A tritrophic analysis of host preference and performance in a polyphagous leafminer

The optimal oviposition theory predicts that oviposition preferences of phytophagous insects should correlate with host suitability for their offspring. As plant host suitability depends not only on its quality as food, but also on its provision of enemy‐free space, we examined the relationship between adult host preference and offspring performance for the leafminer Liriomyza huidobrensis (Blanchard) (Diptera: Agromyzidae) on various host plants, considering also the interaction with natural enemies. Preference and offspring performance were assessed through observational field data and laboratory experiments in central Argentina. Field data suggested a positive host preference – performance linkage, as the leafminer attained larger body size on the crops where it was more abundant. Laboratory trials supported these results: Vicia faba L. (Fabaceae) was the preferred host in the laboratory as well as in the field, performance of L. huidobrensis being also best on this host, with highest survival rates and shortest development time. The actively feeding larval stage showed the largest plant‐related effects. Higher overall parasitism rates were found on plants from which smaller leafminers were reared, reinforcing the preference–performance linkage. On the other hand, the main parasitoid Phaedrotoma scabriventris Nixon (Hymenoptera: Braconidae) reached larger body size, and caused higher mortality rates on crops where the leafminer was larger. Changes in abundance of particular parasitoid species could thus modify overall parasitism trends.     

Host feeding experience affects host plant odour preference of the polyphagous leafminer Liriomyza bryoniae

Herbivorous insects use highly specific volatiles or blends of volatiles characteristic to particular plant species to locate their host plants. Thus, data on olfactory preferences can be valuable in developing integrated pest management tools that deal with manipulation of pest insect behaviour. We examined host plant odour preferences of the tomato leafminer, Liriomyza bryoniae (Kaltenbach) (Diptera: Agromyzidae), which is an economically important agricultural pest widespread throughout Europe. The odour preferences of leafminers were tested in dependence of feeding experiences. We ranked host plant odours by their appeal to L. bryoniae based on two‐choice tests using a Y‐tube olfactometer with five host plants: tomato, Solanum lycopersicum Mill.; bittersweet, Solanum dulcamara L.; downy ground‐cherry, Physalis pubescens L. (all Solanaceae); white goosefoot, Chenopodium album L. (Chenopodiaceae); and dead nettle, Lamium album L. (Lamiaceae). The results imply that ranking of host plant odours by their attractiveness to L. bryoniae is complicated due to the influence of larval and adult feeding experiences. Without any feeding experience as an adult, L. bryoniae males showed a preference for the airflow with host plant odour vs. pure air, whereas females did not display a preference. Further tests revealed that adult feeding experience can alter the odour choice of L. bryoniae females. After feeding experience, females showed a preference for host plant odour vs. pure air. Feeding experience in the larval stage influenced the choice by adults of both sexes: for males as well as females reared on bittersweet the odour of that plant was the most attractive. Thus, host feeding experience both in larval and/or adult stage of polyphagous tomato leafminer L. bryoniae influences host plant odour preference by adults.     

多食性斑潜蝇对寄主植物的选择

综述了有关斑潜蝇对寄主植物选择的研究进展。斑潜蝇对寄主植物的选择表现为对植物组织的选择、对植物不同部位叶片的选择、对同种植物不同品种的选择、对不同寄主植物的选择四个方面。斑潜蝇对寄主植物的选择主要受叶表面的物理性质、植物叶的化学物质成分、成长经历等因素的影响。阐述了多食性斑潜蝇寄主植物范围扩大的原因。     

Evolutionary conservatism of geographic variation in host preference in Callosobruchus maculatus

Abstract.  1. A study of host preference of four pairs of populations of the cowpea weevil Callosobruchus maculates was carried out. The pairs had different geographical origins.
2. One population of each pair had been maintained for about 110 generations on cowpea Vigna unguiculata , the other population had been maintained on mung bean V. radiata . Half of the tested females from each population were raised on cowpea and exposed to this host prior to the assay; the other half was raised on mung bean. This design permitted assessment of the relative contributions of geographical origin, recent host use in the laboratory, and individual experience, to variation in host preference.
3. Host preference was assayed by letting the females oviposit on an equal-weight mixture of cowpea and mung seeds; two experiments were performed six generations apart.
4. Both experiments revealed a strong effect of geographical origin: populations originating from Nigeria laid a much greater proportion (68–86%) of their eggs on cowpea than those originating from Uganda and Yemen (30–42%); those from Cameroon were intermediate (56–60%). These preferences were not affected consistently by about 110 generations of laboratory evolution on one or the other host, or by experience of individual females.
5. These results indicate considerable geographical variation in host preference, and suggest that host preference is behaviourally inflexible and evolutionarily conserved.     

Relationship between oviposition preference and offspring performance in Australian Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae)

Abstract We investigated the oviposition preference and larval performance of Helicoverpa armigera under laboratory conditions to determine if the oviposition preference of individual females on maize, cowpea and cotton correlates with offspring performance on the leaves of the same host plants. The host-plant preference hierarchy of females did not correlate with their offspring performance. Female moths chose host plants that contributed less to their offspring fitness. Plant effects accounted for the largest amount of variation in offspring performance, while the effects of female (family) was low. The offspring of most females (80%, n  = 10) were broadly similar, but 20% (two out of 10), showed marked difference in their offspring performance across the host-plant species. Similarly, there was no relation between larval feeding preference and performance. However, like most laboratory experiments, our experi-mental design does not allow the evaluation of ecological factors (for example, natural enemies, host abundance, etc.) that can play an important role in larval performance in the field. Overall, the results highlight the importance of carrying out preference performance analysis on the individual or family level, rather than pooling individuals to obtain average population data.     

南美斑潜蝇寄主选择性与植物次生化合物及叶毛的关系

采用非自由选择法,以刺伤孔（包括取食孔和产卵孔）数为指标,在室内测定了南美斑潜蝇对6科、16种、22个蔬菜品种的寄主选择性。结果表明,南美斑潜蝇对供试蔬菜品种的选择性存在显著差异,选择性大小依次为豆科>菊科>葫芦科>茄科>伞形科>十字花科; 菜豆和茼蒿是南美斑潜蝇最喜欢产卵和取食的寄主,而甘蓝和番茄是其选择性最差的寄主。化学分析和解剖镜观察表明,叶片单宁酸、黄酮含量及叶毛密度在供试品种间存在显著差异： 苦瓜的单宁酸含量最高,其次为豌豆、番茄和青椒,含量最低的为金丝搅瓜、茼蒿、菜豆和美国西芹; 黄酮含量最高的为甘蓝和番茄,其次为苦瓜,最低为茼蒿、菜豆和美国西芹; 南瓜和小白菜的叶毛密度最高,其次为西葫芦、金丝搅瓜和甘蓝,最低为青椒、莴苣和苦瓜。相关分析表明,南美斑潜蝇寄主选择性与叶片单宁酸和黄酮含量呈显著的负相关关系,相关系数分别为-0.4425（ P=0.0392）和 -0.5728（ P=0.0053）,而与叶毛密度相关关系不显著（ R＝-0.1807,P=0.4211）。说明黄酮和单宁酸对南美斑潜蝇产卵和取食具有明显的抑制作用。     

The influence of host plant variation and intraspecific competition on oviposition preference and offspring performance in the host races of Eurosta solidaginis

1. A series of experiments was conducted to measure the impact of plant genotype, plant growth rate, and intraspecific competition on the oviposition preference and offspring performance of the host races of Eurosta solidaginis (Diptera: Tephritidae), a fly that forms galls on Solidago altissima and Solidago gigantea (Asteraceae). Previous research has shown that both host races prefer to oviposit on their own host plant where survival is much higher than on the alternate host plant. In this study, neither host race showed any relationship between oviposition preference and offspring performance in choosing among plants of their natal host species. 2. The larval survival of both host races differed among plant genotypes when each host race oviposited on its natal host species. In one experiment, altissima host race females showed a preference among plant genotypes that was not correlated with offspring performance on those genotypes. In all other experiments, neither the altissima nor gigantea host race demonstrated a preference for specific host plant genotypes. 3. Eurosta solidaginis had a preference for ovipositing on rapidly growing ramets in all experiments, however larval survival was not correlated with ramet growth rate at the time of oviposition. 4. Eurosta solidaginis suffered high mortality from intraspecific competition in the early larval stage. There was little evidence, however, that females avoided ovipositing on ramets that had been attacked previously. This led to an aggregated distribution of eggs among ramets and strong intraspecific competition. 5. There was no interaction among plant genotype, plant growth rate, or intraspecific competition in determining oviposition preference or offspring performance.     

南美斑潜蝇寄主选择性与植物营养物质及叶绿素含量的关系

采用非自由选择法,在室内研究了南美斑潜蝇寄主选择性及其与植物叶片营养物质和叶绿素含量的关系.结果表明,南美斑潜蝇成虫对不同供试植物的选择性存在显著差异,选择性最强的为菜豆、茼蒿和西葫芦,最差的为番茄和甘蓝.相关分析表明,南美斑潜蝇成虫寄主选择性与叶片可溶性糖含量存在显著的正相关关系(R＝0.5014,P=0.0174,F＝6.717),而与叶片可溶性蛋白和叶绿素含量相关关系不显著.     

Preference ofPholetesor ornigis [Hym.: Braconidae] for the sap-feeding larvae of its host,the spotted tentiform leafminer,Phyllonorycter blacerdella [Lep.: Gracillaridae]

Significantly more eggs were deposited by the braconid,Pholetesor ornigis (Weed), in the sap-feeding larvae (i.e. instars 1–3), than in the tissue-feeding larvae (i.e. instars 4–5), of its host, the spotted tentiform leafminer,Phyllonorycter blancardella (Fabr.).     

番石榴实蝇寄主选择性试验

番石榴实蝇Bactrocera correcta(Bezzi)是一种为害多种水果的害虫,近年来在云南的元江暴发成灾。本试验选择番石榴实蝇的常见寄主:梨、芒果、苹果、桃、橙及芒果的不同品种进行室内试验,比较番石榴实蝇雌虫产卵对寄主的选择性及不同寄主对番石榴实蝇生长发育的影响。结果显示,番石榴实蝇雌虫对5种供试水果的寄生选择性大小依次为桃>梨>芒果>橙>苹果;番石榴实蝇对芒果不同品种的寄生选择性为三年芒>台农行1号>虎豹牙>胭脂芒>鹰嘴芒;番石榴实蝇在不同寄主上的蛀果期(卵及幼虫的历期)具有极显著的差异,以寄生桃和芒果的最短;从梨和苹果上饲养出的蛹最轻,且极显著低于其它寄主上饲养出来的蛹重;苹果上寄生的番石榴实蝇蛹重与蛀果期呈显著的正相关,而寄生梨的番石榴实蝇蛹重与蛀果期呈极显著的负相关。结果表明,番石榴实蝇雌虫产卵对不同寄主或同一寄主不同品种有明显的选择性,且该选择行为与其后代的生长发育有关。     

Effects of leaf nitrogen content on oviposition site selection, offspring performance, and intraspecific interactions in an omnivorous bug

Abstract.  1. Oviposition site preference and its relation to offspring performance have received much attention in studies of herbivorous insects. Although this relationship is of great ecological significance, its presence in predacious and omnivorous insects has hardly been explored. When selecting an oviposition site, omnivores are expected to respond to both prey availability and, even more strongly, to plant traits that affect both females and their offspring.
2. In this study, females of Orius albidipennis (Heteroptera: Anthocoridae) showed a strong preference for oviposition at the vein origin site of cotton leaves. It appears that this site is a limited resource for ovipositing females, because they defend it against conspecific intruders. This defensive behaviour was significantly more pronounced on nitrogen-rich than nitrogen-poor plants.
3. The females' defensive behaviour on nitrogen-rich leaves corresponds with offspring performance; the nymphs developed faster and enjoyed higher survival on nitrogen-rich than on nitrogen-poor leaves. At the between-plant level, oviposition preference was not significantly correlated with offspring performance, and egg hatching rate did not differ significantly between nitrogen treatments.
4. Oviposition site selection by this omnivorous bug appears, however, to be correlated positively with offspring performance at the within-leaf level. This is supported by the significantly higher hatching success of eggs deposited at the preferred vein origin site as compared with those deposited on other parts of the leaf.     

Adaptive learning of host preference in a herbivorous arthropod

Although many publications deal with the effects of experience on behaviour, adaptive learning (i.e. behavioural change with experience resulting in improved reproductive success) is poorly documented. We present direct evidence that learning of host preference improves fitness in the herbivorous mite, Tetranychus urticae . Individual mites from two strains were repeatedly given a choice between two host plants, tomato and cucumber, and then subjected to a performance test on each. For both strains, food experience affected the subsequent choice: individual mites learned to prefer cucumber over tomato. The performance test showed this effect to be adaptive, as the food plant the mites learned to prefer (cucumber) allowed for increased oviposition, survival and development. These findings have important implications for the interpretation of the preference–performance relationship among herbivorous arthropods. The frequently reported absence of such a relationship may be due to experience-dependent preference and/or performance.     

Divergent host plant preference causes assortative mating between sympatric host races of the ladybird beetle,Henosepilachna diekei

Divergent host preference (i.e. host fidelity) plays a significant role in the speciation process in phytophagous insects. However, how and to what extent this divergence reduces gene flow between populations has rarely been measured. Here, we estimated the intensity of assortative mating caused solely by host fidelity in two host races of the phytophagous ladybird beetle Henosepilachna diekei, specialized on Mikania micrantha (Asteraceae) and Leucas lavandulifolia (Lamiaceae) in West Java, Indonesia. These host races mated randomly in the absence of host plants under laboratory conditions, but demonstrated nearly complete assortative mating in field cages with the two host plants, by spending almost all of their time on their respective host plants. The frequency of assortative mating in the field cages was not affected drastically by host plant patch structure. These results suggest that fidelity to the different host plants yields directly almost complete reproductive isolation between the host races by limiting the habitat on the respective host plant. In addition, the high host fidelity also ensures female oviposition on the original host plant. As larvae cannot survive on non‐host plants, a positive association between female oviposition preference and larval performance on the host plant on which the beetles are specialized will further facilitate the evolution of host fidelity. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 606–614.     

Seasonal changes in leaf nutritional quality influence grass miner performance

1. The plant phenological age hypothesis predicts that phytophagous insects should prefer and perform better on phenologically young plants than on old plants because plant nutritional quality decreases with plant phenological age. This hypothesis was tested under field and laboratory conditions with the grass miner Chromatomyia milii on the free‐growth species Holcus lanatus. 2. The field experiment was conducted at four sample sites in Belgium where nutritional quality of H. lanatus leaves and performance of C. milii were monitored throughout the growth season. Foliar nutritional quality was highest early in the season due to high levels of proteins and soluble carbohydrates and low levels of lignin. Offspring performance (pupal size) decreased with plant phenological age, due at least partially to the decreasing foliar nutritional quality. 3. In the laboratory experiment, preference and performance of C. milii were determined on three different age classes of H. lanatus. Multiple‐choice experiments demonstrated that oviposition preference did not differ among age classes. Offspring survival decreased with plant phenological age, while pupal size did not differ among age classes. The relationship between foliar nutritional quality and plant phenological age was equivocal and did not correspond to the predictions of the plant phenological age hypothesis. 4. The results of the field experiment were consistent with the idea that the phenological age hypothesis holds in free‐growth species. The laboratory experiment gave only little support to the plant phenological age hypothesis. Possible causes for the differences between field and laboratory experiments are discussed.     

Intra-plant host selection, oviposition preference and larval survival of Chrysophtharta agricola (Chapuis) (Coleoptera: Chrysomelidae: Paropsini) between foliage types of a heterophyllous host

Abstract 1 Paropsine chrysomelid beetles defoliate commercial eucalypt plantations in Australia. Adults and larvae feed on the same host, with the larval food source determined by the oviposition choice of females. Most eucalypt species are heterophyllous, with their foliage undergoing distinct morphological and chemical changes between adult and juvenile growth. 2 The intra‐plant foliage feeding and oviposition preference adults and the larval development of Chrysophtharta agricola were examined using adult and juvenile foliage of a heterophyllous plantation species, Eucalyptus nitens. The foliage types differ in chemistry, toughness, waxiness and timing of production. 3 In the field, feeding damage caused by adult beetles was 15% more frequent on adult foliage than on juvenile foliage; however, egg batches were three times more common on juvenile than on adult foliage. 4 Oviposition preference for juvenile foliage over adult foliage was confirmed in choice trials in the laboratory, with adult fecundity and longevity not significantly different between foliage types. 5 Larval survival, development time and subsequent pupal weight were also unaffected by foliage type, suggesting that neither foliage type is nutritionally superior for adults or for larvae. However, adult foliage was significantly thicker than juvenile foliage and this may prove a physical constraint to larval establishment. Biotic and abiotic factors (including interactions with natural enemies, competition, microclimate and mate location) that may affect patterns of host plant utilization are discussed.     

Asymmetry of larval diet breadth and oviposition preference in Leptinotarsa undecimlineata

The relationship between preference and performance was studied under laboratory conditions for larvae and adult females of Leptinotarsa undecimlineata (Stål) (Coleoptera: Chrysomelidae). Leptinotarsa undecimlineata feeds exclusively on Solanum lanceolatum L. (Solanaceae) in the centre of Veracruz, Mexico. A common sympatric species, Solanum myriacanthum Dunal, is not used as host. We conducted a series of laboratory experiments to detect reasons for the exclusive use of one host plant. Leptinotarsa undecimlineata larvae were reared under laboratory conditions on both plant species, Solanum lanceolatum and S. myriacanthum . Length of larval stages, pupal weight, and percent adult eclosion were compared, and larval preference for both Solanum species was tested. Upon eclosion, adult females were exposed to each of the plants, and the effect of larval diet on adult preference was determined. Oviposition on both Solanum species was observed in no-choice tests. Larvae showed no significant preferences for either plant species. Pupal weight, however, was significantly higher on S. lanceolatum . Female beetles significantly preferred S. lanceolatum leaves, and no effect of preimaginal conditioning was found. Females reproduced in lower numbers after feeding on S. myriacanthum , oviposition initiated later, and females laid fewer eggs than females fed with S. lanceolatum . No female accepted S. myriacanthum for oviposition. Results obtained suggest that the potential host breadth of L. undecimlineata is broader in larvae than in female beetles.     

Discrimination within and between host species by a butterfly: implications for design of preference experiments

Experiments designed to reveal variation among individual parasites in preference for different host species may generate misleading results. Apparent variation in the order of preference among host species can be generated solely from variation in the strength of discriminations made within host species. We illustrate this with a study of oviposition preference in the butterfly Melitaea cinxia. All butterflies were tested on the same six individual plants, three Plantago lanceolata (P) and three Veronica spicata (V). Some insects repeatedly preferred all individual P over all individual V or vice versa. We designated these as "pure" species ranks. Other insects repeatedly produced "mixed" ranks, preferring some individual V over some P, and some individual P over some V. We show how a "mixed" rank butterfly could differ from a "pure" rank insect by discriminating either more within plant species and/or less between them. Therefore, discrimination within host species can mask or confound discrimination among species. We discuss implications for the design of preference experiments.     

Oviposition preference of Lycaena salustiusfor,and larval performance on,a novel host plant: an example of ecological fitting

1. ‘Ecological fitting’ is the process whereby the suites of traits an organism carries from previous evolutionary relationships are used to enable colonisation of novel environments or resources. 2. The concept has much explanatory power in studies of novel host associations, particularly when data suggest a deviation from optimal foraging theory, but is often overlooked in studies of herbivore host selection behaviour in favour of evolutionary hypotheses. 3. In the present study, the concept was used to explain the unusual host selection behaviour of the New Zealand endemic oligophagous butterfly Lycaena salustius Fabricius, the larvae of which feed on endemic Polygonaceae species and the introduced and closely related Fagopyrum esculentum Moench. 4. In field cage oviposition choice assays involving only endemic plants, females preferred to oviposit on the rare Muehlenbeckia astonii Petrie. However, the novel host F. esculentum was overwhelmingly preferred in an additional greenhouse assay. In larval no‐choice performance assays, fitness indicators were variable for the novel host. 5. This imperfect relationship between oviposition preference and larval performance is discussed as a possible example of ecological fitting and highlights the potential use of the concept as an explanatory tool in novel host selection behaviour studies.     

草地螟对寄主植物的选择性及其化学生态机制

田间自然条件下,草地螟成虫产卵对不同植物具有明显的选择性,其中灰菜上落卵量最多,占调查总卵量的50%以上;甜菜、大豆上次之,玉米和马铃薯上最少。在室内条件下,草地螟成虫产卵对灰菜、大豆、玉米和马铃薯的选择性与田间调查结果完全吻合。而且,草地螟幼虫的嗜食性与成虫产卵对寄主的选择性是一致的。吸附、收集上述植物的挥发物后进行GC-MS测定,并利用风洞进行了生测。其中灰菜和大豆的主要挥发物成分为反-2-己烯醛、青叶醇、青叶酯、1-十八烯等;玉米和马铃薯的挥发物性成分包括青叶酯、芳樟醇、香叶醇、紫罗酮、1-十八烯。生测结果表明,灰菜的挥发物对草地螟雌蛾引诱力最大,选择系数为78%;大豆苗挥发物对其也有较高的诱虫活性;但玉米和马铃薯几乎无引诱作用。自然生长的4种植物的诱虫结果与其挥发物的诱虫结果相似。这些生测结果与成虫和幼虫选择寄主的一致性反映了草地螟选择寄主行为的化学机制。     

Natal origin affects host preference and larval performance relationships in a tritrophic system

Many insects face the challenge to select oviposition sites in heterogeneous environments where biotic and abiotic factors can change over time. One way to deal with this complexity is to use sensory experiences made during developmental stages to locate similar habitats or hosts in which larval development can be maximized. While various studies have investigated oviposition preference and larval performance relationships in insects, they have largely overlooked that sensory experiences made during the larval stage can affect such relationships. We addressed this issue by determining the role of natal experience on oviposition preference and larval performance relationships in a tritrophic system consisting of Galerucella sagittariae, feeding on the two host plants Potentilla palustris and Lysimachia thyrsiflora, and its larval parasitoid Asecodes lucens. We firstly determined whether differences in host‐derived olfactory information could lead to divergent host selection, and secondly, whether host preference could result in higher larval performance based on the natal origin of the insects. Our results showed that the natal origin and the quality of the current host are both important aspects in oviposition preference and larval performance relationships. While we found a positive relationship between preference and performance for natal Lysimachia beetles, natal Potentilla larvae showed no such relationship and developed better on L. thyrsiflora. Additionally, the host selection by the parasitoid was mainly affected by the natal origin, while its performance was higher on Lysimachia larvae. With this study, we showed that the relationship between oviposition preference and larval performance depends on the interplay between the natal origin of the female and the quality of the current host. However, without incorporating the full tritrophic context of these interactions, their implication in insect fitness and potential adaptation cannot be fully understood.