Understanding Heliothine (Lepidoptera: Heliothinae) pests: What is a host plant?

Heliothine moths (Lepidoptera: Heliothinae) include some of the world's most devastating pest species. Whereas the majority of nonpest heliothinae specialize on a single plant family, genus, or species, pest species are highly polyphagous, with populations often escalating in size as they move from one crop species to another. Here, we examine the current literature on heliothine host-selection behavior with the aim of providing a knowledge base for research scientists and pest managers. We review the host relations of pest heliothines, with a particular focus on Helicoverpa armigera (Hubner), the most economically damaging of all heliothine species. We then consider the important question of what constitutes a host plant in these moths, and some of the problems that arise when trying to determine host plant status from empirical studies on host use. The top six host plant families in the two main Australian pest species (H. armigera and Helicoverpa punctigera Wallengren) are the same and the top three (Asteraceae, Fabaceae, and Malvaceae) are ranked the same (in terms of the number of host species on which eggs or larvae have been identified), suggesting that these species may use similar cues to identify their hosts. In contrast, for the two key pest heliothines in the Americas, the Fabaceae contains approximate to 1/3 of hosts for both. For Helicoverpa zea (Boddie), the remaining hosts are more evenly distributed, with Solanaceae next, followed by Poaceae, Asteraceae, Malvaceae, and Rosaceae. For Heliothis virescens (F.), the next highest five families are Malvaceae, Asteraceae, Solanaceae, Convolvulaceae, and Scrophulariaceae. Again there is considerable overlap in host use at generic and even species level. H. armigera is the most widely distributed and recorded from 68 plant families worldwide, but only 14 families are recorded as a containing a host in all geographic areas. A few crop hosts are used throughout the range as expected, but in some cases there are anomalies, perhaps because host plant relation studies are not comparable. Studies on the attraction of heliothines to plant odors are examined in the context of our current understanding of insect olfaction, with the aim of better understanding the connection between odor perception and host choice. Finally, we discuss research into sustainable management of pest heliothines using knowledge of heliothine behavior and ecology. A coordinated international research effort is needed to advance our knowledge on host relations in widely distributed polyphagous species instead of the localized, piecemeal approaches to understanding these insects that has been the norm to date.

Intra-specific variation for host plant use in Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae): Implications for management

The polyphagous moth Helicoverpa armigera (Hubner) is one of the world's most important agricultural pests. A number of existing approaches and future designs for management of H. armigera rely on the assumption that moths do not exhibit either genetically and/or non-genetically based variation for host plant utilization. We review recent empirical evidence demonstrating that both these forms of variation influence host plant use in this moth. The significance of this variation in H. armigera in relation to current and future pest management strategies is examined. We provide recommendations on future research needs and directions for sustainable management of H. armigera, under a framework that includes consideration of intra.-specific variation for host use relevant in this and other similar pest species. (C) 2004 Elsevier Ltd. All rights reserved.

Foraging behaviour of Helicoverpa armigera first instar larvae on crop plants of different developmental stages

Understanding how insect pests forage on their food plants can help optimize management strategies. Helicoverpa armigera (Hubner) (Lep., Noctuidae) is a major polyphagous pest of agricultural crops worldwide. The immature stages feed and forage on crops at all stages of plant development, damaging fruiting and non-fruiting structures, yet very little is known about the influence of host type or stage on the location and behaviour of larvae. Through semi-continuous observation, we evaluated the foraging (movement and feeding) behaviours of H. armigera first instar larvae as well as the proportion of time spent at key locations on mungbean [Vigna radiata (L.) Wilczek] and pigeon pea [Cajanus cajan (L.) Millspaugh] of differing developmental stages: seedling- and mature (flowering/pod fill)-stage plants. Both host type and age affected the behaviour of larvae. Larvae spent more time in the upper parts of mature plants than on seedlings and tended to stay at the top of mature plants if they moved there. This difference was greater in pigeon pea than in mungbean. The proportion of time allocated to feeding on different parts of a plant differed with host and age. More feeding occurred in the top of mature pigeon pea plants but did not differ between mature and seedling mungbean plants. The duration of key behaviours did not differ between plant ages in either crop type and was similar between hosts although resting bouts were substantially longer on mungbeans. Thus a polyphagous species such as H. armigera does not forage in equivalent ways on different hosts in the first instar stage.

Forecasting Helicoverpa populations in Australia: A comparison of regression based models and a bio-climatic based modeling approach

Long-term forecasts of pest pressure are central to the effective management of many agricultural insect pests. In the eastern cropping regions of Australia, serious infestations of Helicoverpa punctigera (Wallengren) and H. armigera (Hübner)(Lepidoptera: Noctuidae) are experienced annually. Regression analyses of a long series of light-trap catches of adult moths were used to describe the seasonal dynamics of both species. The size of the spring generation in eastern cropping zones could be related to rainfall in putative source areas in inland Australia. Subsequent generations could be related to the abundance of various crops in agricultural areas, rainfall and the magnitude of the spring population peak. As rainfall figured prominently as a predictor variable, and can itself be predicted using the Southern Oscillation Index (SOI), trap catches were also related to this variable. The geographic distribution of each species was modelled in relation to climate and CLIMEX was used to predict temporal variation in abundance at given putative source sites in inland Australia using historical meteorological data. These predictions were then correlated with subsequent pest abundance data in a major cropping region. The regression-based and bioclimatic-based approaches to predicting pest abundance are compared and their utility in predicting and interpreting pest dynamics are discussed.

Helicoverpa armigera (Hubner): can wheat stubble protect cotton plants against attack?

When investigating strategies for Helicoverpa armigera (Hubner) control, it is important to understand oviposition behaviour. Cotton (Gossypium hirsutum) was sown into standing wheat (Triticum astivum L.) stubble in a closed arena to investigate the effect of stubble on H. armigera moth behaviour and oviposition. Infrared cameras were used to track moths and determine whether stubble acted as a physical barrier or provided camouflage to cotton plants, thereby reducing oviposition. Searching activity was observed to peak shortly before dawn (03:00 and 04:00 h) and remained high until just after dawn (4 h window). Moths spent more time resting on cotton plants than spiralling above them, and the least time flying across the arena. While female moths spent more time searching for cotton plants growing in wheat stubble, the difference in oviposition was not significant. As similar numbers of eggs were laid on cotton plants with stubble (3.5/plant SE +/- 0.87) and without stubble (2.5/plant SE +/- 0.91), wheat stubble does not appear to provide camouflage to cotton plants. There was no significant difference in the location of eggs deposited on cotton plants with and without stubble, although more eggs were laid on the tops of cotton leaves in wheat stubble. As the spatial and temporal distribution of eggs laid on the cotton plant is a crucial component of population stability, eggs laid on the upper side of leaves on cotton plants may be more prone to fatalities caused by environmental factors such as wind and rain. Therefore, although stubble did not influence the number of eggs laid, it did affect their distribution on the plant, which may result in increased mortality of eggs on cotton plants sown into standing wheat stubble.

Host recognition by a polyphagous lepidopteran (Helicoverpa armigera): primary host plants, host produced volatiles and neurosensory stimulation

An important question in the host-finding behaviour of a polyphagous insect is whether the insect recognizes a suite or template of chemicals that are common to many plants? To answer this question, headspace volatiles of a subset of commonly used host plants (pigeon pea, tobacco, cotton and bean) and nonhost plants (lantana and oleander) of Helicoverpa armigera Hubner (Lepidoptera: Noctuidae) are screened by gas chromatography (GC) linked to a mated female H. armigera electroantennograph (EAG). In the present study, pigeon pea is postulated to be a primary host plant of the insect, for comparison of the EAG responses across the test plants. EAG responses for pigeon pea volatiles are also compared between females of different physiological status (virgin and mated females) and the sexes. Eight electrophysiologically active compounds in pigeon pea headspace are identified in relatively high concentrations using GC linked to mass spectrometry (GC-MS). These comprised three green leaf volatiles [(2E)-hexenal, (3Z)-hexenylacetate and (3Z)-hexenyl-2-methylbutyrate] and five monoterpenes (alpha-pinene,beta-myrcene, limonene, E-beta-ocimene and linalool). Other tested host plants have a smaller subset of these electrophysiologically active compounds and even the nonhost plants contain some of these compounds, all at relatively lower concentrations than pigeon pea. The physiological status or sex of the moths has no effect on the responses for these identified compounds. The present study demonstrates how some host plants can be primary targets for moths that are searching for hosts whereas the other host plants are incidental or secondary targets.

Predicting population dynamics of weed biological control agents: science or gazing into crystal balls?

Various factors can influence the population dynamics of phytophages post introduction, of which climate is fundamental. Here we present an approach, using a mechanistic modelling package (CLIMEX), that at least enables one to make predictions of likely dynamics based on climate alone. As biological control programs will have minimal funding for basic work (particularly on population dynamics), we show how predictions can be made using a species geographical distribution, relative abundance across its range, seasonal phenology and laboratory rearing data. Many of these data sets are more likely to be available than long-term population data, and some can be incorporated into the exploratory phase of a biocontrol program. Although models are likely to be more robust the more information is available, useful models can be developed using information on species distribution alone. The fitted model estimates a species average response to climate, and can be used to predict likely geographical distribution if introduced, where the agent is likely to be more abundant (i.e. good locations) and more importantly for interpretation of release success, the likely variation in abundance over time due to intra- and inter-year climate variability. The latter will be useful in predicting both the seasonal and long-term impacts of the potential biocontrol agent on the target weed. We believe this tool may not only aid in the agent selection process, but also in the design of release strategies, and for interpretation of post-introduction dynamics and impacts. More importantly we are making testable predictions. If biological control is to become more of a science making and testing such hypothesis will be a key component.

IPM in the transgenic era: a review of the challenges from emerging pests in Australian cotton systems

The Cotton Catchment Communities Cooperative Research Centre began during a period of rapid uptake of Bollgard II® cotton, which contains genes to express two Bt proteins that control the primary pests of cotton in Australia, Helicoverpa armigera and H. punctigera. The dramatic uptake of this technology presumably resulted in strong selection pressure for resistance in Helicoverpa spp. against the Bt proteins. The discovery of higher than expected levels of resistance in both species against one of the proteins in Bollgard II® cotton (Cry2Ab) led to significant re-evaluation of the resistance management plan developed for this technology, which was a core area of research for the Cotton CRC. The uptake of Bollgard II® cotton also led to a substantial decline in pesticide applications against Helicoverpa spp. (from 10–14 to 0–3 applications per season). The low spray environment allowed some pests not controlled by the Bt proteins to emerge as more significant pests, especially sucking species such as Creontiades dilutus and Nezara viridula. A range of other minor pests have also sporadically arisen as problems. Lack of knowledge and experience with these pests created uncertainty and encouraged insecticide use, which threatened to undermine the gains made with Bollgard II® cotton. Here we chronicle the achievements of the Cotton CRC in providing the industry with new knowledge and management strategies for these pests.

Revision of the Neotropical moth genera Mallodeta Butler and Erruca Walker, revalidated (Noctuidae, Arctiinae, Arctiini, Euchromiina)

Mallodeta Butler and Erruca Walker, revalidated, are redescribed and revised. Mallodeta henceforth includes only its type-species, Glaucopis (Lycorea) clavata Walker, and Erruca is resurrected with seven species: E. deyrolii Walker (type-species), E. consors (Walker), new combination, E. erythrarchos (Walker), new combination, E. cardinalis (Hampson), new combination, E. hanga (Herrich-Schaffer), new combination, E. cruenta (Perty), new combination and E. sanguipuncta (Druce), new combination. Six new synonyms are established, four specific and two generic (junior synonyms in parentheses): Zygaena capistrata Fabricius (=Mallodeta cubana Gaede), Glaucopis (Lycorea) clavata Walker (=M. simplex Rothschild), Erruca deyrolii Walker (=Laemocharis aecyra Herrich-Schaffer and Glaucopis (Hyda) sortita Walker), and Erruca Walker (=Aristodaema Wallengren and Rezia Kirby). Lectotypes are designated to the following species: Erruca deyrolii Walker, Laemocharis deyrollei Herrich-Schaffer, Laemocharis hanga Herrich-Schaffer, Laemocharis aecyra Herrich-Schaffer, Laemocharis norma Herrich-Schaffer, Cosmosoma cardinalis Hampson and Mallodeta sanguipuncta Druce. Illustrations of adults and male and female genitalia of Mallodeta and Erruca are provided, as well as a key to the species of the latter.

Surface sediment analyses, bivalve and gastropod taxa of marine sediments from the Banc d'Arguin, Mauritania

High-nutrient tropical carbonate systems are known to produce sediments that, in terms of skeletal composition, are reminiscent of their extra-tropical counterparts. Such carbonate systems and associated carbonate grain assemblages in the tropics are rare in the present-day world. Nonetheless, it is crucial to gain a better understanding of those ecosystems, including their drivers and players because such settings potentially represent models for ancient depositional systems as well as for predicted future environmental conditions. One of the modern occurrences of eutrophic tropical carbonate systems is the northern Mauritanian Shelf. The marine environment is characterized by an eastern boundary upwelling system that pushes cool and nutrient-rich intermediate waters onto a wide epicontinental platform (Golfe d'Arguin) where the waters warm up to tropical temperatures. The resulting facies is mixed carbonate-siliciclastic with a dominant foramol association grading into bimol and barnamol grain assemblages in the shallowest areas forming the Banc d'Arguin. Besides this cool water-related heterozoan association, the carbonate sediment is characterized by tropical molluskan species, while chlorozoan biota (e.g., corals and algal symbiont-bearing foraminifers) are entirely absent. We here present a first comprehensive facies analysis of this model example of eutrophic tropical carbonates. Furthermore, we reconstruct the loci of carbonate production and provide a conclusive depositional model of the Banc d'Arguin that received little attention to date due to its poorly accessible nature.

Abiotic characteristics and diatom abundance and biomass in different sectors of the Argentinien Shelf and antartic waters

A large spatial scale study of the diatom species inhabiting waters from the subantarctic (Argentine shelf) to antarctic was made for the first time in order to understand the relationships between these two regions with regard to the fluctuations in diatom abundances in relation with environmental features, their floristic associations and the effect of the Polar Front as a biogeographic barrier. Species-specific diatom abundance, nutrient and chlorophyll-a concentration were assessed from 64 subsurface oceanographic stations carried out during the austral summer 2002, a period characterized by an anomalous sea-ice coverage corresponding to a ''warm year". Significant relationships of both diatom density and biomass with chlorophyll-a (positive) and water temperature (negative) were found for the study area as a whole. Within the Subantarctic region, diatom density and biomass values were more uniform and significantly (in average: 35 and 11 times) lower than those of the Antarctic region, and did not correlate with chlorophyll-a. In antarctic waters, instead, biomass was directly related with chlorophyll-a, thus confirming the important contribution of diatoms to the Antarctic phytoplanktonic stock. A total of 167 taxa were recorded for the entire study area, with Chaetoceros and Thalassiosira being the best represented genera. Species richness was maximum in subantarctic waters (46; Argentine shelf) and minimum in the Antarctic region (21; Antarctic Peninsula), and showed a significant decrease with latitude. Floristic associations were examined both qualitatively (Jaccard Index) and quantitatively (correlation) by cluster analyses and results allowed differentiating a similar number of associations (12 vs. 13, respectively) and two main groups of stations. In the Drake Passage, the former revealed that the main floristic change was found at the Polar Front, while the latter reflected the Southern ACC Front as a main boundary, and yielded a higher number of isolated sites, most of them located next to different Antarctic islands. Such differences are attributed to the high relative density of Fragilariopsis kerguelensis in Argentine shelf and Drake Passage waters and of Porosira glacialis and species of Chaetoceros and Thalasiosira in the Weddell Sea and near the Antarctic Peninsula. From a total of 84 taxa recorded in antarctic waters, only 17 were found exclusively in this region, and the great majority (67) was also present in subantarctic waters but in extremely low (< 1 cell/l) concentrations, probably as a result of expatriation processes via the ACC-Malvinas Current system. The present results were compared with those of previous studies on the Antarctic region with respect to both diatom associations in regular vs. atypically warm years, and the distribution and abundance of some selected planktonic species reported for surface sediments.

Seawater carbonate chemistry, stable carbon isotope fractionation and growth rate during experiments with marine phytoplankton community, 1999

Stable carbon isotope fractionation (%) of 7 marine phytoplankton species grown in different irradiance cycles was measured under nutrient-replete conditions at a high light intensity in batch cultures. Compared to experiments under continuous light, all species exhibited a significantly higher instantaneous growth rate (pi), defined as the rate of carbon fixation during the photo period, when cultivated at 12:12 h. 16:8 h, or 186 h light:dark (L/D) cycles. Isotopic fractionation by the diatoms Skeletonema costatum, Asterionella glacialis, Thalassiosira punctigera, and Coscinodiscus wailesii (Group I) was 4 to 6% lower in a 16:8 h L/D cycle than under continuous light, which we attribute to differences in pi. In contrast, E, in Phaeodactylum tn'cornutum, Thalassiosira weissflogii, and in the dinoflagellate Scrippsiella trochoidea (Group 11) was largely insensitive to day length-related differences in instantaneous growth rate. Since other studies have reported growth-rate dependent fractionation under N-limited conditions in P. tricornutum, pi-related effects on fractionation apparently depend on the factor controlling growth rate. We suggest that a general relationship between E, and pi/[C02,,,] may not exist. For 1 species of each group we tested the effect of variable CO2 concentration, [COz,,,], on isotopic fractionation. A decrease in [CO2,,,] from ca 26 to 3 pm01 kg-' caused a decrease in E, by less than 3%0 This indicates that variation in h in response to changes in day length has a similar or even greater effect on isotopic fractionation than [COz,,,] m some of the species tested. In both groups E, tended to be higher in smaller species at comparable growth rates. In 24 and 48 h time series the algal cells became progressively enriched in 13C during the day and the first hours of the dark period, followed by l3C depletion in the 2 h before beginning of the following Light period. The daily amplitude of the algal isotopic composition (613C), however, was <1.5%0, which demonstrates that diurnal variation in Fl3C is relatively small.