Host selection in phytophagous insects: a new explanation for learning in adults

Insect learning can change the preferences an egg laying female displays towards different host plant species. Current hypotheses propose that learning may be advantageous in adult host selection behaviour through improved recognition, accuracy or selectivity in foraging. In this paper, we present a hypothesis for when learning can be advantageous without such improvements in adult host foraging. Specifically, that learning can be an advantageous strategy for egg laying females when larvae must feed on more than one plant in order to complete development, if the fitness of larvae is reduced when they switch to a different host species. Here, larvae benefit from developing on the most abundant host species, which is the most likely choice of host for an adult insect which increases its preference for a host species through learning. The hypothesis is formalised with a mathematical model and we provide evidence from studies on the behavioural ecology of a number of insect species which demonstrate that the assumptions of this hypothesis may frequently be fulfilled in nature. We discuss how multiple mechanisms may convey advantages in insect learning and that benefits to larval development, which have so far been overlooked, should be considered in explanations for the widespread occurrence of learning.

A general model for host plant selection in phytophagous insects

We develop a general theoretical framework for exploring the host plant selection behaviour of herbivorous insects. This model can be used to address a number of questions, including the evolution of specialists, generalists, preference hierarchies, and learning. We use our model to: (i) demonstrate the consequences of the extent to which the reproductive success of a foraging female is limited by the rate at which they find host plants (host limitation) or the number of eggs they carry (egg limitation); (ii) emphasize the different consequences of variation in behaviour before and after landing on (locating) a host (termed pre- and post-alighting, respectively); (iii) show that, in contrast to previous predictions, learning can be favoured in post-alighting behaviour--in particular, individuals can be selected to concentrate oviposition on an abundant low-quality host, whilst ignoring a rare higher-quality host; (iv) emphasize the importance of interactions between mechanisms in favouring specialization or learning.

Early-season movement dynamics of phytophagous pest and natural enemies across a native vegetation-crop ecotone

There is limited understanding about how insect movement patterns are influenced by landscape features, and how landscapes can be managed to suppress pest phytophage populations in crops. Theory suggests that the relative timing of pest and natural enemy arrival in crops may influence pest suppression. However, there is a lack of data to substantiate this claim. We investigate the movement patterns of insects from native vegetation (NV) and discuss the implications of these patterns for pest control services. Using bi-directional interception traps we quantified the number of insects crossing an NV/crop ecotone relative to a control crop/crop interface in two agricultural regions early in the growing season. We used these data to infer patterns of movement and net flux. At the community-level, insect movement patterns were influenced by ecotone in two out of three years by region combinations. At the functional-group level, pests and parasitoids showed similar movement patterns from NV very soon after crop emergence. However, movement across the control interface increased towards the end of the early-season sampling period. Predators consistently moved more often from NV into crops than vice versa, even after crop emergence. Not all species showed a significant response to ecotone, however when a response was detected, these species showed similar patterns between the two regions. Our results highlight the importance of NV for the recruitment of natural enemies for early season crop immigration that may be potentially important for pest suppression. However, NV was also associated with crop immigration by some pest species. Hence, NV offers both opportunities and risks for pest management. The development of targeted NV management may reduce the risk of crop immigration by pests, but not of natural enemies.

Does Gamma-aminobutyric acid function as a plant resistance mechanism against phytophagous insect activity? /

Gamma-aminobutyric acid (GAB A) is a ubiquitous non-protein amino acid synthesized via the decarboxylation of L-glutamate in a reaction catalyzed by the cytosolic enzyme L-glutamate decarboxylase (GAD). In animals it functions as an inhibitory neurotransmitter. In plants it accumulates rapidly in response to various stresses, but its function remains unclear. The hypothesis that GABA accumulation in leaf tissue may function as a plant resistance mechanism against phytophagous insect activity was investigated. GABA accumulation in response to mechanical stimulation, mechanical damage and insect activity was demonstrated. In wt tobacco (Nicotiana tabacum cv Samsun), mechanical stimulation or damage caused GABA to accumulate within 2 min from mean levels of 14 to 37 and 1~9 nmol g-l fresh weight (FW), respectively. In the transgenic tobacco strain CaMVGAD27c overexpressing Petunia GAD, the same treatments caused GABA to accumulate from 12 to 59 and 279 nmol g-l FW, respectively. In the transgenic tobacco strain CaMVGADilC 11 overexpressing Petunia GAD lacking an autoinhibitory domain, mechanical stimulation or damage caused GABA to accumulate from 180 to 309 and 630 nmol g-l FW, respectively. Ambulatory activity by tobacco budworm (TBW) larvae (Heliothis virescens) on leaves of CaMVGAD27c tobacco caused GABA to accumulate from 28 to 80 nmol g-l FW within 5 min. Ambulatory and leaf-rolling activity by oblique banded leaf roller (OBLR) larvae (Choristoneura rosaceana cv Harris) on wt soybean leaves (Glycine max cv Harovinton) caused GABA to accumulate from 60 to 1123 nmol g-l FW within 20 min. Increased GABA levels in leaf tissue were shown to affect phytophagous preference in TBW larvae presented with wt and transgenic tobacco leaves. When presented with leaves of Samsun wt and CaMVGAD27c plants, TBW larvae consumed more wt leaf tissue (640 ± 501 S.D. mm2 ) than transgenic leaf tissue (278 ± 338 S.D. mm2 ) nine times out of ten. When presented with leaves of Samsun wt and CaMVGAD~C11 plants, TBW larvae consumed more transgenic leaf tissue (1219 ± 1009 S.D. mm2 ) than wt leaf tissue (28 ± 31 S.D. mm2 ) ten times out of ten. These results indicate that: (1) ambulatory activity of insect larvae on leaves results in increased GABA levels, (2) transgenic tobacco leaves with increased capacity for GABA synthesis deter feeding, and (3) transgenic tobacco leaves with constitutively higher GABA levels stimulate feeding.

Contrasting success in the restoration of plant and phytophagous beetle assemblages of species-rich mesotrophic grasslands

Over the last 60 years changes to the management of species-rich mesotrophic grasslands have resulted in the large-scale loss and degradation of this habitat across Europe. Restoration of such grasslands on agriculturally improved pastures provides a potentially valuable approach to the conservation of these threatened areas. Over a four-year period a replicated block design was used to test the effects of seed addition (green hay spreading and brush harvest collection) and soil disturbance on the restoration of phytophagous beetle and plant communities. Patterns of increasing restoration success, particularly where hay spreading and soil disturbance were used in combination, were identified for the phytophagous beetles. In the case of the plants, however, initial differences in restoration success in response to these same treatments were not followed by subsequent temporal changes in plant community similarity to target mesotrophic grassland. It is possible that the long-term consequences of the management treatments would not be the establishment of beetle and plant communities characteristic of the targets for restoration. Restoration management to enhance plant establishment using hay spreading and soil disturbance techniques would, however, still increase community similarity in both taxa to that of species-rich mesotrophic grasslands, and so raise their conservation value.

The importance of sward architectural complexity in structuring predatory and phytophagous invertebrate assemblages

1. Although the importance of plant community assemblages in structuring invertebrate assemblages is well known, the role that architectural complexity plays is less well understood. In particular, direct empirical data for a range of invertebrate taxa showing how functional groups respond to plant architecture is largely absent from the literature. 2. The significance of sward architectural complexity in determining the species richness of predatory and phytophagous functional groups of spiders, beetles, and true bugs, sampled from 135 field margin plots over 2 years was tested. The present study compares the relative importance of sward architectural complexity to that of plant community assemblage. 3. Sward architectural complexity was found to be a determinant of species richness for all phytophagous and predatory functional groups. When individual species responses were investigated, 62.5% of the spider and beetle species, and 50.0% of the true bugs responded to sward architectural complexity. 4. Interactions between sward architectural complexity and plant community assemblage indicate that the number of invertebrate species supported by the plant community alone could be increased by modification of sward architecture. Management practices could therefore play a key role in diversifying the architectural structure of existing floral assemblages for the benefit of invertebrate assemblages. 5. The contrasting effects of sward architecture on invertebrate functional groups characterised by either direct (phytophagous species) or indirect (predatory species) dependence on plant communities is discussed. It is suggested that for phytophagous taxa, plant community assemblage alone is likely to be insufficient to ensure successful species colonisation or persistence without appropriate development of sward architecture.

The role of management and landscape context in the restoration of grassland phytophagous beetles

1. Declines in area and quality of species-rich mesotrophic and calcareous grasslands have occurred all across Europe.While the European Union has promoted schemes to restore these grasslands, the emphasis for management has remained largely focused on plants. Here we focus on restoration of the phytophagous beetles of these grasslands. Although local management, particularly that which promotes the establishment of host plants, is key to restoration success, dispersal limitation is also likely to be an important limiting factor during the restoration of phytophagous beetle assemblages. 2. Using a 3-year multi-site experiment, we investigated how restoration success of phytophagous beetles was affected by hay-spreading management (intended to introduce target plant species), success in restoration of the plant communities and the landscape context within which restoration was attempted. 3. Restoration success of the plants was greatest where green hay spreading had been used to introduce seeds into restoration sites. Beetle restoration success increased over time, although hayspreading had no direct effect. However, restoration success of the beetles was positively correlated with restoration success of the plants. 4. Overall restoration success of the phytophagous beetles was positively correlated with the proportion of species-rich grassland in the landscape, as was the restoration success of the polyphagous beetles. Restoration success for beetles capable of flight and those showing oligophagous host plant specialism were also positively correlated with connectivity to species-rich grasslands. There was no indication that beetles not capable of flight showed greater dependence on landscape scale factors than flying species. 5. Synthesis and applications. Increasing the similarity of the plant community at restoration sites to target species-rich grasslands will promote restoration success for the phytophagous beetles. However, landscape context is also important, with restoration being approximately twice as successful in those landscapes containing high as opposed to low proportions of species-rich grassland. By targeting grassland restoration within landscapes containing high proportions of species-rich grassland, dispersal limitation problems associated with restoration for invertebrate assemblages are more likely to be overcome.

The restoration of phytophagous beetles in species-rich chalk grasslands

This study focuses on the restoration of chalk grasslands over a 6-year period and tests the efficacy of two management practices, hay spreading and soil disturbance, in promoting this process for phytophagous beetles. Restoration success for the beetles, measured as similarity to target species-rich chalk grassland, was not found to be influenced by either management practice. In contrast, restoration success for the plants did increase in response to hay spreading management. Although the presence of suitable host plants was considered to dictate the earliest point at which phytophagous beetles could successfully colonized, few beetle species colonized as soon as their host plants became established. Morphological characteristics and feeding habits of 27 phytophagous beetle species were therefore tested to identify factors that limited their colonization and persistence. The lag time between host plant establishment and colonization was greatest for flightless beetles. Beetles with foliage-feeding larvae both colonized at slower rates than seed-, stem-, or root-feeding species and persisted within the swards for shorter periods. Although the use of hay spreading may benefit plant communities during chalk grassland restoration, it did not directly benefit phytophagous beetles. Without techniques for overcoming colonization limitation for invertebrate taxa, short-term success of restoration may be limited to the plants only.

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.

A phytophagous braconid, Allorhogas conostegia sp nov (Hymenoptera: Braconidae), in the fruits of Conostegia xalapensis (Bonpl.) D. Don (Melastomataceae)

Before 1989 all braconid wasps were thought to be parasitoids, but in that year the first phytophagous species was reported. Subsequently, a few other examples of phytophagy have been discovered, most of which are species of Allorhogas in the subfamily Doryctinae. Until now, all demonstrated examples of phytophagy in this genus have been as gall inducers in the fruits of Fabaceae. Here we describe a new species from Costa Rica, Allorhogas conostegia Marsh and Shaw, and provide evidence that it forms galls in the fruits of Conostegia xalapensis (Melastomataceae). We also provide information on the phenology of the plant and of the galls and the effects of the galls on the host plant, and we discuss the potential species richness of Allorhogas in the Neotropics.

A new phytophagous Bracon Fabricius (Hymenoptera, Braconidae) associated with Protium ovatum Engl. (Burseraceae) fruits from Brazilian savannah

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Population Fluctuation and Damage Caused by Phytophagous Mites on Three Rubber Tree Clones

Our aim was to investigate the population fluctuation and the damage caused by the phytophagous mites Calacarus heveae Feres, Tenuipalpus heveae Baker, and Eutetranychus banksi (McGregor) on clones FX 2784, FX 3864, and MDF 180 in rubber tree crops from southeastern Bahia, Brazil. Moreover, we tested for the influence of climatic variables on occurrence patterns of these species throughout weekly samples performed from October to April. The infestation peaks was between mid-January and late February. The clones FX 2784 and FX 3864 had the highest infestations and more severe damage possibly caused by C. heveae, which was the most frequent and abundant species in all clones. We found that sunlight duration and rainfall were the most important factors for C. heveae while T. heveae was affected by rainfall and temperature. Eutetranychus banksi was only affected by sunlight duration. However, the best models had low goodness of fit. We concluded that the clones FX 2784 and FX 3864 had a higher susceptibility to mite attack, and the association between climatic variables and favorable physiological conditions were determinant for the population increase of the species from January to April. © 2012 Sociedade Entomológica do Brasil.

Tometes camunani (Characiformes: Serrasalmidae), a new species of phytophagous fish from the Guiana Shield, rio Trombetas basin, Brazil

Uma espécie nova de Serrasalmidae, Tometes camunani, é descrita para as drenagens superiores da bacia do rio Trombetas, estado do Pará, Brasil. A espécie nova distingue-se dos congêneres pela presença de uma ligeira concavidade no neurocrânio na altura do frontal (vs. concavidade ausente, perfil dorsal do neurocrânio reto). Também pode ser adicionalmente distinguido dos seus congêneres por possuir dentes com a cúspide central mais alta e cume agudo (vs. cúspide central mais baixa e com cume arredondado em T. trilobatus), a boca terminal (vs. boca orientada para cima em T. lebaili), e 12-26 espinhos pré-pélvicos (vs. 0-9 em T. makue). A espécie nova é estritamente reofílica, como as demais espécies de Tometes, e ocorre exclusivamente nas zonas encachoeiradas dos rios de escudo, biótopos complexos, frágeis e ameaçados por ações antropogênicas. Uma chave de identificação para as espécies do grupo Myleus é apresentada.

Biological control of phytophagous arthropods in the physic nut tree Jatropha curcas L. in Brazil

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)