Enemy-free space and habitat-specific host specialization in a butterfly

The majority of herbivorous insects have relatively specialized food habits. This suggests that specialization has some advantage(s) over generalization. Traditionally, feeding specialization has been thought to be linked to digestive or other food-related physiological advantages, but recent theory suggests that generalist natural enemies of herbivorous insects can also provide a major selective pressure for restricted host plant range. The European swallowtail butterfly Papilio machaon utilizes various plants in the Apiaceae family as hosts, but is an ecological specialist being monophagous on Angelica archangelica in southern Sweden. This perennial monocarp grows in three seaside habitat types: (1) on the barren rocky shore in the absence of any surrounding vegetation, (2) on the rocky shore with some surrounding vegetation, and (3) on species-rich meadows. The rocky shore habitat harbors few invertebrate generalist predators, whereas a number of invertebrate predators abound in the meadowland habitat. Here, we test the importance of enemy-free space for feeding specialization in Papilio machaon by assessing survival of larvae placed by hand on A. archangelica in each of the three habitat types, and by assessing the habitat-specificity of adult female egg-laying behavior by recording the distribution of eggs laid by free-flying adult females among the three habitat types. Larval survival was substantially higher in the rocky shore habitat than in the meadowland and significantly higher on host plants without surrounding vegetation on the rocky shore. Eggs laid by free-flying females were found in all three habitat types, but were significantly more frequent in the rocky shore habitat, suggesting that females prefer to lay eggs in the habitat type where offspring survival is highest. These results show that larval survivorship on the same host plant species can be strongly habitat-specific, and suggest that enemy-free space is an underlying factor that drives feeding specialization in Papilio machaon.     

Beyond preference and performance: host plant selection by monarch butterflies,Danaus plexippus

The connection between adult preferences and offspring performance is a long‐standing issue in understanding the evolutionary and ecological forces that dictate host associations and specialization in herbivorous insects. Indeed, decisions made by females about where to lay their eggs have direct consequences for fitness and are influenced by interacting factors including offspring performance, defence and competition. Nonetheless, in addition to these attributes of the offspring, a female's choices may be affected by her own prior experience. Here we examined oviposition preference, larval performance and the role of learning in the monarch butterfly, Danaus plexippus, which encounters diverse milkweed host species across its broad range and over the course of migration. Monarch females consistently preferred to oviposit on Asclepias incarnata subspecies pulchra. This plant, however, was associated with poor caterpillar growth, low sequestration of toxins and the highest plant defences (latex and trichomes). We examined flexibility in this apparently maladaptive preference by testing the impact of previous experience and competition on preference. Experience laying on an alternative plant species enhanced preference for that species in contrast to A. i. pulchra. In addition, presence of a (competing) conspecific caterpillar on A. i. pulchra had a strongly deterrent effect and reversed host plant preferences. Thus, monarch butterflies exhibit preferences contrary to what would be expected based on offspring development and sequestered defences, but their preferences are altered by learning and competition, which may allow butterflies to shift preferences as they encounter diverse milkweeds across the landscape. Learning and perception of threats (i.e. competition or predation) may be critical for most herbivorous insects, which universally experience heterogeneity among their potential host plants.     

Plant responses to butterfly oviposition partly explain preference–performance relationships on different brassicaceous species

Oecologia - The preference–performance hypothesis (PPH) states that herbivorous female insects prefer to oviposit on those host plants that are best for their offspring. Yet, past attempts to...     

The ‘mother knows best’ principle: should soil insects be included in the preference–performance debate?

Abstract.  1. Few entomological studies include soil-dwelling insects in mainstream ecological theory, for example the preference–performance debate. The preference–performance hypothesis predicts that when insect herbivores have offspring with limited capacity to relocate in relation to a host plant, there is a strong selection pressure for the adult to oviposit on plants that will maximise offspring performance.
2. This paper discusses the proposition that insect herbivores that live above ground, but have soil-dwelling offspring, should be included in the preference–performance debate. Twelve relevant studies were reviewed to assess the potential for including soil insects in this framework, before presenting a preliminary case study using the clover root weevil ( Sitona lepidus ) and its host plant, white clover ( Trifolium repens ).
3. Maternal S. lepidus preferentially oviposited on T. repens plants that had rhizobial root nodules (which enhance offspring performance) rather than T. repens plants without nodules, despite plants having similar foliar nutritional quality. This suggests that adult behaviour above ground was influenced by below-ground host-plant quality.
4. A conceptual model is presented to describe how information about the suitability for offspring below ground could underpin oviposition behaviour of parental insects living above ground, via plant- and soil-mediated semiochemicals. These interactions between genetically related, but spatially separated, insect herbivores raise important evolutionary questions such as how induced plant responses above ground affect offspring living below ground and vice versa.     

Oviposition site preference and developmental performance of a gall‐inducing leafhopper on galled and non‐galled host plants

Oviposition preferences of herbivorous insects affect offspring performance. Both positive and negative links between oviposition preference and offspring performance have been reported for many species. A gall‐inducing leafhopper, Cicadulina bipunctata Melichar (Hemiptera: Cicadellidae), feeds on various Poaceae plants and induces galls of enhanced nutritional value for their offspring. Although gall induction by C. bipunctata improves nymphal performance, the oviposition preference of females between galled and non‐galled host plants is still unclear. In this paper, the nymphal performance and oviposition and feeding‐site preference of C. bipunctata were investigated using galled wheat, Triticum aestivum L., and non‐galled barley, Hordeum vulgare L., as host plants. The survival rate of C. bipunctata on wheat was significantly higher than on barley. In the choice test, significantly more eggs were laid into barley, whereas the number of eggs deposited on both hosts was not significantly different in the no‐choice test. The number of settling individuals per leaf area was not significantly different between wheat and barley, suggesting no clear preference for oviposition between these plants. Experience as a nymph with a growing host did not affect oviposition preference as adult female. The inconsistent correspondence between offspring performance and oviposition preference of C. bipunctata may reflect the high mobility of nymphs and/or differences in leaf area between host plants. The results indicate that the previous finding that oviposition preference and offspring performance are not always positively correlated in herbivorous insects is applicable to gall‐inducing insects.     

Can mechanism help explain insect host choice?

Evolutionary theory predicts that herbivorous insects should lay eggs on plants in a way that reflects the suitability of each plant species for larval development. Empirical studies, however, often fail to find any relationship between an adult insect's choice of host-plant and offspring fitness, and in such cases, it is generally assumed that other 'missing' factors (e.g. predation, host-plant abundance, learning and adult feeding sites) must be contributing to overall host suitability. Here, I consider an alternative theory - that a fitness cost inherent in the olfactory mechanism could constrain the evolution of insect host selection. I begin by reviewing current knowledge of odour processing in the insect antennal lobe with the aid of a simple schematic: the aim being to explain the workings of this mechanism to scientists who do not have prior knowledge in this field. I then use the schematic to explore how an insect's perception of host and non-host odours is governed by a set of processing rules, or algorithm. Under the assumptions of this mechanistic view, the perception of every plant odour is interrelated, and seemingly bad host choices can still arise as part of an overall adaptive behavioural strategy. I discuss how an understanding of mechanism can improve the interpretation of theoretical and empirical studies in insect behaviour and evolution.     

Plant secondary substances and insects behaviour

Allelochemicals are storing in different location in plant tissues as inactive form. Number of identified compounds may now exceed 100,000. Environmental factors have an effect on allelochemicals concentration in plants. Many allelochemicals classified as toxins or deterrents for herbivorous insects. Allelochemicals play a major role in feeding or ovipositing stimulants for some specialist insects. Consumption and assimilation of herbivorous insects had affected by the type of allelochemicals in host plants. Allelochemicals have an acute or chronic toxicity on herbivorous insects. Most specialist herbivorous insects rely heavily of ingested plant allelochemicals. Plant allelochemicals may influence an insect's susceptibility to pathogens such as bacteria, fungi and nematode. Specialists herbivorous insect can be using the allelochemicals in their host plants as protection against natural enemies. Some herbivorous insects are synthesising the aggregation, attracting, alarm or mating pheromone from the allelochemicals in their host plants.     

Maternal effects in the soft scale insect Saissetia coffeae (Hemiptera: Coccidae)

Effects of maternal environment on offspring performance have been documented frequently in herbivorous insects. Despite this, very few cases exist in which exposure of parent insects to a resource causes the phenotype of their offspring to be adjusted in a manner that is adaptive for that resource, a phenomenon called adaptive transgenerational phenotypic plasticity. I performed a two-generation reciprocal cross-transplant experiment in the field with the soft scale insect Saissetia coffeae (Hemiptera: Coccidae) on two disparate host plant species in order to separate genetic effects from possible transgenerational plasticity. Despite striking differences in quality between host species, maternal host had no effect on overall offspring performance, and I detected no "acclimatization" to the maternal host species. However, there was a significant negative association between maternal and offspring development times, with potentially adaptive implications. Furthermore, offspring of mothers reared in an environment where scale densities were higher and scales were more frequently killed by fungi were significantly less likely to suffer from fungal attack than were offspring of mothers reared in an environment where densities were low and fungal attack was rare. Although S. coffeae does not appear to alter offspring phenotype to increase offspring fitness on these two distinct plant species, it does appear that offspring phenotype may be responding to some subtler aspects of maternal environment. In particular, the possibility of induced transgenerational prophylaxis in S. coffeae deserves further investigation.     

植食性昆虫寄主植物嗅觉搜寻述介

植食性昆虫与寄主植物关系的本质是化学。植食性昆虫搜寻寄主的嗅觉媒介是植物气味即化学信息物质。在介绍植物气味构成及其扩散模型基础上,阐述了植物气味在地上植食性昆虫成虫、幼虫和地下植食性昆虫搜寻寄主过程中的嗅觉导向作用,并指出了今后相关研究需要注意的问题。从植物与环境因子的关系来看,植物气味包括构成性气味和诱发性气味两类,这两类气味的概念既相联系而又不同。构成性气味组分及构成因植物分类地位等而不同。诱发性气味组分因植食性昆虫取食、植物病原微生物、机械致伤等因子的胁迫而变化,这种变化性状随植物属和/或种、植株生长发育阶段、胁迫因子性质及其作用方式而不同。无论是哪种植物气味,其释放均具有节律性。气味扩散过程比较复杂,扩散状态可用数学模型表征。对于地上植食性昆虫成虫,植物气味对其寄主搜寻行为具有导向特异性,重点分析了这种特异性形成的两个假说;鳞翅目昆虫幼虫,能够利用植物化学信息物质趋向寄主植物或回避非寄主植物;地下植食性昆虫搜寻寄主,既与寄主植物地下组织释放或分泌的次级代谢物有关,又与一些初级代谢物有关。初级代谢物中的CO₂,起着“搜寻触发器”作用。有助于增强人们对昆虫与植...     

Similarity and specialization of the larval versus adult diet of European butterflies and moths

Many herbivorous insects feed on plant tissues as larvae but use other resources as adults. Adult nectar feeding is an important component of the diet of many adult herbivores, but few studies have compared adult and larval feeding for broad groups of insects. We compiled a data set of larval host use and adult nectar sources for 995 butterfly and moth species (Lepidoptera) in central Europe. Using a phylogenetic generalized least squares approach, we found that those Lepidoptera that fed on a wide range of plant species as larvae were also nectar feeding on a wide range of plant species as adults. Lepidoptera that lack functional mouthparts as adults used more plant species as larval hosts, on average, than did Lepidoptera with adult mouthparts. We found that 54% of Lepidoptera include their larval host as a nectar source. By creating null models that described the similarity between larval and adult nectar sources, we furthermore showed that Lepidoptera nectar feed on their larval host more than would be expected if they fed at random on available nectar sources. Despite nutritional differences between plant tissue and nectar, we show that there are similarities between adult and larval feeding in Lepidoptera. This suggests that either behavioral or digestive constraints are retained throughout the life cycle of holometabolous herbivores, which affects host breadth and identity.     

Nonadaptive radiation: Pervasive diet specialization by drift in scale insects?

At least half of metazoan species are herbivorous insects. Why are they so diverse? Most herbivorous insects feed on few plant species, and adaptive host specialization is often invoked to explain their diversification. Nevertheless, it is possible that the narrow host ranges of many herbivorous insects are nonadaptive. Here, we test predictions of this hypothesis with comparative phylogenetic analyses of scale insects, a group for which there appear to be few host‐use trade‐offs that would select against polyphagy, and for which passive wind‐dispersal should make host specificity costly. We infer a strong positive relationship between host range and diversification rate, and a marked asymmetry in cladogenetic changes in diet breadth. These results are consonant with a system of pervasive nonadaptive host specialization in which small, drift‐ and extinction‐prone populations are frequently isolated from persistent and polyphagous source populations. They also contrast with the negative relationship between diet breadth and taxonomic diversification that has been estimated in butterflies, a disparity that likely stems from differences in the average costs and benefits of host specificity and generalism in scale insects versus butterflies. Our results indicate the potential for nonadaptive processes to be important to diet‐breadth evolution and taxonomic diversification across herbivorous insects.     

The role of nectar sources for oviposition decisions of the common blue butterfly Polyommatus icarus

Neural limitations on information processing have been shown to play an important role for host plant specialization in herbivorous insects. The necessity of fast and accurate decisions favors the adoption of a few high-contrast signals, which selects against the use of multiple resources. Many species face a similar problem when searching for adult food sources and the simultaneous need to fulfill both search tasks can lead to a potential conflict. Some insects use the same host plant species for both adult and larval nutrition, which makes it possible to decrease the number of search images and thus potentially increase efficiency of the choices. The aim of this study was to investigate if there is a connection between choice of nectar sources and choice of oviposition host plant. In a laboratory experiment, females of Polyommatus icarus preferred to oviposit on Lotus corniculatus plants with flowers over those without flowers. Observations of behavioral sequences also revealed that oviposition often followed immediately after nectaring. The results suggest that nectar availability could play an important role in oviposition decisions of P. icarus and can provide one explanation to why some phytophagous insects not always choose the host plant that gives the best offspring performance.     

Have Genetic Trade-Offs in Host Use been Overlooked in Arthropods?

A popular hypothesis to explain the high degree of host specialisation observed among mites and insects is the existence of host-associated fitness trade-offs. According to this theory, adaptation to a host results in a relatively poorer performance on alternative hosts due to the antagonistic pleiotropic action of one or more genes. Evidence in favour of the genetic trade-off hypothesis is however scarce. Recent ecological work has shown that the optimisation of adult performance drives the evolution of host choice in at least some phytophagous insects. Yet, most ecological and evolutionary studies on host choice assume that females maximise their fitness by optimising offspring performance. In this paper, we investigate whether a general lack of attention for the role of adult performance in host choice may have diminished the chance of detecting genetic trade-offs. We reviewed the literature on genetic trade-offs and showed that most studies neglected host specific variation in adult performance. Moreover, studies that considered both adult and offspring performance had a higher chance of detecting genetic fitness trade-offs. Our results also suggested that studies on asexual reproducing species tend to detect trade-offs more often than studies on obligate sexual reproducing species. We argue that future studies on genetic trade-offs should take all fitness parameters into account in order to be conclusive. This approach may reveal (i) that genetic trade-offs are more common than hitherto reported and/or (ii) that genetic trade-offs are more common, or more easily detected among asexual reproducing species like mites and aphids. Co-ordinating editor: Dr. F. Stuefer     

Phytophagous insect fauna tracks host plant responses to exotic grass invasion

The high dependence of herbivorous insects on their host plants implies that plant invaders can affect these insects directly, by not providing a suitable habitat, or indirectly, by altering host plant availability. In this study, we sampled Asteraceae flower heads in cerrado remnants with varying levels of exotic grass invasion to evaluate whether invasive grasses have a direct effect on herbivore richness independent of the current disturbance level and host plant richness. By classifying herbivores according to the degree of host plant specialization, we also investigated whether invasive grasses reduce the uniqueness of the herbivorous assemblages. Herbivorous insect richness showed a unimodal relationship with invasive grass cover that was significantly explained only by way of the variation in host plant richness. The same result was found for polyphagous and oligophagous insects, but monophages showed a significant negative response to the intensity of the grass invasion that was independent of host plant richness. Our findings lend support to the hypothesis that the aggregate effect of invasive plants on herbivores tends to mirror the effects of invasive plants on host plants. In addition, exotic plants affect specialist insects differently from generalist insects; thus exotic plants affect not only the size but also the structural profile of herbivorous insect assemblages.     

The pollen beetle, Meligethes aeneus, changes egg production rate to match host quality

Motivation-based models dominate current theory concerning host plant selection for oviposition by herbivorous insects. A female searching for a host plant will be more likely to accept a host which is of inferior quality for her offspring if motivation is high, e.g. a large eggload or long time since last oviposition. This implies that insects will accumulate eggs if exposed to hosts of low acceptability and after a time lay eggs on such hosts. An alternative strategy for insects when exposed to less acceptable hosts is to stop producing, instead of accumulating, eggs. Thus, resources would be saved until a more acceptable host is found. If this hypothesis is true, a herbivorous insect would cease egg production when exposed to hosts of low acceptability and resume egg production when exposed to hosts of high acceptability. Previous exposure should not affect oviposition rate when an insect encounters a new host of a different quality. In an earlier study pollen beetles, Meligethes aeneus (F.) (Coleoptera: Nitidulidae), did not accumulate eggs in the absence of high quality hosts. In this study we monitored the daily oviposition rate of female pollen beetles on hosts plants of low (Sinapis alba L.), intermediate (Brassica nigra Koch) or high (B. napus L.) acceptability over a 5-day period. Individuals were then switched to an oviposition resource of a different acceptability. Beetles moved from high- to low-acceptability plants reduced their oviposition rate considerably. In the opposite case, low to high acceptability, the rate of oviposition increased markedly after the switch. When M. aeneus were moved from the high-acceptability host to that of intermediate acceptability oviposition rate was modified accordingly. However, when moved to the intermediate host from a host of low acceptability oviposition on B. nigra was much less than would normally be expected. A possible mechanism for this finding is discussed. M. aeneus, by adjusting oviposition rate to host acceptability, maximizes the average host quality for offspring, even at the cost of a lower egg-laying rate. Received: 5 October 1998 / Accepted: 20 April 1999     

Host plant specialization driven by sexual selection

We propose a new mechanism based on sexual selection to explain the evolution of diet breadth in insects. More specifically, we show that mate choice in females for certain diet-derived male pheromones can be exploited by maternal effect genes that preferentially place offspring on a specific host plant, resulting in specialization. Our analytical model also suggests that the process is more likely to occur with species that show male-congregating mating strategies, such as lekking and hilltopping. The model offers a new explanation for the similarity between the composition of male lepidopteran pheromones and the chemistry of their host plants and also suggests a novel mechanism of host plant shift. This is the first time that sexual selection has been proposed to drive host plant specialization and the first time that a mechanism with selection acting solely on the adult stage has been shown to be capable of determining larval feeding habits.     

Resource concentration hypothesis: effect of host plant patch size on density of herbivorous insects

The resource concentration hypothesis (Root 1973) predicts that specialist herbivorous insects should be more abundant in large patches of host plants, because the insects are more likely to find and stay longer in those patches. Between August 1989 and January 1990 we experimentally tested Root's hypothesis by analyzing the numerical response of four species of herbivorous insects associated with patches of 4, 16, 64 and 225 cabbage plants, Brassica oleracea var. capitata. In addition, we studied the colonization of patches by adults of Plutella xylostella (L.) (Lepidoptera: Plutellidae), and the migration of their larvae in patches of different sizes. No herbivorous insect densities differed significantly with patch size. Adults of P. xylostella colonized all kind of patches equally. Larvae did not migrate between patches, and their disappearance rate did not differ between patches. The resource concentration hypothesis is organism-dependent, being a function of the adult and juvenile herbivore dispersal behavior in relation to the spatial scale of patchiness.     

Plant and insect cuticular lipids serve as behavioral cues for insects

The roles of plant and insect cuticular lipids in insect and plant interactions are reviewed. Emphasis is given to the influence that the host plant and the surface lipids of the host plant have upon insect herbivores and the predators and parasitoids of these herbivores. Variations in cuticular lipids of herbivorous insects are dependent upon the host plant, and these variations may affect the behavior of predators and parasitoids. The cuticular lipids of species which interact on multiple trophic levels are compared. Similarities were found between the hydrocarbons of herbivorous insects, their host plants, and their predators or parasitoids.     

大气CO_2浓度升高对植物-植食性昆虫的作用机制

大气二氧化碳浓度升高及其伴随的全球变暖引起国内外科学家的极大关注。CO2浓度升高主要通过改变植物的初级和次级代谢产物,影响以之为食的昆虫。本文结合作者近年来的研究成果,着重于以CO2浓度升高为作用因子,以植物和植食性昆虫的相互关系为对象,比较了咀嚼式口器昆虫与刺吸式口器昆虫对大气CO2浓度升高的响应特征,分析了不同取食类型昆虫-植物对大气CO2浓度升高的响应机制。     

Effects of host plant and maternal feeding experience on population vital rates of a specialized leaf beetle

In herbivorous insects, the interaction between adult preference and progeny performance on specific host plants is modified by maternal feeding experience and host plant quality. Ultimately, changes in the strength of this interaction can affect insect population dynamics. In this study, we hypothesized that adult host plant preference influences progeny performance through a maternal feeding experience × host plant interaction, that is, the effect of adult feeding experience on progeny performance will depend on the host plant. Second, that decoupling of the preference–performance relationship due to host switching results in different population vital rates changing population dynamics. An increase in development time and a decrease in body size of individuals in the alternate host should decrease population growth. We tested these hypotheses using two lines of the tortoise beetle Chelymorpha varians Blanchard fed with two hosts (Convolvulus arvensis and Calystegia sepium). Maternal feeding experience treatments were crossed with host plant species, and the offspring’s developing time and adult size were measured. The host plant influence on the beetle’s population vital rates was tested using stage-structured matrix population models and life table response experiments. Host plant preference affected offspring body size through a host plant effect that contributed to adaptive life history responses only in the better quality host. C. varians’ population growth was positive when fed with either host; comparatively, however, C. sepium had a negative effect on growth by reducing all transition probabilities of the life cycle stages of the beetle. Here, we show that individuals of C. varians prefer and perform differently on distinct hosts and that these patterns influence population vital rates in different ways. When beetles prefer the host plant where their progeny performs best, life history responses and life stage transitions lead to higher population growth; otherwise, growth rate decreases.