on leaf litter decomposition

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光柱长鳞杜鹃——四川杜鹃花属一新变种

描述了四川杜鹃花属一新变种——光柱长鳞杜鹃 Rhododendron　longesquamatumSchneid. var. glabristylum　Y. Y. Geng et Z. L. Zhao。该变种与原变种长鳞杜鹃R. longesquamatum　Schneid. var. longesquamatum的主要区别是叶片和花较小,叶片及叶脉成熟后无毛或仅见零星残存毛,花萼仅有绒毛而无腺毛,花柱光滑。     

Leaf pubescence mediates intraguild predation between predatory mites

Plant morphological traits such as leaf pubescence may affect herbivores and their natural enemies at the individual, population and community levels. Leaf pubescence has been repeatedly shown to mediate predator‐herbivore interactions whereas the influence of leaf pubescence on predator–predator interactions such as intraguild predation (IGP) has seldom been investigated. Using a three‐pronged approach we assessed the influence of leaf pubescence on the predatory mites Kampimodromus aberrans and Euseius finlandicus. Both predators occur on broad‐leaved trees in Europe. Euseius finlandicus is mostly found on trees with glabrous leaves whereas K. aberrans mainly occurs on trees with pubescent leaves. We hypothesized that leaf pubescence mediates IGP between K. aberrans and E. finlandicus and thereby determines their dominance and proportional abundance. A field survey on apple revealed that the abundance of K. aberrans and E. finlandicus is negatively correlated, with the former predominating on cultivars with strongly pubescent leaves and the latter predominating on cultivars with little pubescent or glabrous leaves. Microhabitat choice tests showed that K. aberrans preferentially resides on pubescent leaves whereas E. finlandicus preferentially resides on glabrous leaves. The effects of leaf pubescence on survival and development of immature IG predators and IG prey were reversed for K. aberrans and E. finlandicus. In the presence of the IG predator E. finlandicus, immature K. aberrans had higher survival probabilities on pubescent leaves than on glabrous ones. In contrast, the survival chances of immature E. finlandicus were higher on glabrous leaves than on pubescent ones when the IG predator K. aberrans was present. Artificial leaf pubescence enhanced IG prey capture by immature K. aberrans and prolonged their longevity but impaired IG prey capture by immature E. finlandicus and shortened their longevity. We conclude that leaf pubescence mediates IGP strength and symmetry and discuss the implications to natural and biological control.     

Dimorphism in trichome production of <Emphasis Type="Italic">Persicaria lapathifolia</Emphasis> var. <Emphasis Type="Italic">lapathifolia</Emphasis> and Its multiple effects on a leaf beetle

Polymorphisms in plants are main factors that determine the diversity of associated animal communities and their population dynamics. Typically, Persicaria lapathifolia var. lapathifolia (Polygonaceae) has no trichomes on leaf surfaces (glabrous type), but a hairy type does sometimes occur. Based on a cultivation experiment, the presence or absence of trichomes is clarified to be under genetic control. To reveal the defensive function of trichomes against herbivores, laboratory experiments were conducted using a major herbivore, Galerucella grisescens (Coleoptera: Chrysomelidae). In both choice and no-choice feeding tests, the glabrous type was significantly more consumed by G. grisescens adults, while the hairy type was not consumed. In the hairy leaf treatment, larval duration tended to become longer, the adult body weight became significantly lower, and adults laid significantly more eggs than in the glabrous leaf treatment. Hairy leaves contained significantly more total phenolics and condensed tannins than glabrous leaves, suggesting that the hairy type allocates more resources for physical and chemical defence. Because no significant differences in leaf consumption were detected in the feeding experiment using powdered host leaves, G. grisescens seems to have adapted to the chemical defences of P. lapathifolia var. lapathifolia. These results clearly indicate that leaf trichomes of P. lapathifolia var. lapathifolia effectively act as a physical defence against G. grisescens.     

Correspondence of trichome mutations in diploid and tetraploid cottons

Quantitative variation for leaf trichome number is observed within and among Gossypium species, varying from glabrous to densely pubescent phenotypes. Moreover, economically important cotton lint fibers are modified trichomes. Earlier studies have mapped quantitative trait loci (QTLs) affecting leaf pubescence in Gossypium using allotetraploids. In this study, we mapped genes responsible for leaf trichome density in a diploid A genome cross. We were able to map 3 QTLs affecting leaf pubescence based on trichome counts obtained from young leaves (YL) and mature leaves (ML). When the F(2) progeny were classified as pubescent versus glabrous, their ratio did not deviate significantly from a 3:1 model, suggesting that glabrousness is inherited in a simple Mendelian fashion. The glabrous mutation mapped to linkage group A3 at the position of major QTL YL1 and ML1 and appeared orthologous to the t1 locus of the allotetraploids. Interestingly, a fiber mutation, sma-4(ha), observed in the same F(2) population cosegregated with the glabrous marker, which indicates either close linkage or common genetic control of lint fiber and leaf trichomes. Studies of A genome diploids may help to clarify the genetic control of trichomes and fiber in both diploid and tetraploid cottons.     

Interaction of bean leaf pubescence with rust urediniospore deposition and subsequent infection density

The role of dense straight trichomes on bean leaves associated with race-nonspecific resistance to rust was studied using a settling tower method for uniform inoculation. Abaxial leaf trichomes, which occur only on adult leaves, trapped significant numbers of urediniospores, especially those deposited in clusters, and resulted in reduced effective inoculum reaching the leaf surface. In the field where infection conditions are often less than optimum, leaf pubescence would partially explain the observed resistance. However, the observed increase in total number of spores as well as similar infection densities on fully expanded leaves of glabrous and pubescent genotypes cannot be explained by the trichomes. Other physical or chemical features of the leaf surface such as waxes or cations may play a role. The ecological fit of leaf pubescence is discussed.     

Maximizing feeding efficiency and minimizing time exposed to predators: a trade-off in the black-capped chickadee

Summary Animals often must feed away from protective cover, sometimes at a considerable risk of being preyed upon. Feeding at the maximum rate while away from cover may simultaneously minimize the time spent exposed to predators, but this is not always the case. Under some circumstances, carrying prey items to protective cover before they are consumed will minimize the time spent exposed to predators, whereas feeding at maximum efficiency (staying to eat prey where they are found) will actually increase the time spent exposed to predators. Whether or not there is a conflict between maximizing foraging efficiency and minimizing exposure time, depends upon the travel time to cover relative to the handling time of a prey item; short handling times and/or long travel times are associated with the no-conflict situation, whereas the conflict situation is associated with long handling times and/or short travel times to cover. Free-ranging chickadees foraging at an artificial patch at various distances from cover can distinguish between these two foraging situations. When there is no conflict, they stay and eat at the patch. Their behavior in the conflict situation indicates that they are tradingoff foraging considerations against the risk of predation. When the cost of carrying is low and the benefit gained is high, the chickadees elect to carry items to cover; they tend to stay and eat at the patch when the relative magnitudes of costs and benefits are reversed.     

Functional response of Neoseiulus californicus (Acari: Phytoseiidae) to Tetranychus urticae (Acari: Tetranychidae) on strawberry leaves

Neoseiulus californicus (McGregor) is a predatory mite employed for biological control of the agricultural pest Tetranychus urticae (Koch). We explored whether environmental differences, in this case the trichome densities of abaxial leaf surfaces of strawberry cultivars (‘Maehyang’ and ‘Sulhyang’ varieties) affect the functional response of adult female N. californicus preying on immature stages (egg, larva and nymph) of T. urticae. We also evaluated the functional response of N. californicus to eggs of T. urticae at different temperatures (15, 20, 25, 30 and 35°C). We conducted a logistic regression of the proportion of prey consumed as a function of initial prey density to identify functional response types, and used nonlinear least‐squares regression and the random predator equation to estimate attack rates and handling times. The functional response of adult female N. californicus to T. urticae was not influenced by non‐glandular trichomes on abaxial leaves but was affected by temperature. Overall, adult female N. californicus exhibited a type 2 functional response to T. urticae. The handling time of N. californicus was highest (1.9970 h) against T. urticae nymphs. The attack rate did not change much at 15–30°C, but was significantly higher at 35°C. The handling time decreased significantly with increasing temperature at 15–35°C. At 35°C, the attack rate was highest (0.2087) and the handling time was lowest (0.9511 h).     

Dynamics of refuge use: diurnal, vertical migration by predatory and herbivorous mites within cassava plants

Plants may protect themselves against herbivorous arthropods by providing refuges to predatory arthropods, but they cannot prevent herbivores from taking countermeasures or even from reaping the benefits. To understand whether plants benefit from providing self‐made refuges (so‐called domatia), it is not only necessary to determine the fitness consequences for the plant, but also to assess (1) against which factors the refuge provides protection, (2) why predatory arthropods are more likely to monopolise the refuge, and (3) how herbivorous and predatory arthropods respond to and affect each other in and outside the refuge. In this article, we focus on the last aspect by studying the dynamics of refuge use of a predatory mite (Typhlodromalus aripo) and its consequences for a herbivorous mite (Mononychellus tanajoa) on cassava plants in Benin, West Africa. The refuge, located in‐between the leaf primordia of the cassava apex, is thought to provide protection against abiotic factors and/or intraguild predators. To test whether the predator waits for prey in the apex or comes out, we sampled predator‐prey distributions on leaves and in the apex at 4 hour‐intervals over a period of 24 hours. The predatory mites showed pronounced diurnal changes in within‐plant distribution. They were in the apices during the day, moved to the young leaves during night and returned to the apices the next morning. Nocturnal foraging bouts were more frequent when there were more herbivorous mites on the leaves near the apex. However, the foraging predators elicited an avoidance response by mobile stages of their prey, since these were more abundant on the first 20 leaves below the apex during late afternoon, than on the same leaves during night. These field observations on cassava plants show that (1) during daytime predatory mites monopolise the apical domatia, (2) they forage on young leaves during night and (3) elicit avoidance by within‐plant, vertical migration of mobile stages of the herbivorous mites. We hypothesize that cassava plants benefit from apical domatia by acquiring protection for their photosynthetically most active, young parts, because predatory mites (1) protect primordial leaves in the apex, (2) reduce the densities of herbivorous mites on young leaves, and (3) cause herbivorous mites to move down to less profitable older leaves.     

Host plant effects on the behaviour and performance of Amblyseius swirskii (Acari: Phytoseiidae)

Biological control in ornamental crops is challenging due to the wide diversity of crops and cultivars. In this study, we tested the hypothesis that trichome density on different host plants influences the behavior and performance of the predatory mite Amblyseius swirskii Athias-Henriot (Acari: Phytoseiidae). Behavioural observations of this predator in the presence or absence of prey (western flower thrips, Frankliniella occidentalis Pergande) (Thysanoptera: Thripidae) were done on leaf squares of ornamental plant species differing in trichome density (rose, chrysanthemum and gerbera) and compared to a smooth surface (plastic). Tomato leaves were used to observe the influence of glandular trichomes. The performance of A. swirskii was assessed by measuring predation and oviposition rate. Behaviour of A. swirskii was influenced by plant species. Up to a certain density of trichomes, trichome number had a negative effect on walking speed. It was highest on plastic, followed by rose. No differences were found among chrysanthemum, gerbera and tomato. Walking speed was slightly higher on disks without prey. Proportion of time spent walking was the same on leaf disks of all plant species, with and without prey. No effect of glandular trichomes on tomato leaves was seen. Most thrips were killed and consumed on gerbera, and least on rose. Predation rates on chrysanthemum and plastic were intermediate. In contrast, no differences in oviposition rate were found among plant species. The results of this study indicate that trichome density can explain some of the variability in efficacy of A. swirskii on different crops. Release rates of A. swirskii may need to be adjusted depending on the crop in which it is used.     

中国特有属藤山柳属(猕猴桃科)植物的叶表皮形态及其分类学意义

由于形态特征变异和地理分布区域存在重叠,中国特有属藤山柳属(猕猴桃科)的物种划分问题长期以来存在争议,曾被分为20种或修订为含1种4个亚种的单型属。该研究选取了在形态和地理分布上有代表性的29个居群的184份标本,利用光学显微镜和扫描电镜观察了叶表皮形态和微形态特征,以探讨它们的分类学意义。结果表明:(1)藤山柳属植物叶表皮毛被的形态和微形态特征有3类,即光滑-短柱状毛、刚毛-长柱状毛/长刺毛、绒毛-单列多细胞毛,且这些特征在居群间差异明显,并各具明显的地理分布格局,支持把藤山柳属分为3类,即光滑类、刚毛类和绒毛类。(2)3类藤山柳植物在个别居群表现出部分同域分布现象,在峨眉山不同海拔高度的3个居群存在垂直地带性分布特点。(3)藤山柳属植物叶表皮的其他形态特征,如不规则型表皮细胞、6类气孔器、叶表皮初级蜡质纹饰以网状隆起为主,伴随着2～4类次级纹饰,在居群间变化多样等均没有明显的分类学意义。(4)由于具有相同的叶表皮形态特征和地理分布,建议把繁花藤山柳合并到绒毛藤山柳,故支持藤山柳属是1个正处于分化进程中的单型属,包括1个种3个亚种。     

The functions of plant feeding in the omnivorous predator Dicyphus hesperus: water places limits on predation

1. Dicyphus hesperus Knight (Heteroptera: Miridae) nymphs were fed from egg hatch to the adult stage on Ephestia kuehniella eggs provided either alone or in combination with tomato leaves or with a supplementary water source. 2. Only 6% of individuals completed nymphal development on a diet of eggs alone. In contrast, a high proportion of nymphs completed development on a diet of eggs when either tomato leaves (97%) or a supplementary water source (88%) were provided. 3. The development times of nymphs given access to leaves were significantly shorter than those of nymphs given access only to supplementary water. 4. Adult female D. hesperus that were given access to tomato leaves prior to feeding trials consumed significantly more eggs in a 4‐h period than females that were dehydrated before trials. Dehydrated females that were allowed access to water for 3 h before trials consumed an intermediate number of eggs. 5. Plant feeding or access to some other water source is required for prey feeding, growth, and development in D. hesperus, and acquisition of water is proposed as a primary function of plant feeding. In addition, D. hesperus derives nutrients from plant feeding that increase the rate of nymphal development, although nymphs cannot complete development when provided only with tomato leaves. 6. Three simple models are presented of feeding behaviour in predatory Heteroptera where the amount of plant feeding either decreases, increases, or is constant as a function of the amount of prey feeding. The models are discussed with reference to the results and the probable multifunctional nature of plant feeding in predatory Heteroptera.     

Plant species modifies the functional response of Phytoseiulus persimilis (Acari: Phytoseiidae) to Tetranychus urticae (Acari: Tetranychidae): implications for biological control

The functional response of the predatory mite Phytoseiulus persimilis Athias-Henriot to eggs of its prey, the spider mite Tetranychus urticae Koch was examined on three plant species. Experiments were done to determine whether differences in the functional response on the three plant species were due to the morphological features of the crop directly on the predator or through an effect of the plant species on the prey. The results show that crop morphology is the only factor influencing the predatory ability of P. persimilis on the three plant species. Fewer eggs were eaten on Ceanothus thyrsiflorus var. 'Autumnal Blue', the plant species with hairy leaves, and greater numbers of prey consumed on Choisya ternata, a species with smooth leaves. However, similarly few eggs were eaten on the smooth, but waxy leaved Euonymus japonicus as on Ceanothus thyrsiflorus, demonstrating that morphological characters of leaves other than the possession of hairs and trichomes may affect the rates of predation. The implications of these results for the tritrophic interactions between plant, predator and prey, and the development of suitable biological control strategies are discussed.     

PATTERNS OF VARIATION IN FOLIAR TRICHOMES OF EASTERN NORTH AMERICAN QUERCUS

Scanning electron microscopy of leaf trichomes of the forty two native species of oaks in eastern North America indicates five patterns of variability: 1) Eight trichome types are evident among the species and each species possesses a definite complement of trichome types. Certain trichomes are restricted to particular subgenera and series. 2) An obvious seasonal loss of trichomes occurs during leaf maturation. This loss may be both quantitative in terms of trichome density and qualitative in terms of trichome type. 3) There is an obvious difference between the adaxial and abaxial surfaces. The adaxial side of most oak leaves is dark green, lustrous, and glabrous or glabrate. The abaxial surface either remains pubescent, becomes glabrate or glabrous, or maintains trichomes along the midrib or in the axils of major secondary veins. There are also initial quantitative and qualitative trichome differences between the two sides. 4) Geographical and ecological variations are due in part to non-genetic ecophenic modifications, ecotypic differentiation, and random genetic differences not necessarily correlated with environmental conditions. Trichome types are considered to be less affected by environment than is trichome density. 5) Hybridization and introgression within a subgenus leads to localized variability. Trichomes of hybrids are usually a combination of the parental types. These five patterns of variation are predictable and appear to be held within rather narrow limits. The complement of foliar trichomes, therefore, is a reliable character in the taxonomy of the oaks.     

Threat of Infection and Threat-Avoidance Behavior in the Predator <Emphasis Type="Italic">Dicyphus hesperus</Emphasis> Feeding on Whitefly Nymphs Infected with an Entomopathogen

The nature and severity of intraguild interactions between predators and entomopathogens will be determined, in part, by a combination of threat of infection, and avoidance of that threat by the predator. We determined the threat of infection posed by the entomopathogen, Paecilomyces fumosoroseus (as PFR-97™) to the generalist predator, Dicyphus hesperus. We then asked if D. hesperus displays behavioral avoidance of infection while foraging for whitefly nymphs at different stages of infection by the pathogen. When exposed to leaf surfaces treated with the pathogen, 28% of adult female predators died due to infection. Consumption of Ephestia kuehniella eggs by surviving predators over 6 d was significantly reduced, suggesting effects of a sublethal infection. Whitefly nymphs that had been treated with P. fumosoroseus 3 d prior were acceptable as prey to D. hesperus but whitefly nymphs that had been treated with P. fumosoroseus 5 days prior were not. When foraging for whitefly nymphs, adult D. hesperus females rejected infected nymphs 96% of the time, compared to 39% of non-infected nymphs. Paecilomyces fumosoroseus therefore presents a measurable threat to D. hesperus through mortality and reduced prey consumption. Dicyphus hesperus does not avoid initial contact with infected prey but does not feed on such prey. The mechanism underlying these rejections could be due to either avoidance of infection or rejection of prey already consumed by the infectious agent. These results suggest that predation by D. hesperus foraging among infected whitefly nymphs under greenhouse or natural conditions could be reduced through a combination of pathogenicity and reduced efficiency of foraging.